Protist, Vol.

153, 177190, June 2002 Urban & Fischer Verlag

http://www.urbanfischer.de/journals/protist
Published online 29 May 2002

Protist

FROM THE ARCHIVES

A Salute to Antony van Leeuwenhoek of Delft,

Most Versatile 17th Century Founding Father
of Protistology
John O. Corliss1
P.O. Box 2729, Bala Cynwyd, OA 19004, USA

Im well aware that these my writings will not be accepted by some, as they judge it to be impossible to
make such discoveries [but] I will say once more that tis my habit to hold fast to my notions only until Im
better informed or till my observations make me go over to others
A. van Leeuwenhoek in letters to the Royal Society
(excerpts taken from English translations given by Dobell 1932)

During the third and last century of the Renaissance,

that great revival of art, literature, and learning all
across Europe, the amateur scientist Antony van
Leeuwenhoek (Fig. 1) was unobtrusively born in October 1632 in the quiet but thriving Dutch town of
Delft. Also born that same year were other men destined for eternal fame in quite different fields: for example, John Locke, Baruch de Spinoza, Christopher
Wren, Nicolaes Maes, and in the very same town
Jan Vermeer. That date, interestingly enough, marked
the middle of the drawn-out Thirty Years War, with all
its religious and political machinations and implications, an off-again-on-again conflict sometimes pitting England against Holland (The Netherlands). Recall that the 17th century also witnessed the beginning of the Enlightenment Period of history.
Although the man who is the subject of this brief
tribute died in 1723, two months shy of his 91st birthday, he and his impressive scientific contributions
were very little known and seldom publicized and appreciated until their major revelation in the 20th century more than 200 years after his death. It is true that

fax 1-610-664-4904
e-mail jocchezmoi@aol.com

Figure 1. Antony van Leeuwenhoek, in his mid-fifties.

A photograph of a portion of Dutch painter Johannes
Verkoljes celebrated mezzotint engraving of 1686,
taken in 1970 for the authors private collection of pictures of great protistologists.
1434-4610/02/153/02-177 $ 15.00/0

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J. O. Corliss

W. Saville Kent (1880) and Otto Btschli (1887), and

succeeding protozoologists knowing of the tremendous monographs on protists of these two prolific
19th century giants in the field (Btschli 18801889,
Kent 18801882), were and have been quite aware of
some of the Leeuwenhoek protistological discoveries, but not always accurately or in detail.
The most comprehensive treatments to emerge of
Leeuwenhoeks life and letters, and of his numerous
contributions to the many fields dependent in large
measure on precise microscopical observation,
have been the following: Cole (1937), Dobell (1932),
Ford (1985, 1991), Schierbeek (1959, 1963; and as a
major editor, 1939?, of a long-continued series of
volumes published in both Dutch and English), and a
number of papers (not cited here) by several 20th
century Dutch microscopists or microbiologists
written in their own language only. Shorter biographies, but of some comparative historical value and
to be noted for their accuracy, include accounts in
Beltrn (1974), Bulloch (1938), Cole (1926, 1938),
Corliss (1975, 19781979, 1992), Dobell (1923),
Lechevalier and Solotorovsky (1965), Locy (1935),
Meyer (1937), Nordenskild (1928), Rooseboom
(1956), Singer (1959), and Zuylen (1981).

Popular Father of
Titles for the Man from Delft
Before examining the life and works of Leeuwenhoek, the amateur biologist who, with his simple but
high-powered single-lens microscope made such
early astounding observations on the theretofore
largely unseen world of cells, protists, animal and
plant tissues, I should mention briefly the discoverer titles generously and usually deservedly bestowed upon him during the past century. If we concede that Father of does not confer sole or absolutely very first discovery in a field, then the following
accolades are quite accurate: Father of Protistology,
Father of Protozoology, Father of Phycology (at the
microscopic level), Father of Microbiology, Father of
Bacteriology, Father of Hematology or Serology (he
saw erythrocytes and extended Harveys landmark
observations of 1618 to include capillary circulation
in tissues), Father of Plant (Microscopic) Anatomy,
and even Father of Microbial Ecology.
Additional titles that may be a little exaggerated
or a bit over-extravagant would include Father of
Parasitology (but he was indeed the first to see, describe, and determine the sizes of symbiotic bacteria and protozoa from the digestive tracts or feces of
several invertebrates and vertebrates, including humans); Father of Crystallography (a superb glass-

blower, he ground his own lenses with unusual precision, sometimes from grains of pure sand); Father
of (simple) Stain and (razor-blade) Sectioning Technologies; and Father of Pesticide Control (he experimentally demonstrated the effects of certain chemicals on microorganisms and small invertebrates) .
Further accolades can be found, ones occasionally still in usage although they are or may be technically quite incorrect. An outstanding example is the
persistent but falsely bestowed Father of Microscopy title. Large-bodied compound microscopes, with eyepiece, draw tube, multiple lenses, a
mirror, etc., were in existence and usage a full half
century before Leeuwenhoeks work and have
served as models for modern equipment (see Bradbury 1967, Singer et al. 1957, Woodruff 1939, and
numerous other compendia on microscopy and its
history). Other examples of mostly inaccurate labels
include Father of Cytology (although his comparative observations on spermatozoa, for example,
stand as landmarks in the history of biology); Father
of Histology (but both plant and animal tissues did
come under his early scrutiny); Father of Embryology (yet he did observe early cell division stages in
eggs of some invertebrates and in spores of some
plant species, not to mention reproduction in the
alga Volvox, and an account of the reproductive organs of ants; and he described budding in the
cnidarian Hydra some four to five decades before
the classical, long-well-known papers by Trembley,
Baker, and other zoologists); Father of (microscopic)
Mycology (he did offer early descriptions of yeast
and some mold cells/spores); and Father of (microscopic) Neurology (he described hand-sectioned
bovine optic nerve material in 1674 and 1675). He
even contributed substantially to the field of microcrustacean anatomy, perhaps inspired by his long
and close friendship with Jan Swammerdam (born
1637, died 1680) whose celebrated work on Daphnia (then known simply as a water-flea or waterlouse) first appeared in 1669 (in Dutch).
We certainly may conclude that if there were a
section in the Guinness Book of World Records entitled Most Fathers of Scientific Fields, the versatile Mr. van Leeuwenhoek would definitely rate a
first-place position in it!

Brief Account of Major Periods

in Leeuwenhoeks Life
It may be helpful to arbitrarily divide Leeuwenhoeks
long life span into roughly three periods, each of
about 30 years duration. This biographical and
overview section is followed by more detailed atten-

Antony van Leeuwenhoek

tion to his beloved magnifying glass (as he himself

called it) and to his recorded descriptions, including
reproduction of some of his (or his draftsmens) sufficiently accurate drawings to allow taxonomic identifications to be made with considerable certainty
today.

His First 30 Years

Born into a large family (but the only son) in Delft,
Holland, on 24 October, 1632, young Antony was
sent away for schooling until about age 15 or 16. He
may have learned some rudimentary mathematics
and physics but didnt take Latin or foreign languages or other subjects which we in America today
call requisite parts of a classical or college preparatory curriculum. At the age of 16, he was sent to
Amsterdam to master the linen-draper trade. Six
years later, he returned to Delft where he set up his
own shop and remained for the rest of his life.
In 1654, Antony married his first wife (Barbara)
and they had five children, only one of whom, Maria,
survived beyond early years. In fact, she became
her fathers faithful right-hand man for many
decades, passing away in 1745 at the ripe old age of
(nearly) 89, surviving her nonagenarian dad by some
22 years. The year 1654 was historically a very important one for Delft: a huge explosion at its arsenal
caused fires killing thousands of its residents and
destroying much of the town. As fate would have it,
spared among others were two of its young (22year-old) inhabitants of particular interest to us here,
Leeuwenhoek and Vermeer.
Incidentally, in 1664, to look beyond the first third
of Antonys life for a moment, his mother passed
away; both his father and his stepfather had died
much earlier. In 1666, his wife Barbara died; he remarried (to Cornelia) three years later; childless, she
passed away in 1694, leaving Leeuwenhoek with
only his loving daughter Maria as close companion
for the rest of his life.

His Second 30 Years

The middle third of Leeuwenhoeks full life is full of
significant happenings. On the local scene, he became a well recognized citizen of the town, was
made a municipal surveyor, named the official winegauger (did assaying all wines and spirits entering
Delft contribute to his longevity!?), and given the
post of Chamberlain to the Sheriffs Office (not a janitors job, as is sometimes suggested, but really an
honorary sinecure with a salary to help support
along with his drapery business income his various
living expenses while he did what he wished as a

179

naturalist of the microbial world, the latter a hobby

of his that was increasingly becoming known to others by the 1670s). In 1676, he was appointed the executor of the estate of his departed close friend Jan
Vermeer, whose art and philosophical outlook on life
so much appealed to him.
Twas his only trip to London, in 1668 (during a lull
between new sets of Anglo-Dutch Wars and two
years after the terrible London fire), that undoubtedly
heavily influenced how he was to spend the rest of
his life. Although Dobell (1932) and Dutch historians
have uniformly insisted that Leeuwenhoeks passion
for objects microscopical was entirely self-generated, modern investigator and biographer Brian Ford
(1991) has offered the surprising but convincing supposition that Robert Hookes (1665, 1667) Micrographia was seen and read (with the help of English
friends as translators) by him and excited him
greatly during and well after that fateful trip. Recall
that Hooke described the intimate structure of a
number of textiles in his highly popular monograph,
using a single lens magnifying glass; he also figured
cells (the spaces left by them) in razor-blade sectioned cork, the discovery of his that biologists are
apt to remember best. And, among other observations made, Hooke postulated what causes chalk to
appear white. Now Leeuwenhoek himself, as a textile merchant, was well acquainted with the weaving
patterns, etc. in different kinds of cloth. And on his
trip across the English Channel he became curious
about the nature of the white cliffs of Dover, independently wondering about their reflection of light, etc.
The careful drawings by Hooke whose second edition of Micrographia had become available very likely
by the time of the Dutchmans visit to London could
certainly have stimulated Antony to delve further into
such matters (while Hooke himself mostly went off
into other fields of investigation).
It is known that the two men corresponded,
though they never met, and Hooke (born 1635, died
1703) retained a lifetime high admiration for
Leeuwenhoeks subsequent microscopical observations. In fact, it was Hooke who was behind the latters nomination to the Royal Society, an honor bestowed in 1680. But it was a friend and fellow resident of Delft, the eminent anatomist Reinier de
Graaf, who first brought Henry Oldenburgs (first
Secretary of the Society) attention to Leeuwenhoek
by letter in 1673 (the very year of de Graafs untimely
death at the age of 32); and later that same year a
delegation from the Society visited Delft and returned with glowing reports of the simple but effective microscopes they had seen and the multiple
uses of them by their ingenious Dutch maker. This
resulted in an arrangement whereby Leeuwenhoek

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J. O. Corliss

would write to Oldenburg of his discoveries over the

many years following that important visit. He was
still penning and posting epistles (totaling > 200) to
the Royal Society, most always handwritten and in
his native language, during his last days on earth in
the month of August 1723, some 45 years after the
death of editor Oldenburg and of several others succeeding him as Secretary of the Society.
On nine occasions during the period 16741687,
incidentally, the Delftman included with his letters
packets of samples of materials he had examined.
These were discovered and their dried contents
meticulously reexamined and described, for the first
time, by Ford (1985, and earlier notices cited
therein). It might also be mentioned here that
Leeuwenhoeks devoted daughter Maria, a few
weeks after his death and on his wishes, sent to that
august Society in London more than two dozen of
his best homemade microscopes. Alas, they (mis)managed to lose every single one of those precious
instruments over time.
In this paper, quite appropriately, I shall limit most
of our attention to the Letters, some quite lengthy,
that are concerned mainly with protistological or
bacteriological material, giving them the numbers
first assigned by Dobell (1932). Leeuwenhoeks principal communications wholly or in part on these
subjects were given numbers 6150+ and ran from
the year 1674 to 1716. Recall that numerous notes
were posted to other people or other destinations
than the Royal Society. Their writer, with very little
formal or classical higher education, nonetheless
truly became a man of letters!
Leeuwenhoeks observations on free-living protists and prokaryotes (bacteria) and on symbiotic or
parasitic species of these two major assemblages of
microorganisms were considered separately by Dobell (1932), but since information on both is intermixed in many letters I shall not make such a distinction in the present paper. In the second 30-year
period of the Father of Protistologys long life, Letters No. 6 through 71 were penned. During his last
30 years (see below), numbers 75 through 150 (plus
two more, numbered by Dobell as VII and XXIX) appeared.
In 1686, this man of growing fame sat for an oil
portrait by the distinguished painter Johannes
Verkolje, who had taken up permanent residence in
Delft a dozen years before then. A mezzotint engraving, made later the same year by the same artist and
introducing several small changes plus reversal of
the image in the print, is the colored picture most
often reproduced in major biographies and textbooks alike that have appeared during the past century (and see Fig. 1).

His Third 30 Years

In the last third of his life, Leeuwenhoek finally commenced to slow down a wee bit, but he still made
numerous contributions to the literature (again
mostly via publication of his Letters, in English
translation, in the Philosophical Transactions of the
Royal Society). The bulk of the relatively few drawings he made were published during the first half of
this period, with the remainder from the last third of
the preceding one (see Fig. 4). As mentioned above,
we are concerned here principally with the observations on protists and bacteria. The reader is referred
to Ford (1991) for discussion of his remarkable contributions to other fields similarly requiring microscopical techniques.
His last letter on protozoa (not illustrated) was
published in 1716, when he was 84 years old. But
recall that Leeuwenhoek was still sending scientific
letters to London on other topics during the final
year of his life, 1723!
By now, the Delft protistologist had achieved international (well, European) fame, and he not only
received many letters from admirers in all stations of
life but also requests to call on him personally.
Among the latter, ones most flattering to him personally were visits from royalty. For example, he entertained Queen Mary II of England, an Emperor of
Germany, and Peter I, Czar of Russia, bestowing microscopes upon all of them. To his joy, the learned
Peter the Great conversed fluently with him in the
Dutch language. Various distinguished scholars of
the time numbered among other visitors, as well as
several scoundrels who intended to copy his works
or steal his jealously guarded magnifying glasses.
Leeuwenhoek diagnosed his own terminal illness
more accurately than his physician did, who could
suggest only excessive palpitations of the heart. The
dying man himself concluded that he was suffering
from bronchopneumonia and so indeed he was,
passing quietly away in bed, with only daughter
Maria as family survivor, on 26 August 1723.

Leeuwenhoeks
Refined Single-Lens Microscope
The simple, postage-stamp size instruments with
which the Father of Protistology made his amazing
observations over a 60-year period in the 17th and
early 18th centuries were hardly complex ones (see
Figs. 2, 3), but they afforded a resolution and a clarity in magnification that clearly surpassed that of the
bulkier multi-lens compound microscopes of the
times (and for scores of subsequent years). But it is

Antony van Leeuwenhoek

181

Figure 2. The simple Leeuwenhoek hand-lens microscope, shown natural size. a. The front face (notice the
tiny aperture behind which is located the single lens). b. The back face, revealing the mechanics of the instrument. c. Side view, showing further aspects of its construction. See text for details. These pictures are copies of
original photographs taken of the authors personal brass replica of a typical Leeuwenhoek microscope and kept
in his collection of protistological memorabilia.

the latter kind of microscope that ultimately, with

many refinements, gave rise to our superior modern
instruments manufactured today by Zeiss, Leitz,
Spencer (American Optical), Bausch & Lomb, Olympus, Leica, and other companies. Thus there is good
reason (as stressed on a preceding page) not to
label Leeuwenhoek as the Father of Microscopy, despite his excellent results with his own special magnifying glass, so superior to other single-lens
scopes of then or ever since (see how/why
below).
Dobell (1932) and especially Ford (1985, 1991)
have offered such thorough descriptions of the
Leeuwenhoek scope that there is little need for me
to give many details (of its structure, etc.) here. But
review of a few facts will perhaps help the reader appreciate its uniquenesses and explain why its early
critics were completely in error to judge it to be impossible to make such discoveries as the Delft amateur microscopist did with this primitive tool (as
they imagined it).
Figure 3. Drawing of the front face of the authors authentic brass replica of a Leeuwenhoek microscope,
natural size, held in the hand of artist Lois Reid. Notice
its postage-stamp size!

Numbers and Sizes of the Microscopes

It is not generally realized that a great number of
these instruments were made by the Dutch draper

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J. O. Corliss

from Delft: hundreds is a safe estimate. Over the

years of his long life, dozens must have been given
away, with 26 alone to the Royal Society. How many
were donated or bequeathed to museums and such
repositories in The Netherlands, his own country,
does not seem to be accurately known. Some 250
completely finished forms were auctioned off to
Hollanders in 1747, almost 24 years after his death.
How many might have been discarded or disposed
of for various reasons, by their inventor or others, is
unknown. Simple though they may have been,
fashioning even one of them was not a short-time
task. Replicas (how many? hundreds?) have appeared over the years, some more faithful than others to original designs, and/but their whereabouts
are also not completely known or easily determinable. I am pleased to possess a fine example of
the latter kind myself, a personal gift from Dr. A.
Schierbeek when I was privileged to make his acquaintance briefly in 1951 in Amsterdam (see Fig.
2).
Sizes and shapes were not identical throughout
all the instruments known or still surviving (as originals or presumably faithful copies) either on purpose or sometimes supposedly by chance mostly
because an exact configuration of the frame (the
two body-plates, riveted together or held together
by screws) is not really essential from the point of
view of obtaining the best images of objects under
study. For example, using the nine surviving scopes
seen and measured by Ford (1991), lengths of the
always oblong plates ran from 32 to 47 mm, with
widths varying from 16 to 28 mm. Sometimes corners were squared, sometimes quite rounded; and
bottom halves of plates might be tapered to a
greater or lesser extent. My own facsimile has basically square corners, and both plates are roughly the
same size, measuring 42 mm (= 1 5/8 in.) by 23 mm
(= 7/8 in.), with no posteriad tapering (Figs. 2, 3).
It might be noted that one of last years U.S.A.
rectangular commemorative postage stamps, at
hand as I write this paper, measures 37 X 24 mm.
Except for the long coarse-threaded screw (for lowering and raising the stage on which the specimen is
located, to bring the latter into position opposite the
lens-hole), which extends a variable distance beyond the lower ends of the plates, the Leeuwenhoek
magnifying glass is indeed postage-stamp size. This
apt comparison, first suggested by Ford (1991), is
an excellent one, especially since most writers (including Dobell, and even Ford himself) publish photographs and drawings considerably and misleadingly larger than the actual instruments being portrayed. I have attempted to avoid that here (see Figs.
2, 3).

Major Parts and Functions of the Apparatus

Details are not needed here. Suffice it to say that the
clever craftsman Leeuwenhoek equipped his simple microscopes with simple parts that can nevertheless be used to carry out the very same functions
expensively taken care of today by accoutrements
supplied to our sophisticated compound scopes:
for examples, a mechanical stage and a manipulative object-carrier, focusing and positioning gears
and screws (for both fine and coarse adjustments),
and a path to a source of light (sun- or daylight; and
see below). The material to be examined could be
affixed directly onto the apical end of the object-carrier; or if living material required an aqueous
medium, a tiny handmade capillary glass tube could
be mounted on or glued to the carrier. For viewing,
the operator need only hold the tiny scope in his/her
hand and position an eye opposite the viewing aperture on the front face/side, working the positioning
screws as needed for sharpest focus. But notice
that the focal length, the working distance between
the eye and the object, has to be a very short one indeed, a disadvantage which even Leeuwenhoek
complained of as tiring to maintain properly over a
long period of time.
Left out above has been the single most important part of the entire instrument: the lens itself, of
course! For best results, it should be a tiny wellground and skillfully polished biconvex lens with a
resolving power approaching 1.0 m and a magnifying power (without spherical or chromatic aberration) of 200300+ diameters. It is held in place in a
socket between the appressed pair of body-plates
at the level of the pre-prepared aperture. No mirror is
necessary. Focus, and make and record your observations: cest tout! The lens in my replica allows a
magnification of only 30X, but that is sufficient to
observe and roughly identify quite a few species of
ciliates.
The kind of metal utilized for the plates is not of
importance. Leeuwenhoek employed all kinds available. The more expensive instruments, including the
ones often offered to select people as gifts, were
made of gold or, more likely, silver: both very pretty!
Copper was sometimes used, but probably most
commonly the metal chosen by the frugal inventor
himself was brass. My own copy is of brass.

Kinds of Lighting Employed

While Leeuwenhoek always advised visitors to look
towards a natural source of light (best would be simply daylight) or a lighted candle placed an appropriate distance away, it is now believed (see Dobells

Antony van Leeuwenhoek

1932 excellent reasoning and conclusions drawn on

the subject) that the Delft experimenter himself must
have discovered some simple means of realizing
darkfield illumination. Secretive man that he was, he
never imparted such information to anyone, but
some of his observations (e.g., seeing flagella on
bacteria) would seem to have required such a technique. Once again, the simple man from Delft had
brilliantly devised a way to enhance his critical viewing of organisms comprising the unseen world of microbiology.

Nature and Identification

of Small Organisms Observed
From the many accounts contained in the marvelous
sets of Letters sent to the Royal Society (and sometimes elsewhere), we can get a feel for the incredible
breadth of materials, biological and otherwise, so
painstakingly studied and so accurately described
by our amazing pioneer in the use of microscopy. In
the present paper, however (as stated elsewhere), I
am limiting mention primarily to microorganisms
today identifiable and classifiable as protists (mainly
the eukaryotic algae and protozoa) and bacteria
(prokaryotes), with emphasis on the former assemblage.
A strong note should be made of the fact that
Leeuwenhoek was by no means a nomenclaturist;
that is to say, he assigned no scientific names to the
wee organisms he described. This is not surprising,
of course (although people forget it), because he
lived and worked many decades before the entrance of the great Swedish botanist and nomenclaturist Carolus Linnaeus (also known as Carl von
Linn) upon the biological scene. The official dates
for the nomenclatural beginning of proper (Latin or
latinized) binomial descriptors for species of plants
and animals (together, at that time, embracing all
protists and prokaryotes as well as the higher eukaryotes) ultimately became 1753 and 1758, respectively (see current Codes of Nomenclature).

Celebrated Letter No. 6

In one of his earliest communications (dated 7
September 1674) to the Royal Society, our budding
protistologist gave clear, if brief, descriptions of the
phytoflagellate Euglena viridis (green in the middle
and before and behind white: Leeuwenhoek 1674),
the filamentous algal protist Spirogyra, and some
unidentifiable ciliates and flagellates, along with rotifers and other micro-inhabitants of a small freshwater lake not far outside Delft. This Letter No. 6

183

heralds the discovery of diverse protists, none ever

seen before, thus marking the birth of protistology itself. We owe knowledge of the existence of the note,
as well as its careful translation from the original
Dutch language, to the persistent and arduous twodecades-long investigative labors of the Englishman
Clifford Dobell (see Dobell 1932).

No. 18: Famous Letter on the Protozoa

This very lengthy multi-section epistle of Leeuwenhoeks (1677, but written in the fall of 1676), less
than half of which was published (in translation) by
Oldenburg, the editor, in the Philosophical Transactions, became the presumed First Letter on freeliving protists by the perceptive Delft observer, perhaps primarily because of Kents (1880) attention to
it as the first, in some five pages of his own highly influential leading volume on the Infusoria. But Kent
was familiar only with the parts that had appeared in
the Transactions. Dobell (1932) has supplied us filling more than fifty-four(!) full pages of his book with
an English translation of and comments on all sections of that early and indeed famous Letter on the
Protozoa, although mostly restricting his coverage
to paragraphs containing specific mention of protozoa, algae, and bacteria. Today we also have Schierbeeks (1960) illustrated 31-page booklet devoted
solely to that Letter 18, but it is written in Dutch.

Published Illustrations
While in the above and many subsequent Letters,
diverse species of protists and prokaryotes were described verbally with sufficient precision (including
helpful measurements) to allow their probable identification by Dobell or other protistological specialists, published figures appeared rather rarely; thus
we are often not allowed to have additional clues as
to the organisms structural composition. Unlike his
contemporary Robert Hooke and his biographer
Clifford Dobell, Leeuwenhoek was no draftsman. He
hired local artists, surely several over the last 60
years of his life, and/but they are never named in his
letters. He often had them look through his microscopes to confirm the images he himself was seeing; he also drew crude sketches for them to embellish. A few of his own direct attempts at providing
original drawings with his notes to the Royal
Academy may be seen occasionally along the margins of his script-pages housed in the Academys
files. A number of his illustrations (by draftsmen) are
reproduced in my Figure 4; and individual genera are
mentioned by their modern taxonomic names in the
immediately following section.

Figure 4. Leeuwenhoeks draftsmens figures (not drawn to a single scale) of diverse eukaryotic protists and prokaryotic bacteria. See text for detailed explanations, but brief identifications of the protists are given here. Under A (all figures redrawn and renumbered from Dobell 1932 and Corliss
1975): a, Anthophysa (chrysomonad); b, Volvox (chlorophyte); c, Coleps (prostome ciliate); d, Cepedea (opalinid); e, Nyctotheroides (heterotrich ciliate); f, Vorticella (solitary peritrich ciliate); g, Cothurnia (loricate peritrich); h, Carchesium (colonial peritrich); i, Elphidium (foraminiferan). Under B (all
figures redrawn from Dobell 1932): a-e, various spirochaetes and motile and non-motile bacilli and rods, all difficult to assign to exact genera with
certitude (see discussion in text).

184
J. O. Corliss

Antony van Leeuwenhoek

Identification and Taxonomy

of Wee Animalcules
This is not the place to treat the probable classification of Leeuwenhoeks protists and prokaryotes
(free-living or symbiotic, generally living material) in
any detail, but it might be interesting to note the
broad distribution of the forms that he described adequately enough for our later probable identification.
The higher-level taxonomic scheme of classification
employed below is based primarily on quite recent
overview papers of Cavalier-Smith (1993, 1998:
bacterial phyla included in the second) and/or of the
writer (Corliss 1994, 1998, 2000), with only minor
modifications; and for the ciliated protozoa, see
Corliss (1979) and Lynn and Small (1997, 2002).
Generic (and higher-level) names are indicated in
place. In some cases, the Delftmans precise descriptions have allowed us today to recognize the
very species he managed to see by means of his
simple scope; a few of these are included in the
present paper. Phyletic names are purposely given
in boldface script in the following paragraphs.
Kingdom BACTERIA. The prokaryotes are perhaps better broken up into several kingdoms, but
this is not of relevance to us here. Organisms in Figure B, a-e, are all from the human mouth. Phylum
Spirochaeta: several genera, but today exact
names of those seen and figured (e.g., see Fig. 4 B,
d) by our Delft microbiologist are difficult to assign;
quite possibly Treponema or Leptospira species
were involved. Additional motile spirochetes were
apparently noted in samples of polluted water; they
are similarly difficult for assignment of exact generic
names, although there is little doubt of their having
been sighted. Phylum Proteobacteria: members of
the enteric genera Escherichia and Spirillum? Phylum Selenobacteria: Selenomonas (Fig. 4 B, b,
showing path of movement through the medium).
Phylum Endobacteria: species of various genera
(including Bacillus and Streptococcus?) may have
been seen, even figured, but difficult to recognize
with taxonomic precision today from the supplied
information (Fig. 4 B, a, c, e). The anaerobic forms
he tantalizingly and inadvertently discovered in
sealed vials of soil-water and/but never studied
further (see below) may likely have included
Clostridium. Cyanobacteria (= the blue-green algae
in much of the literature, still today): some probably
seen, including Spirulina or the very common filamentous Oscillatoria. It should be noted and admitted! that seldom do microbiological textbooks
credit Leeuwenhoek with discovering any true
bacteria, free-living or symbiotic forms, that are recognizable nomenclaturally (at the generic or specific

185

level) today. Perhaps their caution is commendable.

The home laboratory of the Father of Microbiology
was obviously not equipped to isolate and cultivate
strains under aseptic conditions or to carry out the
sophisticated staining or physiological/biochemical/molecular tests required nowadays to determine
exact taxonomic or phylogenetic relationships of
any minute microorganisms under study.
Most of the organisms reported by the skilled single-lens microscopist from Delft are representatives
of all five of the eukaryotic kingdoms recognized by
Cavalier-Smith and the writer (see citations given
above). Many such species, even at the level of protists (essentially the only organisms treated below),
are at least one or two orders of magnitude larger or
lengthier than most of the common species or cells
of the prokaryotic bacteria mentioned above.
1. Kingdom PROTOZOA. Phylum Euglenozoa:
Euglena and the colorless kinetoplastid Bodo, both
free-living forms; plus Crithidia or Leptomonas (of
the trypanosomatids), parasitic in tabanid horseflies
examined. Phylum Dinozoa: likely some species
seen, but unnameable from information provided us.
Phylum Sporozoa (syn. Apicomplexa): Eimeria, in
the oocyst stage, in rabbit feces. Phylum Rhizopoda (exclusive of foraminiferans, next entry,
below): None! This is surprising, because diverse
freshwater and soil amoebae sometimes abound in
field collections. Perhaps most amoebae (that might
have been in Leeuwenhoeks samples) did not move
fast enough to attract his attention? Or were not
free-floating in the fluid in his tiny vials? Or were too
fragile to withstand his treatment? Nor did he seem
to note any free or encysted stages (too small?) of
symbiotic amoebae in the many samples of fecal
materials he examined at various times. Phylum
Foraminifera (if accepted as a separate phylum):
Elphidium, empty tests (see Fig. 4 A, i), found in
1700 in the stomach of a shrimp. Leeuwenhoek
seemed to be unaware or had forgotten Hookes
(1665) earlier account of a very small foram (but a
fossil form and of a different genus) found in sand.
But Hooke, in turn, did not cite Gesners (1565) fine
description of a similar although much larger fossil
form (described as a snail, a microcephalopod) a full
century earlier.
Phylum Ciliophora: Specimens were seen that
today are assignable to perhaps a dozen different
classes or subclasses of this phylum, the largest in
the kingdom Protozoa (if we exclude fossil protists).
Class Heterotrichea: Nyctotheroides (called Nyctotherus by Dobell and others: but split off from that
genus well after Dobells lifetime), from frog large intestine (see Fig. 4 A, c). Class Spirotrichea, subclass
Hypotrichia: Euplotes; subclass Stichotrichia: Oxy-

186

J. O. Corliss

tricha, Stylonychia, and Kerona, the last a very common ectocommensal found on the cnidarian Hydra;
subclass Oligotrichia: perhaps Halteria, a small
freshwater ciliate known for its rapid erratic, jumpy
movements through its medium. Class Litostomatea: Dileptus and Enchelys. Class Phyllopharyngea: Chilodonella. Class Colpodea: surely Colpoda,
so common as a soil or terrestrial ciliate, was seen
yet was never recognizably described. Class Prostomatea: Coleps (see Fig. 4 A, c). Class Oligohymenophorea, subclass Peniculia: Paramecium,
which Leeuwenhoek noted in conjugation, correctly
interpreted as a sexual copulation not longitudinal
fission as later workers (except O.F. Mller 1786)
kept insisting on for another 200 years; subclass
Scuticociliatia: Cyclidium; subclass Hymenostomatia: Colpidium and very likely the ubiquitous Tetrahymena; subclass Peritrichia: Carchesium, probably C.
polypinum (see Fig. 4 A, h); Cothurnia (see Fig. 4 A,
g) and Vaginicola, in their loricae; Trichodina, a mobiline peritrich that is another ectocommensal of
Hydra; and Vorticella (see three individuals in Fig. 4
A, f), whose attachment stalks were considered
tails. Regarding solitary vorticellids, our patient
observer first, in 1676, described them as wretched
creatures struggling in vain to disengage their bodies from debris in which their tails had become entangled; later, in 1713, he appreciated better the
lives of these attached protozoa, even ascertaining
the function of their peristomial ciliature (wheelwork, in his words) in feeding.
Members of phyla of zooflagellates, both free-living and symbiotic forms, were often noted by our
Father of Protistology and Parasitology, but exact
identifications, even at the generic level, are difficult
today. Some were probably colorless phytoflagellates that belong to the following kingdom (see
below). What the Delft microscopist saw in fecal material from frogs, rabbits, and chickens and in his
own diarrheric stools as well was of great interest
to Dobell, a recognized authority in parasitological
protozoology born 254 years after Leeuwenhoek
and also an avid and perceptive observer of wee
animalcules. Dobell (1932) insisted that his great
predecessor had certainly seen such tiny flagellates
as at least Chilomastix, Giardia, Hexamita, Trichomastix, and Trichomonas (of phyla Metamonada and Parabasala).
2. Kingdom CHROMISTA. This is essentially the
Stramenopila of some workers. Phylum Chrysophyta: Anthophysa vegetans (see Fig. 4 A, a),
Spumella (formerly Monas), and Cercomonas, all
colorless chrysomonad flagellates with the first a
colonial species with its stalks typically encrusted
with brown particles of ferric hydroxide (Leeuwen-

hoeks accurate depiction is even better than that of

the great O.F. Mller 1786 its first rediscoverer
some 80 years later: contrast Fig. 3 of Corliss 1986
with Fig. 4 A, a of the present paper). Other very
small colorless flagellates, unnameable, may also
belong here. Phylum Diatomeae: unnameable diatoms. Phylum Opalinata: Cepedea, from frog large
intestine (see Fig. 4 A, d).
3. Kingdom PLANTAE. Only its algal groups are
considered here. Phylum Charophyta (class Conjugatophyceae): Spirogyra. Phylum Chlorophyta:
Volvox (see Fig. 4 A, b), Chlamydomonas, Haematococcus, and Polytoma.
4. Kingdom FUNGI. Members of protist groups
that are placeable here, all parasitic/symbiotic forms
(chytrids, Microspora), werent described by
Leeuwenhoek. But he did observe minute fungal
spores and some hyphal forms having unicellular
stages in their life cycles; for example, Phylum Ascomycota (class Saccharomycetes): Saccharomyces, some species of which are the common
brewers yeast.
5. Kingdom ANIMALIA. Members of protist groups
that are placeable here, for example, the all-parasitic
Myxozoa, were not seen by Leeuwenhoek. Many
very small, sometimes microscopic, invertebrate animals, however, were observed, including Hydra,
various rotifers, worms, insects (larvae and adults),
etc., multicellular organisms of phyla well beyond
coverage in the present paper.

Misconceptions of Leeuwenhoek
and his Works
Some of the misconceptions or misunderstandings
of our modest microbiologist from Delft have already
been alluded to on various preceding pages. During
his own lifetime his observations were, perhaps understandably, often met with skepticism. Being the
first person to reveal the theretofore hidden world of
the microcosm and the unlikely discoverer not
even a learned man with advanced degrees and/or a
professorship in a great university (tsk, tsk!) was
not a particularly enviable position in which to expect to find oneselfs being taken seriously or credibly. Fortunately for posterity, Leeuwenhoek was in
due time accepted, through backing by the eminence of the Royal Society of London and by such
contemporary recognized scientists as Robert
Hooke and, much later, such distinguished and authoritative protozoologists as Btschli and Kent. But
as the Delftman philosophically wrote in one of his
letters in the autumn of his life, It doesnt strike me
as odd that I meet with contradictions. This was in

Antony van Leeuwenhoek

response to the comment by an elderly acquaintance in his town who had sadly but kindly prophesied, My friend, youve got the truth but it wont be
received in your lifetime (from translations in Dobell
1932).
Leeuwenhoek was definitely a man ahead of his
time. Perhaps saddest have been the widespread
misunderstandings still present some 200 years
after his death and even occasionally still today,
despite the interest rekindled in recent decades. Dobell (1932) often rather scathingly brings up scores
of mistakes and inaccuracies, even ones quite minor
in nature, and he lists >25 variants in spellings of the
mans names (for which careless Dutch, English,
French, and German writers are blamed)!
One amusing example of a postdobellian slipup can be found in the deservedly popular book (its
title inspired by a well-known ditty of Jonathan
Swifts, who, by the way, lived in the time of
Leeuwenhoek) by the distinguished American protozoologist-parasitologist Robert Hegner (1938), who
depicted our good Father of Protistology peering intently through an elaborate compound microscope
set-up used by Robert Hooke and seeing amoebae,
parasitic trypanosomes, and other protists that were
favorites of Hegners. But actually neither these organisms nor that particular compound scope were
ever seen by the man from Holland! Somewhat similarly and Dobell would turn over in his grave if he
knew of this one the Dover unabridged paperback
edition of Dobell (1932) has superimposed on its
cover-portrait of Leeuwenhoek a parade of amoebae pseudopoding their way across the page: but
our hero never saw (or at least, never described)
such wee animalcules.

Leeuwenhoeks Legacy and His Honors

I have briefly reviewed the life, works, and letters of
the Discoverer of Protists on preceding pages. We
have yet to consider a bit further the qualities of his
character and the overall breadth of his scientific interests and personal goals. The appropriate words
Leeuwenhoek legacy, used in the titles of the insightful book by the super-sleuth Brian Ford (1991)
and of the recent article by the ecologically oriented
team of Finlay and Esteban (2001), pay deserved
tribute to the natural philosophy of the kind
Leeuwenhoek practiced [which] was rooted in a desire to understand, to clarify, and to communicate
(Ford, 1991). No flowery language, no obscurantism,
no exaggerated sense of scientific superiority, which
qualities, in Fords opinion, underpin much [of]
modern science. [Ouch!]

187

Honest and unassuming himself, Leeuwenhoek

abhorred deceit, and if he has been remembered as
a cranky old man it was because he did not suffer
contemporary fools or rogues gladly. He was not a
person intent on hypothesizing. Instead, he was
much like our diligent Philadelphian Joseph Leidy of
the 19th century, in this as well as other respects
(Corliss 2001) content with finding facts and letting
others make wild speculations or draw lofty conclusions.
The above point is well illustrated in the Delftmans long overlooked, even if minor, contribution to
the much later eventual overthrow of the abiogenesis doctrine (= spontaneous generation), that life
could arise from inanimate materials. Having heard
of the treatise by Francesco Redi (published in Italian in 1668), he decided, in 1680, to carry out some
trials himself. Working patiently, he discovered that
no (metazoan) life arose in periodically examined
tubes filled from various muddy or polluted aquatic
sources. But if such tubes in which seeds were
present were sealed, then the seeds themselves
could multiply into teeming populations of tiny organisms. He thus, unwittingly, discovered anaerobic
bacteria, but, alas, he never followed up on such
work. The reader will recall that Spallanzanis celebrated experiments on spontaneous generation did
not take place until nearly a full century later, and it
was not until still another 100 years had passed before Pasteur and Tyndall convincingly put the final
nails to the coffin of abiogenesis (although with the
passage of yet another 150 more years, we do find
today a few uneducated folk clinging to the attractive ancient myth of instant life spontaneously reoccurring or arising or being generated from non-living
things). In fairness to Leeuwenhoeks enterprising
contemporary Louis Joblot (born 1645, died 1723) in
Paris, I should mention that the Frenchman, working
independently a bit on abiogenesis himself, was the
first to boil some infusions and then keep them in
closed containers: neither microorganisms nor other
living material were ever found on his later (re)examination of those infusions. This little experiment (described in Joblot 1718), like the earlier one by our
Dutchman, has been widely overlooked by historians of science.
Remember that our first inquisitive microbiologist
was not in the business, as it were, of confirming
earlier observations of predecessors: in any case,
almost always, there were no predecessors! To the
category of originality, beyond his morphological
descriptions of protists, we must also assign his
works on feeding by protozoa, the coiling and uncoiling of stalks of attached ciliates, conjugation in
protozoa as a sexual phenomenon, reproduction in

188

J. O. Corliss

Volvox, asexual fission in a number of protistan

forms, budding in Hydra, discovery of parthenogenesis in aphids, avoidance reactions of unicellular organisms confronted with certain chemicals, ecological preferences of microorganisms, capillary circulation of blood cells, comparative morphology of invertebrate spermatozoa, structural histology of
plant and animal tissues, etc.
In protistology he was interested in and contributed to knowledge of his organisms biodiversity,
several full centuries before that term became a
fashionable household buzz word, following E.O.
Wilsons (1992, and see earlier papers cited therein)
relatively recent introduction of it into our modern
ecological literature at all levels in plant/animal systematics.
Another important way in which Leeuwenhoek
made original contributions to the world of the invisible: he was the first conscientious measurer or
metrologist (Dobells word) of his observed very
tiny objects. Perhaps he got into the spirit of such a
task with microbes from his experiences as the
town surveyor (he accurately figured out the height
of a local church steeple) or even more so as a
drapery merchant, which involved much careful
measuring of goods for customers. We have to remember that meters (and thus micrometers) had
not yet been invented; the lowest unit of measurement in those times was the inch (with his inch
0.75 mm longer than ours of today, according to
Dobells careful calculations). The Delftman divided
his homemade inches-ruler into tenths. The next
problem was to select tiny measurable-by-thenaked-eye objects that had quite a uniform length
or diameter: his most popular choices were sand
grains (fine and coarse), millet seeds, vinegar eels (a
nematode), and the eye of a louse (the one commonly found, still today, on the human head).
Armed with such measurements, expressed in precise fractions of an inch, he was thus enabled to
state that a given protist or cell measured a still
tinier fraction of that fraction. One finds that such
final figures are often very close approximations to
todays known average lengths or diameters (in micrometers) of the protists in question. Two examples will suffice in illustration of this point. A common species of the small colorless chrysophyte
genus Spumella (called Monas by Dobell and others) comes out with a diameter of about 1215 m;
and a stichotrichian spirotrich ciliate of the genus
Stylonychia, with a length of about 140 m. Remarkably accurate measurements!
In preceding pages, I have mentioned many of the
honors mostly very belatedly bestowed on Antony
van Leeuwenhoek for example, his Father of ti-

tles. During his busy lifetime of so long ago, his

prime recognition as an early microscopist of truly
unbelievable ability came with his appointment, at
the age of 47 (in 1680), as a Fellow of the esteemed
Royal Society of London, for which he was always
grateful. Reproductions of his simple hand lens
from time to time have reminded us of the marvels of
his inventiveness, and we need to thank Brian Ford
(1985, 1991) for his brilliant detective work in gathering data on and even demonstrating its effectiveness in revealing the hidden world of invisible sights.
In his 84th year, 1716, the devoted amateur scientist received a silver medal from the University of
Louvain, now in Belgium, in recognition of his never
yet properly appreciated and celebrated discoveries
in Natural Philosophy. Quite emotionally, the aged
but still active recipient replied (in the introductory
part of a lengthy thank-you letter: see Dobells 1932
translation), When I think on the flatteries expressed in your letter I dont only blush, but my eyes
filled with tears too, especially because my work
was not pursued in order to gain the praise I now
enjoy, but chiefly from a craving after knowledge,
which I notice resides in me more than in most men.
And there-withal, whenever I found out anything remarkable, I have thought it my duty to put down my
discovery on paper, so that all ingenious people
might be informed thereof.
There are other continuing reminders of the Father of Microbiology sensu lato in addition to inclusion of his name and portrait in biological textbooks
and histories of science. A microbiological journal,
Antonie van Leeuwenhoek, has been regularly published in The Netherlands. And a coveted Leeuwenhoek Medal has been awarded every decade or so
during the past 125 years by the Royal Academy of
Sciences of Amsterdam to distinguished microbiologists around the world. The list is too long to cite
here in full. The first recipient was, fittingly, C. G.
Ehrenberg (see Schlegel and Hausmann 1996); and
about 65 years ago again highly appropriately
the awardee was C. Dobell, the man who so
poignantly brought Leeuwenhoek back to life for all
protistologists today.
Thus lived, died, and finally immortalized that
17th century indefatigable Dutchman, a gifted and
most versatile single-lens microscopist with such
tremendous patience and dexterity and such incredible eyesight. Above all, he was a person with an insatiable curiosity about the natural wonders of the
fascinating microcosm surrounding us. In conclusion of this brief tribute to Leeuwenhoek, allow me
(inspired by a term used by Woodruff 1938) to propose one more accolade for him, viz., First Philosopher in Little Things.

Antony van Leeuwenhoek

References
Beltrn E (1974) Notas de historia protozoolgica. III.
Leeuwenhoek y el tricentenario del descubrimiento de
los protozoarios. An Soc Mex Hist Cienc Tecnol 4:
225258
Bradbury S (1967) The Evolution of the Microscope.
Pergamon Press, London
Bulloch W (1938) The History of Bacteriology. Oxford
University Press, London (reprinted in 1960)
Btschli O (18801889) Protozoa. Abt. I-III. In Bronn
HG (ed) Klassen und Ordnung des Their-Reichs, CF
Winter, Leipzig, 1: 12035
Btschli O (1887) Historische Entwickelung der Infusorienforschung. Pp 11001195 of Abt. III (see the immediately preceding reference, above)
Cavalier-Smith T (1993) Kingdom Protozoa and its 18
phyla. Microbiol Rev 57: 953994
Cavalier-Smith T (1998) A revised six-kingdom system of life. Biol Revs 73: 203266
Cole FJ (1926) The History of Protozoology. University
of London Press, London
Cole FJ (1937) Leeuwenhoeks zoological researches.
Parts I, II. Ann Sci 2: 146, 185235
Cole FJ (1938) Microscopic science in Holland in the
seventeenth century. J Quekett Micros Club (series 4)
1: 5978
Corliss JO (1975) Three centuries of protozoology: a
brief tribute to its founding father, A. van Leeuwenhoek
of Delft. J Protozool 22: 37

189

Corliss JO (2000) Biodiversity, Classification, and

Numbers of Species of Protists. In Raven PH, Williams
T (eds) Nature and Human Society: the Quest for a
Sustainable World. National Academy Press, Washington DC, pp 130155
Corliss JO (2001) Two most remarkable Amoeba men:
Joseph Leidy (18231891) of Philadelphia and Eugne
Penard (18551954) of Geneva. Protist 152: 6985
Dobell C (1923) A protozoological bicentenary: Antony
van Leeuwenhoek (16321723) and Louis Joblot
(16451723). Parasitology 15: 308319
Dobell C (1932) Antony van Leeuwenhoek and His
Little Animals: Being Some Account of the Father of
Protozoology and Bacteriology and His Multifarious
Discoveries in These Disciplines. Swets and Zeitlinger,
Amsterdam (reprinted in an unabridged paperback
edition in 1960 by Dover Publications, Inc., New York)
Finlay BJ, Esteban GF (2001) Exploring Leeuwenhoeks legacy: the abundance and diversity of protozoa. Int Microbiol 4: 125133
Ford BJ (1985) Single Lens: the Story of the Simple Microscope. Harper and Row, Publishers, New York
Ford BJ (1991) The Leeuwenhoek Legacy. Biopress
and Farrand Press, Bristol and London
Gesner K (1565) De Omne Rerum Fossilum Genere.
Tiguri
Hegner RW (1938) Big Fleas Have Little Fleas or
Whos Who Among the Protozoa. Williams and Wilkins,
Baltimore (reprinted in an unabridged paperback edition in 1968 by Dover Publications, Inc., New York)

Corliss JO (19781979) A salute to fifty-four great microscopists of the past: a pictorial footnote to the history of protozoology. Parts I and II. Tr Am Microsc Soc
97: 419458; 98: 2658

Hooke R (1665) Micrographia or Some Physiological

Descriptions of Minute Bodies Made by Magnifying
Glasses with Observations and Inquiries Thereupon.
1st ed. Martyn and Allestry, London

Corliss JO (1979) The Ciliated Protozoa: Characterization, Classification, and Guide to the Literature. 2nd ed.
Pergamon Press, London and New York

Hooke R (1667) Micrographia 2nd ed. Martyn and

Allestry, London (reprinted in an unabridged paperback edition in 1961 by Dover Publications, Inc., New
York)

Corliss JO (1986) The 200th anniversary of O.F.M.,

1786: a tribute to the first comprehensive taxonomic
treatment of the protozoa. J Protozool 33: 475478
Corliss JO (1992) Historically important events, discoveries, and works in protozoology from the mid-17th
to the mid-20th century. Rev Soc Mex Hist Nat 42 (year
1991): 4581
Corliss JO (1994) An interim utilitarian (user-friendly)
hierarchical classification and characterization of the
protists. Acta Protozool 33: 151
Corliss JO (1998) Classification of Protozoa and Protists: the Current status. In Coombs GH, Vickerman K,
Sleigh MA, Warren A (eds) Evolutionary Relationships
Among Protozoa. Kluwer Academic Publishers, Dordrecht, pp 409447

Joblot L (1718) Descriptions et Usages de Plusieurs

Nouveaux Microscopes avec de Nouvelles Observations Paris
Kent WS (1880) Introductory General History of the
Infusoria Pp 130 of Vol. I (see the immediately following reference, below)
Kent WS (18801882) A Manual of the Infusoria: Including a Description of All Known Flagellate, Ciliate,
and Tentaculiferous Protozoa, British and Foreign, and
an Account of the Organization and Affinities of the
Sponges. Vol. I-III. David Bogue, London (vol. III consists solely of his plates of figures)
Lechevalier HA, Solotorovsky M (1965) Three Centuries of Microbiology. McGraw-Hill, New York

190

J. O. Corliss

Leeuwenhoek A van (1674) More observations from

Mr. Leewenhook, in a letter of 7 September 1674, sent
to the publisher. Philos Tr Roy Soc 9: 178182

Schierbeek A (1959) Measuring the Invisible World:

the Life and Works of Antoni van Leeuwenhoek.
Abelard-Schuman, London and New York

Leeuwenhoek A van (1677) Observations communicated to the publisher by Mr. Antony van Leewenhoeck, here Englishd, concerning little animals by him
observed in rain-, well-, sea- and snow-water, as also
in water wherein pepper had lain infused. Philos Tr Roy
Soc 12: 821831

Schierbeek A (1960) De van Leeuwenhoek-Brief van 9

Oktober 1676, de Geboorte van de Microbiologie.
Konink Nederl Gist- en Spiritusfabr. N.V. Delft (in Dutch)

Locy WA (1935) Biology and Its Makers. 3rd ed. Holt,

Rinehart and Winston, New York

Schierbeek A (ed) (1939?) The Collected Letters of

Antoni van Leeuwenhoek. Vols 1?. Swets and
Zeitlinger, Amsterdam (possibly not yet completed set
of the Letters, being published simultaneously in English and Dutch)

Lynn DH, Small EB (1997) A revised classification of

the phylum Ciliophora Doflein, 1901. Rev Soc Mex Hist
Nat 47: 6578

Schierbeek A (1963) Antoni van Leeuwenhoek en Zijn

Voormaamste Ontdekkingen. Kruseman, Den Haag (in
Dutch)

Lynn DH, Small EB (2002) Phylum Ciliophora Doflein,

1901. In Lee JJ, Leedale GF, Bradbury P (eds) An Illustrated Guide to the Protozoa: Organisms Traditionally
Referred to As Protozoa, or Newly Discovered Groups.
2nd ed. Society of Protozoologists, Lawrence, Kansas,
pp 371656

Schlegel M, Hausmann K (eds) (1996) Christian Gottfried Ehrenberg-Festschrift. Leipziger Universittsverlag, Leipzig

Meyer AW (1937) Leeuwenhoek as an experimental biologist. Osiris 3: 103122

Singer CJ, Holmyard EJ, Hall AR (eds) (1957) A History of Technology. Vol. 3. Clarendon Press, Oxford

Mller OF (1786) Animalcula Infusoria Fluviatilia et Marina, Quae Detexit, Systematice Descripsit et Ad Vivum
Delineari Curavit. Havniae et Lipsiae (a posthumous
publication)

Wilson EO (1992) The Diversity of Life. Harvard University Press, Cambridge

Nordenskild E (1928) The History of Biology: A Survey. AA Knopf, New York and London (reprinted in
1935 and 1946)

Woodruff LL (1939) Microscopy before the nineteenth

century. Am Nat 73: 485516

Rooseboom M (1956) Microscopium. Leiden

Singer CJ (1959) A History of Biology to about the Year

1900: A General Introduction to the Study of Living
Things. 3rd ed. Abelard-Schuman, London and New York

Woodruff LL (1938) Philosophers in little things. Univ

Okla Bull No. 739: 2133

Zuylen J van (1981) The microscopes of Antoni van

Leeuwenhoek. J Microsc 121: 309328