The Journal of

Protozoology

August, 1912

Volume 19

Number 3

J. PROTOZOOL.
19(3), 389-400(1972).

A Man to Remember, E. FaurC-Fremiet (1883-1971):

Three-Quarters of a Century of Progress in Protozoology*
Department

of

JOHN 0. CORLISS
Zoology, University of Maryland, College Park, Maryland 20742

SYNOPSIS. The late Professor Emmanuel FaurC-Fremiet, nearly 88 years old at the time of his passing in November
1971, was not only an outstanding leader in ciliate protozoology but also a general cytologist, experimental embryologist,
and electron microicopist of note, and, in earlier years, a sculptor and painter of merit. Both born and marrying into celebrated families, Monsieur FaurC led a career of his own filled with honors and distinction, but he remained a sensitive,
modest man, remembered, too, for his ever-present sense of humor. Author of nearly 500 books and papers in diverse fields
of biology, FaurC-Fremiet made his most lasting contributions in the realms of synthesis, innovative hypotheses, and
heuristic ideas.
Index Key Words: Faur&Fremiet, E., in memoriam.

C E.

YTOLOGIST, embryologist, electron microscopist, Professor

FaurC-Fremiet was foremost and above all a protozoologist, with a distinguished scientific career spanning over
70 years, from the discovery of mitochondria to elucidation of
finer aspects of their ultrastructural organization. Brilliant
theorist, inquisitive naturalist, keen observer, patient and persistent, vivacious yet sensitive in spirit, endowed with a well
known sense of humor but at the same time a model of
modesty, this great man made contributions to every field of
endeavor which he entered, including areas well outside those
of strictly biological research.
The purpose of the present tribute is to emphasize the impact
of this remarkable Frenchman on the whole science of protozoology, noting in passing that his professional involvement in
this area included activities within the relatively youthful Society of Protozoologists (currently celebrating its 25th anniversary). But some attention must also be given to other
aspects of his very full life and to the influence he has had on
many persons working in diverse fields around the world.

Some Biographical Notes

Emmanuel F a d - F r e m i e t was born 29 December 1883, the
elder son of the celebrated composer and organist Gabriel
F a d , a man particularly remembered today for his moving
Requiem Mass, his many songs and chamber music, his pupil
Ravel, and his friendship with Saint-Saens, his teacher. FaurC
the father often had to be absent from Paris on various musical
journeyings, so young Emmanuel spent many enjoyable hours
in the studio of his maternal grandfather and namesake, the
famous animal sculptor Emmanuel Fremiet, whose perhaps bestknown work-the golden equestrienne statue of Jeanne dhcstands in the Place des Pyramides near the Louvre, with copies
in various parks around the world, including one in the center
of Philadelphia. By his fathers arrangement, little Emmanuel
and his brother Philippe legally adopted the compound surname
The publication of this homage to the late Professor E. F a d Fremiet was supported by the Society of Protozoologists.

FaurC-Fremiet, particularly touching since Fremiet had had

no s0ns.l
In March 1913, E. FaurC-Fremiet married Jeanne Henneguy,
one of two daughters of the distinguished cytologist and microscopist Louis-FClix Henneguy, long-time Professor at the Colltge de France and himself son of the author and playwright
FClk Henneguy. O n their mothers side, the sisters Jeanne
and Suzanne were grandchildren of Pierre-Joseph Proudhon,
fiery and prolific 19th century revolutionary socialist leader
and philosopher.
Alas that the union of Emmanuel and Jeanne FaurC-Fremiet
was not blessed with children: the end of generations of genius.
After a very long and happy life together (and, in later years,
sharing their apartment with unmarried sister Suzanne), Madame
Faurt-Fremiet passed away in June 1967; Suzanne, in March
1970.
I n June 1971, at the age of 87 and scarcely 5 months before
his death, Monsieur FaurC, as he was widely and fondly
known by scientific colleagues, his staff, and former students
(and as he often will be called subsequently in the present
paper), remarried. The second Madame Fad-Fremiet, who
survives him, had been for many years his faithful secretary,
research collaborator, and devoted friend of the family (as
was her mother, who passed away in October 1968), Mademoiselle Marie Hamard, a name well-known to ciliate protozoologists.
Brother Philippe, biographer, playwright, essayist, and critic,
passed away in 1954, at a relatively youthful age when compared with other members of the family lineages mentioned
above. Also childless, he is survived by his wife Blanche (nCe
Felon, a family distinguished in the arts in its own right), who

Interestingly enough, Emmanuel FaurC-Fremiet resembled his

grandfather Emmanuel Fremiet in certain physical features much
more closely than he did his father, although considerably shorter
in stature and apparently more animated in facial expression (a
livelier twinkle in the eye). On the other hand, he seemed to
possess his fathers dark complexion, short wiry body, and, sadly
enough, susceptibility to bronchitis,

389

390

E. FAURE-FREMIET (1883-1971)

Fig. 1. Professor FaurC-Fremiet at his desk in the Coll&gede France (ca. 1950), just before his second major trip to the United
States. Notice the pipe, a constant companion, and the little vase, with three flags, on the bookcase over his shoulder. (Photo by
Dr. Jean Dragesco.)
still resides in the ancestral Fremiet (and, later, FaurC) treasurefilled home at 32 Rue des Vignes in the western outskirts of
Paris (arr. XVIe).
FaurC-Fremiet died on 6 November2 1971, a little over a
month before his 88th birthday. Confined to his long-occupied
apartment, rich in memories, at 9 Rue ThCnard (less than 75
feet from the entrance to the College de France) for only a
few weeks before his death, suffering principally from emphysema and bronchitis, his passing was a mercifully quiet one.
Dear Madame Marie FaurC-Fremiet, patient and self-sacrificing, was constantly by his side, as she had been ever since their
marriage, giving him great comfort to the very end. He is
buried in the Henneguy family cemetery at Prtcy-sur-Oise.

Professional Career
For essentially the whole of his scientific life, Monsieur FaurC
was associated with the College de France, that now long-established and well-known institution which formally broke away
in 1543 from the prestigious Sorbonne (the original University
of Paris) on the other side of the Rue St. Jacques. He suc I t is curious that the father and both of his sons all died in
the same month and within a few days of one another. Gabriel
passed away on 4 November 1924; Philippe on 19 November
1954.

ceeded his father-in-law, L. -F. Henneguy, to the Chaire 8Embryogenie ComparCe of the College de France and to Directorship of the Laboratoire, in 1928, holding those important
posts until his obligatory retirement in 1955, at the age of 72.
(When Professor Etienne Wolff, embryologist from Strasbourg,
next assumed the Chair, incidentally, the name of the laboratory
was changed to Laboratoire dEmbryologie Exptrimentale.)
A modest, unaggressive man, far more interested in his microscope, his experiments, and his cells or organisms than in details of protocol, administration, or so-called professional advancement, Monsieur Faurt never bothered with acquisition of
a Docteur es Sciences Naturelles until rather firmly (as the
story goes) urged to do so by his Maitre and administrative
superior, Professor Henneguy. He took his doctoral degree
in 19243 from the University of Paris with ease, having already
published 189 papers in diverse fields of biology! Thus he was
eligible, a scant 4 years later, to accept the Chair and the
Directorship of the Laboratory.
Little total time in his long career was spent outside France.
After 8 or 9 months every year in Paris, FaurC-Fremiet typically
fled to coastal biological stations, especially Concarneau in
Brittany, in the summertime. He worked on various occasions

He dedicated his thesis ( 7 ) to the memory of his grandfather

Fremiet who passed away in 1910.

E. FAURB-FREMIET (1883-1971)

391

special guest of honor at Americas first all-Faurt piano recital, given by Miss Alstadter under the patronage of Alice B.
Tully. In Washington he visited the National Academy of
Sciences, greeted by President Handler; and he was given a
memorable reception at Howard University through the efforts
principally of Dr. Harold E. Finley (see Fig. 3).
After his official retirement from the CollPge de France,
FaurC-Fremiet was, in due time, given the title Professeur
honoraire (in 1957) and became, if possible, more active in
research than ever, particularly excited by his thorough immersion in the budding field of electron microscopy. Practically
until the month of his death, he worked either at the College
or at his nearby home or at the CNRS center for hydrobiological research at Gif-sur-Yvette, with summers frequently
at Concarneau.

Principal Honors

Fig. 2. Monsieur FaurC as he appeared (note pipe in hand)

about 1930, just after return from his first trip to America.
(Photographs in Figures 2, 4, and 6 were obtained through the
kindness of Mme Marie FaurC-Fremiet.)
at Roscoff, Banyuls-sur-Mer, and other stations. He also
attended meetings and gave invitational seminars at various
times in certain of the major centers of biological research in
such European countries as England, Poland, Italy, and nearby
Belgium and Holland, but relatively rarely over his long
career.
In 1929, 1933, and 1950, with the aid principally of Rockefeller Foundation and Commonwealth Fund grants, extensive
trips were made to North and South America, with considerable
time spent at the Marine Biological Laboratory, Woods Hole,
Massachusetts (summers 1929, 1950) and at SHo Paulo, Brazil
(1933). (It was during his 1950 visit to the States that I had
an opportunity to meet and talk with this great man, through
the kindness of my professor at New York University, the late
Professor R. P. Hall. It was also then that Monsieur FaurC
extended me the kind invitation to spend a full postdoctoral
year, 1951-52, in his laboratories, an experience which turned
out to be a most rich and unforgettable one.)
A young New York City pianist who specializes in playing
Gabriel Faur& compositions (Judith Alstadter) and I arranged
a short, final, and highly nostalgic visit for FaurC-Fremiet to
the United States in March 1970. His itinerary included only
New York and Washington, D. C. areas, but a number of old
1929 Woods Hole acquaintanccs, as well as a host of more
recent colleagues and admirers, met with him and paid him
homage. At the Lincoln Center, New York City, he was the

An accurate measure of this modest but highly intelligent

mans impact on the scientific world may be seen in the long
list of honors conferred on him during his lifetime. He was
active in a number of biological societies in France and abroad,
often an officer or member of an editorial board, but space
does not permit listing all of these. He was involved in international groups, too, from early work as Secretary of the International Union of Biological Sciences (see 6 ) , during the
formative years of IUBS, to association with various international societies or congresses on cell biology, embryology, and
protozoology.
His highest honors were the unsolicited (with the exception
of the French Academy itself, whose system he found quite
distasteful) elections to membership in the most prestigious
academies and learned societies in the world (see Fig. 4).
Principal ones among these are cited briefly below in chronological order, with indication of the year of Monsieur FaurCs
election to them: Membro Correspondente dii Academia Brasileira dis Ciencias (Rio de Janeiro)--1933; Honorary Foreign
Member of the American Academy of Arts and Sciences
(Boston)-1938; Membro Straniero della Accademia Nazionale
dei Lincei (Rome)-1952;
Membre Etranger de 1AcadCmie
Royale Flamande des Sciences, des Lettres et des Beaux Arts
(Brussels)-l955; Membre de 1Institut de France, AcadCmie
des Sciences (Paris)-l959; Foreign Fellow of the Royal Society (London)-1963; Foreign Associate of the National Academy of Sciences (Washington)-1969.
In the field of protozoology, as might be expected, FaurtFremiet was active in both national and international societies
and congresses. This included membership on the Executive
Committee (years 1953-1955) and Editorial Board (years 1954,
date of launching of The Journal of Protozoology, through 1967)
of the Society of Protozoologists; and, in 1958, election to Honorary Membership in that society. He was Honorary President
of the 2nd International Conference on Protozoology, held in
London in 1965. In France he was a founding father, in 1961,
of the Groupement des Protistologues de Langue Franqaise.
In August 1971, belatedly in more than one respect, the American Microscopical Society elected him to Honorary Membership, principally and appropriately in recognition of his 65
years of outstanding work on the cytology of protozoa.

Breadth of Contributions
Leaving works in nonscientific fields aside here (but see a
subsequent section, below), the contributions of FaurC-Fremiet
are still amazingly broad, even within the biological sciences.
His publications alone (nearly 500 in number) may be classified

392

E. FAURfi-FREMIET (1883-1971)

Fig. 3. Monsieur FaurC with Dr. Harold E. Finley and the author, on the occasion of a reception in his honor at Howard
University (March 1970), during his last trip to the United States. (Photo obtained through the kindness of Dr. Donald M.
Spoon.)
into several diverse subfields of biology, the scheme of the
classification depending on the interest of the person doing it
and/or the fineness of the subdivisions. Monsieur FaurCs own
scheme (in his unpublished Titres et Travaux Scientifiques)
will not be followed here because its 8 subdivisions involve so
much overlapping, though some is obviously inevitable in any
arrangement.
In his own research, in that of his students and laboratory
guests, and in his 32-year-long series of cours professCs at the
Coll&ge de France, FaurC-Fremiet may be said to have made
major contributions to 4 principal areas of biology, simply defined as follows:
Cytology, with emphasis on cellular constituents and the
protoplasmic structure of living matter as revealed by light and,
later in his career, transmission electron microscopy. Here, too,
one might include work on species of bacteria, algae, and nonciliate Protozoa, for want of a more appropriate place in the
present scheme, but exclude the ciliates, which are given a
separate area below. Selected examples of major contributions in this vast field are included among the following references: 2, 4, 15, 18-20, 27, 31, 36, 43, 61-63, 67, 69, 70, 72,
118, 132, 133, 172.
Developmental biology, including experimental embryology,
tissue culture, physiology, X-radiation, diverse problems of
growth and development, and the kinetics of living matter.

Arbitrarily excluded are the ciliate Protozoa; thus metazoan

material is emphasized (e.g., see 5, 8, 9, 12, 43, 60, 61, 64, 68,
69, 71, 76, 77, 81, 118).
Fibrous structures, the physical chemistry of fibrous proteins.
Wherever such structures may be found in biological materials,
they held an enduring fascination for Monsieur FaurC throughout the 70-odd years of his active scientific life. Most of .his
observations were made on protozoan material (see following
section), but, among those which were not, 4 representative
works are included in the bibliography (17, 80, 82, 83).
Ciliate protozoology, an area so predominating, especially
during the last 25-30 years of his research output, that it requires its own subdivisions and thus more extended treatment
(see below).
In all of his works, though no shirker in the gathering of
massive quantities of original data via both field and laboratory
observations, FaurC-Fremiets most memorable contributions lie
in the realms of synthesis, of innovative hypotheses, of heuristic
ideas. Great embryologists, for example, men such as the late
Ross Granville Harrison of Yale University and Paul Weiss of
Rockefeller, have mentioned how excited they were, turned
on, in modern parlance, by such treatises as his little-publicized
but profound book La Cinitique du Dtveloppement, when
it first appeared in 1925 (8). And other great giants of biology
in America in even earlier days-E. B. Wilson, F. R. Lillie, E.

E. FAURfi-FREMIET (1883-1971)

393

leagues such as those whose names are given above, joined by

some protozoological associates, several other distinguished
French scientists, and Monsieur FaurCs successor in the directorship of the laboratory, Etienne Wolff. At this memorable reception, the Patron, with Mme FaurC-Fremiet proudly bv his
side, was presented with a volume of manuscripts of dedicatory
scientific papers which were subsequently published (year 1965)
as 4 fascicles of the well-known French journal PAnnde Biologique.

lmpact on Ciliate Protozoology

Fig. 4. FaurC-Fremiet at the time of his election to the prestigious French Academy in 1959 (he wore his formal attire just
once and without joy).
G. Conklin, T. H. Morgan, H. S. Jennings, C. E. McClung, R.
Chambers, M. F. Guyer, F. M. Child, G. N. Calkins-were apparently similarly impressed by the imaginativeness and acuity
in the prolific output of their modest young colleague on the
other side of the Atlantic.
I have mentioned above that a gauge of this mans impact
on fields of science may be seen in his list of unsolicited honors.
Still another approach to judgment of his influence may be
made by recognition of the accomplishments of students or
close associates of Monsieur FaurC, men and women considering
him their Maitre or even Patron. Such people number
nearly a hundred. A few among the nonprotozoologists of the
group who have gone on to establish outstanding reputations
of their own will serve as examples: Jean AndrC, Charles Devillers, Jean-P. Ebel, Bernard Ephrussi, the late Louis Rapkine,
and Charles Rouiller.
The essentially spontaneous arrangement of a touching 80th
birthday celebration4 in the halls of the College de France, in
February 1964, was carried out by former students and col4Not to mention a similar occasion of honor, held about 10
years earlier, in recognition of his official retirement from the
College de France.

The bulk of FaurC-Fremiets numerous publications is concerned with problems relating to various aspects of the biology
of the ciliate Protozoa, problems on which he brought to bear
the experience and perceptiveness of his earlier training in
microscopy, cytochemistry, and experimental embryology.
Leaving to one side his not inconsiderable contributions to
methods of collecting and culturing ciliates,5 to technics of
fixing and staining (his hundreds of permanent slide preparations will long persist as beautiful proof for posterity of his
skill and precision in such procedures), and to use of microcinematography (in his remarkable studies of feeding methods in
gymnostomes, for example 113a), his works on these protozoa
may be subdivided into at least 5 major areas-still with inevitable degrees of overlapping, of course. They are treated
very briefly below, with citation of representative selected
papers.
T o parallel the mention of his students and disciples in nonprotozoological fields of biology, acknowledgment should be
made here of their protozoological (especially ciliate) counterparts, many of whom have maintained very close ties over
the years with their Patron. A dozen of the students or research collaborators best known in this respect include: Anna
Czapik, Jean Dragesco, Jeanine Ducornet, Marie-Claude Ganier,
Michele Gauchery, Yvette Guilcher-Skreb, Marie Hamard,
Michele Laval, HCltne Mugard-Rouyer (always teasingly called
opaline by Monsieur FaurC), R. H. Pottage, Madeleine
Roque, John B. Tucker, Michel Tuffrau, and the writer.6
Cytology and Ultrastructure
A pioneer in many of his observations, FaurC-Fremiet gained
a lasting reputation among cytologists when he discovered
mitochondria in ciliates in 1907, all the more remarkable in
view of the now tremendously outmoded microscope with which
he carried out such early work. At an even earlier date, 1904,
he discovered and properly interpreted the scopula of peritrichs,
an organelle which continued to fascinate him for the following
67 years. The bulk of his work, throughout his lifetime, involved
making cytological or microscopical observations on natural,
and also experimental, material to understand better the structural diversity so apparent to him in ciliates. Key works on
organelles, exclusive of research on the infraciliature and his
ultrastructural investigations, are: 48-52, 54, 56-59, 84, 88, 89, 95,
101, 125, 129.
He was long ago the inventor, for example, of the protozoological car, the set of submerged slides for collecting species
at various depths in a pond or stream (74) ; and, in more recent
years, he maintained a notable collection of living strains of
diverse species of ciliates in his laboratory at Gif-sur-Yvette ( 146).
Some of his earlier microplankton collecting techniques have also
been described in a publication (see 139).
Actually Messieurs AndrC and Rouiller might well be included
in this as well as the previous list, since their contributions to
protozoan ultrastructure remain formidable even if they might
deny being protozoologists.

394

E. FAURfi-FREMIET ( 1883-1971)

FaurC-Fremiets favorite technic, among many, for study of

cortical characteristics of most species of ciliates, following the
setting of the stage by Bruno Klein in Vienna, Joseph von Gelei
in Budapest, and especially Chatton and Lwoff in France, was
the latters wet method of silver impregnation. Precise revelation of the principal components of both the Silberliniensystem and the infraciliature, viz., the kinetosomes, revolutionized approaches to investigations of many ciliate groups and
thus affected areas of morphogenesis, systematics, and evolution
as well as cytology and comparative morphology. Examples of
contributions in this sub-area, largely exclusive of ultrastructural observations and deliberately omitting here most of the
many morphogenetic and systematic papers, cited subsequently,
in which the French silver impregnation technic played an
indispensable role, are: 47, 48, 94, 100, 103, 107, 160-162,
169.

Over 70 years old when he first took up electron microscopy

in earnest, Monsieur Fauri instantly became a world leader in
ultrastructural biology, once again comprehending the overall
picture as well as producing superb electron micrographs (many
still unpublished) of formerly unknown or poorly understood
cytoplasmic structures. This serves as just one more example
of an enviable trait so characteristic of this astonishing Frenchman: the remarkable ability to adjust to new ideas and technics, even foresee them, and to welcome them with open arms,
with no lingering grief over data earlier gathered with such
devotion and then eclipsed by the newer findings. Representative papers on fine structure of protozoan organelles, fibrillar
structures, etc. are cited in the bibliography (27, 30-32, 36, 37,
44, 45, 48, 121-124, 126, 127, 131, 135-138, 140, 144, 145,
150, 151, 154, 158-161, 164, 165, 167, 168, 170, 171, 173,
175, 176).

FaurC-Fremiets over-riding goal in all of his cytological research was to throw light on the nature of the underlying
organization of the cell or organism concerned. Several examples
of general or review papers of this sort, or of other work not
included in the immediately preceding paragraphs, are given
among the citations at the end of this paper (1, 25, 27, 30-32, 36,
37,44, 48, 107, 109, 113).

Morphogenesis
This was perhaps Monsieur FaurCs favorite area, if he had
any at all, for here he could apply both theory and fact of
the developmental biologist, cellular physiologist, physical chemist, and experimental embryologist to the ever-fascinating problems of the ontogenetic development of a ciliate in its full life
cycle.
Polarity, symmetry, de- and redifferentiation, polymorphism,
auxomorphy, behavior, teratology, cytoarchitectural diversity,
allometric growth, homothety, enantiotropy, inducers, organizers,
regulation, reorganization, regeneration, axial gradients, morphogenetic movements, stomatogenesis, en- and excystation, polystomy, senescence and rejuvenescence, growth and differentiation
of colonies, continuity of subcellular structures, paracrystalline
organization of macromolecules, anisotropy, form-and-function
i la DArcy Thompson (whom he greatly admired)-all
of these notions, concepts, physiological phenomena, and
theories are amenable to straightforward observation or an
experimental approach, both of which Monsieur Faurt brought
to bear on such seemingly simple events as binary fission or
new mouth-formation in a ciliate.
So it was with genuine joy that FaurC-Fremiet tackled such
intriguing challenges, a happiness undiminished, indeed augmented, by the advent of still finer tools of investigation via

the sophisticated technics of molecular biology and electron

microscopy. Some 32 selected samples of his research in fields
of morphogenesis, papers the titles of which clearly delineate
the topics covered in them, are included in the bibliography
(21, 22, 26-28, 35, 41, 45, 47, 48, 53, 55, 73, 75, 90-92, 97,
98, 100, 101, 103, 104, 108, 114, 116, 117, 121, 130, 156,
167, 169).

Ecology
Ordinarily it would be surprising to find a passionate and
productive interest in ecology and natural history field work
demonstrable in a benchman experimentalist, but such was
the case with Monsieur FaurC. He made numerous contributions
to general aspects of protozoan ecology, and took imaginative
approaches to the several specific areas mentioned in the following paragraphs.
FaurC-Fremiet was long interested in planktonic ciliates,
mainly freshwater forms (recall his thesis research, 7) but also
species found in salt-marshes, intertidal pools, sand, and other
unusual habitats (48, 59, 65, 102, 146). Sand-dwelling forms
of the intertidal zone intrigued him from several points of view:
their adaptability to a specialized habitat, the effect of this on
their body form, locomotion, etc., and their possession of a
possibly primitive nuclear apparatus (24, 48, -105, 111, 115,
120. 142. 153).
I

Tidal rhythms among protistan forms also piqued the curiosity

of this insatiable philosopher in little things, and he made
several noteworthy contributions before the days of sophisticated
experimental laboratory approaches to study of such phenomena
(23, 48, 96, 102, 106, 112). He was knowledgeable in the field
of chemical ecology and prepared an important review on
the subject just a few years before his death (42).
Although Monsieur FaurC left to fidouard Chatton most of
the vast area of parasitic ciliate protozoology (see 46), he
retained a lively interest in problems of symbiosis, of adaptability (both of ciliates acting as parasites or commensals or as
hosts themselves to other microorganisms), of facultative parasitism, and of possible preadaptations to a symbiotic way of
life, for example, as seen in histophagous forms (48, 86, 87,
93, 99, 109, 113, 134, 147, 167).

T o me, the work he initiated on community structure

(microbiocenoses) was perhaps the most innovative and heuristic
of all his ecological investigations (23, 48, 110, 152). Unfortunately, few protozoologists or ecologists have had the
patience or ability (or perhaps the insight) to apply his approach to other similar protistan associations in nature; thus
there remains much still to be learned about such interesting
interrelationships.

Phylogeny and Evolution

Since the ciliate fossil record is essentially nonexistent (the
few preserved folliculinid and tintinnid loricae discovered to
date throw no real light on major questions of ancestor-descendant relationships), the area of phylogeny and evolution is
perfect for the inquisitive mind which loves to postulate or
tangle with unprovable hypotheses and theoretical concepts.
Nevertheless, it was with both serious concern and laudable
caution that Monsieur FaurC entered the field, seeking out
strenuously the most reliable criteria on which to base his proposed phylogenetic trees. He was intrigued by the now
generally unacceptable concept of orthogenetic lines and by the
possible clue to phylogenirs to be found in ontogenies. Though
not a teleologist, he was fascinated by pattern and sought evi-

E. FAURB-FREMIET (1883-1971)
dence of affinity through comparisons of the arrangement and
distribution of infraciliary structures.i
FaurC-Fremiet was one of the first to realize the great potential significance of dynamic events in the morphogenesis of
a ciliate with respect to possible phylogenetic interrelationships
of the higher level taxonomic groups to which thr spccics belonged. For nearly 70 years he studied the prritrichs (44, 4852, 66, 84, 87, 97, 107, 126, 131, 137, 140, 161), fascinated
and challengcd to the utmost by their seemingly unique characteristics, but I believe he never was completely satisfied with
explanations offrred by others or by himself with respect to
their most likely affinities Mith other groups of ciliates, or
with how thcy got the way thry arc.
Major contributions to this primarily highly theoretical arcd
include the following: 27, 48, 50, 94, 107, 136, 157, 161, 163,
165.
Systematics
FaurC-Fremiet had a profound appreciation for thr necessity
of schemes of classification and for naming new taxonomic
groups (species and above) within the general guidelines laid
down by the International Code of Zoological Nomenclature.
At the same time, he possessed highly limited patirnce with
strictly armchair nomenclaturists, was not well informed on
details of the Code, and cared little for involved or timeconsuming solutions to complicated taxonomic or nomenclatural problrms. He also had no desire to cope with the
specialized jargon of the several schools of modern evolutionary
biologists, numerical taxonomists, and systematists of the higher
organisms, For rxample, he was disinterested in tcrms or concepts like homoplasy, monophyly, cladistics, patristic distances,
phenetics, phcnograms, a posteriori weighting of characters,
OTUs (operational taxonomic units), positional, compositional,
and serial homologies, genetic distance coefficient, hypodigm,
polytypy, mosaic evolution, hierarchical priority, populational
character divergence, successional species, parataxa, seventyfive per cent rule, symbolic logic and set theory, compatibility
matrices, optimality criteria, and clustering algorithms.
Nevertheless, Monsieur FaurC was constantly obliged to set
up or revise taxonomic schemes and to name new species,
albeit reluctantly, since many of his discoveries inevitably involved ciliates new to science. An uncomplicated phylogenetkist in outlook, he also endeavored to apply his ideas concerning evolutionary relationships within or among groups to
creation of more sensible classificational arrangements. The key
to his philosophy in this area of research was carrying out
detailed comparative morphological studies, supplemented as
much as possible by data from other fields (such as ecology,
physiology, and niorphogenesis) , as a basis for his systematic
conclusions.
Revolutionary though some of his rrsulting proposals may
seem or may have seemed, his viewpoint was nonetheless a rclatively conservative one. He was more a lumper than a splitter
and particularly decried (along with kindred souls such as the
late Libbie Henrietta Hyman) the post-war rash of erection of
new familial taxa. For him the family-group was a rather
sacred unit and could (and should) support considerable diversity among its included genera.
FaurC-Fremiet was appreciativc of the past and very knowledgeable of the great works carried out in the 19th and early

But cytoplasmic or pellicular characters were not used exclusively. FaurC-Fremiet also appreciated the value of nuclear
properties, especially in attempts to understand relationships
among the allegedly most primitive groups of ciliates ( 115,
128, 142).

Fig. 5. An example of FaurC-Fremiets talent as sculptor: an

image of one of his ciliate models, the hymenostome Urocentrum
turbo, complete with inked-in infraciliature. (Photo by Mr. Frederick Dickson, University of Maryland.)
20th centuries. Thus he was not eager to discard lightly the
carefully arrived at taxonomic conclusions of such admirable
prcdrcessors as Stein, Riitschli, Claparsde and Lachmann,
Wallengren, Kent, Penard, and such contemporaries as Chatton, von Gelei, and Kahl.s I have his copy of the latters
prodigious monographs of thc 1930s, bound into one wellworn volume, a surviving testament of Monsieur FarirCs high
regard for the taxonomic work of his German contemporary.
Several taxonomic works are included in the bibliography
(48, 49, 65, 78, 59, 93, 107, 119, 141, 143, 144, 148, 149,
155, 157, 160, 165, 166, 174). Surely the single paper with
the greatest and longest-lasting impact is that rather modest,
succinct work published in 1950, which heralded a new era in
ciliatc systematics (107).

Still Other Talents

While universally rccognizcd professionally as a biologist and
protozoologist of great renown, FaurC-Fremiet also left his mark
in other areas of human endeavor, although this is not so widely
known (primarily bccausc of his modesty).
Apparently inheriting the artistic abilities of his grandfather
Fremiet, Monsieur FaurC produced scvrral sculptures of value,
including busts of Fremiet, Gabriel FaurC, and L.-F. Henneguy.

Chatton passed a\vay in 1947, at the age of 63; von Gelei, in

1952, at 66; Kahl, in 1946, at 69.

E. FAURE-FREMIET (1883-1971)
Fellow protozoologists have long envied the drawings in
FaurC-Fremiets scientific papers. His graceful, life-like sketches
of protozoan species (many unpublished) are, again, reminiscent
of animal drawings made by the talented E. Fremiet. An
outstanding example are the illustrations in his published
doctoral dissertation (7; and see Fig. 6 on this page).
Monsieur F a d s ability with prose, plus his conscientious
appreciation of the historical perspective, and his own longevity,
earned him the rather onerous task of authoring in memoriam
notices for a considerable number of colleagues in biology whom
he outlived (11, 13, 29, 34, 38, 38a, 39). He also was the
author of many papers on other assorted historical topics (3,
10, 14, 1.5, 33, 40, 46).
Often I have been asked if FaurC-Fremiet inherited any of
the musical talent of his father, Gabriel FaurC. Apparently not,
in the sense of being a composer or a pianist (brother Philippe
did play quite well). But his sensitivity and compassion for
life, his appreciation and feeling for the arts and humanities,
and his profound respect, admiration, and love for his father
were striking characteristics of Monsieur FaurC. We have lasting
evidences of this, most recently in a posthumous publication in
Scherzo (the December 197 1 number, dedicated to Saint-SaEns),
reporting on an interview with FaurC-Fremiet concerning
memories of his father; and in a beautiful radio broadcast made
in 1944, the 20th anniversary of Gabriels death, by his elder
son. A quotation9 from the introductory words of that address
follow :

Fig. 6. An example of FaurC-Fremiets artistic talent: a copy

of his original drawing, made in ca. 1913 (see references 7 and
6 5 ) , of the celebrated Tontonia appendiculariformis, a planktonic
oligotrich ciliate.
The originals of these fine statues are located in appropriate
institutions or parks in Paris or other cities. The one of his
father is the central figure in the beautiful park at Foix, the
picturesque small village in Arihge (southern France) near
where Gabriel FaurC grew up, now the site of an annual music
festival. He also produced some smaller objects, such as a very
life-like and inquisitive mouse which long adorned a bit of a
shelf in the living room of his apartment. And at the laboratory
he made a number of splendid models of ciliates (e.g., see
Fig. 5 ) or of their organelles (e.g., the complex denticulate
ring of the basal disc of trichodinids).
In his younger days, FaurC-Fremiet did considerable painting
(while concomitantly managing to put in many hours of research
at the laboratory). His excellent works, generally in oils, include
an often exhibited detailed full-length portrait of grandfather
Fremiet in dress regalia, a painting willed to the Museum at
Dijon; a touching head-and-shoulders of his wife Jeanne; one
of his sister-in-law Suzanne and one of his mother-in-law;
another of his father-in-law, showing Henneguy sitting in his
garden at PrCcy in front of another canvas himself; one of
Jeanne and Suzanne together, standing by the ancient well at
Prtcy; and a delightful one-surely one of his earliest-of little
brother Philippe earnestly practicing at the piano.

In memory of my father, permit me to evoke some recollections of my childhood : the incomparable privilege of having
been rocked to sleep near him by nocturnes, barcaroles, and
newly born melodies; then the emotion I felt while timidly
listening to a rehearsal of the Sonata in A, the quartets, or the
first quintet; finally, the confused revelation of an inexpressible
world, which I believe now, having become a man of the
laboratory, surpasses in richness all human knowledge, when
the music, this music, uncovers deep within us a mysterious
realm, mysterious because of its impossible verbal transcription.
In a much lighter vein, I must relate the following tale which
indicates that being the son of a great celebrity has its risks as
well as pleasures. Monsieur FaurC himself loved to tell this
story, and with a little more elaboration than space here permits.
Some 10 or 15 years ago, when Faurt-Fremiet was already
about 75 years old, he was journeying to Warsaw (or was it
Vienna?) to give a major address at a scientific conference
and was, rare for him, travelling from Paris via an express
wagon-couchette, rather than his usual frugal second class
sitting-up-all-night mode of rail travel. Just as, with a sigh of
contentment, he was preparing to enter his (lower) berth, an
ample-bosomed lady bore down on him in the aisle, with an
enraptured look on her face. Ah, Monsieur, cried she, 5s it
true that you are indeed the son of Gabriel FaurC, that superbly
wonderful man whose music is so heavenly [etc., etc., quite
literally into the night!]? Barely allowing poor Monsieur FaurC
to admit to that filial status, she continued on and on, with her
captive audience desperately wondering if hed ever get a chance
to sleep at all before the very active day ahead of him at the
next stop. Finally, a couple of hours later, emotionally
exhausted, the lady returned her attention more specifically to
her polite listener, staunchly standing by, Oh, just imagine my
being with a living son of that great master of music!! Please,
kind sir, embrace me . . . but chastely!
Typescript of the full address generously supplied by Mme
Marie FaurC-Fremiet; translation of the paragraph quoted here
kindly made by Mrs. Barbara Whitney.

E. FAURfi-FREMIET (1883-1971)
Some Personal Characteristics
A number of the personality traits of Monsieur Faur6s have
been revealed, here and there, in preceding sections of this paper.
But a few additional comments must be made. Great and often
abstract scientist though he was, the man had an abiding interest
in and generous love for young people and animals-all
the
more pity that he had neither children nor pets. He had a
magic way of making the other person, no matter what his or
her generation, feel important, at least partly through his way
of giving undivided attention to the words of his companion
of the moment.
There was never a more gracious host to visiting firemen
coming through Paris. During my year there, I was introduced
to more distinguished Americans (not to mention other nationalities) in his home than Id met in four years of graduate
school in New York City. After Monsieur FaurC and poor
Mme Jeanne and Mlle Suzanne had wined, dined, and entertained the guests royally in his little apartment, he frequently
would, regardless of the hour, return to his desk to work into
the night on some manuscript or other.
At the Coll?ge de France he always had time for the questions
of students and colleagues, though, at the same time, he was
impatient with inefficiency, irresponsibility, and out-and-out
laziness. He practically adopted some of his students, especially
intellectually. An outstanding example was his deep feeling for
young Jean Filhol, a brilliant embryologist crippled in body but
certainly not in mind. When Filhol passed away, at the tender
age of 28, Monsieur FaurC delivered a moving eulogy (16).
He seemed to have an especially warm spot in his heart for
Americans and Americana, from cokes and sliced bread to
plaid shirts and saddle shoes. And he loved certain expressions,
sometimes rather quaint, in English, for example, Oh my
goodness! On his infrequent trips (about every 20 years!) to
the United States, he always managed to find some souvenir,
often an article of clothing, to take back home with him as a
long-lasting addition to his meager wardrobe. Perhaps the only
object more characteristic of him than some conspicuous article
of American clothing was his beloved pipe, a constant companion (see Figs. 1, 2 ) until the last few years of his life.
Modest and frugal (for example, he seldom ordered more
than 100 reprints of any of his papers), sensitive, gentle,
generous, creative, Monsieur Faurt was also tremendously selfdisciplined and held to the highest standards of quality in his
work. I n fact, sometimes his passion for perfectionism worked
to the distinct disadvantage of posterity-two examples of this
will suffice.
The story is told that, in 1929, the late Maurice Caullery
(whose place F a d - F r e m i e t took in the Institut, incidentally)
sent off to press the manuscript of a book on cytology prepared
by his young colleague while Faur6-Fremiet himself was abroad
for several months. When the younger man returned, he was
greatly upset and immediately withdrew the work from the
publisher, with the typical comments that, first, he must add
a little new information to it; and second, the entire draft
would have to be put into better shape. The book never did
appear.
Then, in the 1940s, when Pierre-P. Grass6 was lining up
contributors for his Traite de Zoologie, keen on being the first
European zoologist to ever start and finish such a series, FaurtFremiet was asked to prepare Tome 11, on the cell and the
ciliate Protozoa, with a deadline set for early in 1949. Alas,
by the time of his passing, in 1971, Monsieur Faurt was still
not satisfied with his draft of the text (it occupied several file
drawers, thus still requiring much heart-rending condensation),

397

having spent the 22 years since the first deadline concentrating

on electron microscopy of the ciliates, firm in his conviction
that the work would have been immediately out of date had it
not contained comparative ultrastructural data involving a t least
a representative number of the more than 5,000 known species
of thcsc protozoa.10
There is ample evidence, finally of a keen sense of humor
and a quick wit in this man to remember. This is attested by
many a witness, young and old, French and American, but space
does not permit recalling specific anecdotes here. There must
have been many a time when his physical condition (for
example, when he had to undergo the painful and rather
long-drawn-out operation on his face about 10 years before his
death) or his mental concentration on a particularly demanding
task would have caused most men to become grim or shorttempered or at least preoccupied. But Monsieur FaurC seemed
always, if the occasion were appropriate, to have a smile or a
joke or even a practical prank on his faux or up his sleeve, to
the delight of all, surely including himself!
This fun-loving spirit must have been a factor in keeping the
man full of life and energy and keen in perceptiveness through
nearly fourscore years and eight. I t is a nice way to remember
him, as a person. As a scientist, he will livr on via his published
works and through the many younger biologists inspired by the
high example set by their Patron.
SELECTED BIBLIOGRAPHY
Explanation
It is customary to end such an in memoriam tribute as this
with a listing of all the works published by the man being
honored. But Professor FaurC-Fremiet contributed so many papers
(nearly 500) to the progress of protozoology (and developmental
biology) over a span of nearly three-quarters of a century that
it would not be feasible to follow that custom here.
Therefore, I am offering a selected bibliography in 2 parts,
each with its included works arranged strictly chronologically.
Part Z lists books or monographs, major reviews, important general
articles of an historical nature, selected in memoriam notices,
and review chapters in books edited by others. This is followed
by Part 11, a longer section, which is comprised of references to
FaurC-Fremiets major biological (largely protozoological) papers
considered to be of special significance. Neither part is intended
to be exhaustive in its coverage; in Part ZZ, emphasis has been
given to papers published during the last 25 years of his career;
but references in both parts cover a period of more than 65 years.
Although this section is entitled A Selected Bibliography, it
is also Literature Cited, for no work is included to which a
specific reference does not exist in the text of this paper. Thus
I have been spared the nearly impossible task of grouping papers
here by the appropriate field or category: for, by the numbers
cited in the text, the reader can gain an appreciation of the contribution to some specific area of biological research; and yet the
same work can be cited more than once, if it is related to more
than a single field or area (as many of his papers are).
Only about a third of FaurC-Fremiets own works (alone or with
collaborators) are included here; and totally excluded are some
250 other papers considered as travaux du Laboratoire. The
latter are contributions by students and associates working directly
or indirectly under his general supervision, publications to many
of which Monsieur Faurt, if he had been a man of less magnanimity, might quite justifiably have added his own name as
coauthor.
A final note of explanation is needed concerning the references
presented below. The citations comprising this section are presented in a style which is, in part, not typical for this Journal.
10 A group of younger men, including the writer, will attempt
to finish the preparation of Tome 11, preserving Monsieur FaurCs
ideas and, as frequently as possible, his very words (and, of
course, his illustrations), so that the work (48) will stand as an
enduring tribute to him, the kind of memorial he would most
cherish.

398

E. FAURR-FREMIET (1883-1971)

The numbers are given in parentheses, followed by the date of

the work cited, then its title, etc., with an occasional comment
(and names of co-authors) placed in parentheses at the end of
the citation.

(33) 1960. Notice sur la vie et les travaux de Victor Coste

1807-1873. Institut France, Acad. Sci., Paris 13, 1-47.
(34) 1960. Notice sur la vie et les travaux de Maurice Caullery
(1868-1958). Institut France. Acad. Sci.. Paris 14, 1-52.

Part I Books, Monographs, Review and Historical Articles

( 1 ) 1907. Les organismes monocellulaires. Rev. Idies, March
1907, pp. 204-221.
( 2 ) 1908. La structure des matikres vivantes. Bull. Soc. 2001.
France 33, 104.
( 3 ) 1908. La protistologie. Histoire de quelques de ses rCcents progres. Rev. Ghn. Sci. 19, 313-21.
( 4 ) 1911. La dCcouverte des mitochondries et les idCes biologiques. Rev. Zdees, No. 92, 125-36.
( 5 ) 1919. Revue de nos connaissances sur les lois mathCmatiques de la cicatrisation des plaies. Ann. Biol. 24, 5-26 (with
F. Vles).
( 6 ) 1922. Rapport sur letat actuel des publications bibliographiques pour les sciences biologiques et mCdicales. Deuxisme
rCunion du Conseil International de Recherches A Bruxelles (1922).
Union des Sciences Biologiques. (Actually not published until
1925, in the IUBS Proceedings of that year.)
( 7 ) 1924. Contribution A la connaissance des infusoires planktoniques. Bull. Biol. France Belg. Suppl. 6, pp. 1-171. (Doctoral
Dissertation. )
( 8 ) 1925. L a Cinetique du Develo$pement. Presses Universitaire France, Paris. 335 pp.
( 9 ) 1926. Oi en est lEmbryologie? Gauthier-Villars et Cie.
249 pp.
(10) 1929. La Chaire dEmbryogCnie ComparCe du CollBge
de France et ldvolution des sciences du dCveloppement. Rev. Gkn.
Sci. 41), 37-45.
1 1 1 ) 1929. Louis-FClix Henneauv
Arch. Anat.
- , (1850-1928).
.
Micro;. 25, 1-28.
( 12) 1930. The kinetics of living matter. Trans. Faraday SOC.
26, 779-93.
(13) 1931. Albert Brachet. Presse Med., No. 12, 1-11.
(14) 1932. La crCation de la Chaire dEmbryogCnie ComparCe au College de France (Extrait du Livre jubilaire compose
d Poccasion d u quatrieme centenaire d u Coll2ge de France).
Presses Universitaire France, Paris. pp. 193-207.
( 1 5 ) 1935. Les recherches de Felix Dujardin et la notion de
protoplasma. Protoplasma 23, 250-69.
(16) 1938. Jean Filhol (1910-1938). Masson et Cie, Paris,
1 1 PP.
( 1 7 ) 1944. Le problkme de lorganisation et ses aspects physicochimiques. Rev. Sci. 81, 433-46.
(18) 1945. La mati& au point de vue biologique. Centre
International d e Synth2se (May 1939). Presses Universitaire
France, Paris. pp. 115-30.
(19) 1946. La cellule et son organisation. Revue des travaux
recents de cytologie (1940-1946). Ann. Biol. 22, 57-1 18.
( 2 0 ) 1946. Application du microscope Clectronique A 1Ctude
des problemes de la biologie cellulaire et bacthrienne. Rev.
Optique, year 1945, pp. 55-84.
( 2 1 1948. Les mkcanismes de la morohoaenhse
chez les cili6s.
. Fok-Biotheor. 3,-25-58.
( 2 2 ) 1949. Problhmes molCculaires de la morphogenbse. Ann.
Biol. 26, 361-79.
(23) 1950. Ecology of ciliate infusoria. Endeavour 9, 183-

(37) 1961. Cils vibratiles et flagelles. Biol. Rev. 36, 464-536.

(38) 1962. Notice nbcrologique sur Pol Bouin (1870-1962).
C. R . Acad. Sci.. Paris 254. 1361-4.
(38a) 1962. Notice nCcrologique sur Christian Champy (18851962). C . R . Acad. Sci., Paris 254, 3773-5.
(391 1964. Notice nCcrologiaue sur Louis Faae ( 1883-1964).
C . R . Acad. Sci., Paris 258, 6632-5.
(40) 1966. Les origines de 1AcadCmie des Sciences de Paris.
Notes Records Roy. SOC.London 21, 20-31.
141) 1967. La rCeCnkration chez les Drotozoaires. Bull. SOC.
Zool. France 92, 249--72.
(42) 1967. Chemical aspects of ecology, in Kidder, G. W., ed.,
Protozoa. in Florkin. M. & Scheer. B. T.. eds.. Chemical Zooloev.
Academic Press, New York 1,21-54.
(43) 1967. Les prCmices de la biologie molCculaire, in Institut
de France, AcadCmie des Sciences, Troisikme Centenaire, 16661966, 11, pp. 266-91.
(44) 1969. Organites et appareils de fixation chez les Ciliata.
3rd Int. Cong. Protozool., Leningrad, July 1969, Progress in
Protozoology, pp. 7-8.
(45) 1970. Microtubules et mCcanismes morphopoietiques. Ann.
Biol. 9, 1-61.
(46) 1971. Aux croisements de nos chemins, in Monod, J. &
Borek, E., eds., Of Microbes and Life, Columbia University Press,
New York. pp. 36-49.
(47) MS. La differenciation des structures buccales au cours
de 1Cvolution des Ciliata (Unpublished).
(48) 1973. Tome I1 of Trait6 de Zoologie, edited by P.-P.
GrassC, Paris. In press.
I

-.
I

Part I1 Selected Research Papers (mainly protozoological)

(49) 1904. Note sur un groupe nouveau dopercularia. Arch.

Anat. Micros. 7, 181-97. (First major publication.)
(50) 1904. Appareil fkateur des discotriches et ses indications
au point de vue de la phylogenkse. C . R. Soc. Biol. 57, 464-6.
(51) 1904. Sur la structure du pkdoncule chez les Vorticellidae. C . R . Soc. Biol., Paris 57, 506-8.
(52) 1905. La structure de lappareil fixateur chez les Vorticellidae. Arch. Protistenk. 6, 207-26.
(53) 1906. Sur un cas de monstruositC chez Stentor coeruleus.
Arch. Anat. Micros. 8. 660-6.
(54) 1907. Mitochondries et sphCroplastes chez les infusoire;
ciliCs. C . R . SOC. Biol., Paris 62, 523-5. (Discovery of mitochondria in protozoa.)
(55) 1907. Organisation, fonctionnement et rCactions individuelles chez les cytozoaires. Bull. Znst. G i n . Psychol., year 1907,
pp. 75-6, 305-34.
(56) 1910. Appareil nucleaire, chromidies, mitochondries. Arch.
Protistenk. 21, 186-208.
(57) 1910. etude sur les mitochondries des protozoaires et des
cellules sexuelles. Arch. Anat. Micros. 10, 457-648.
(58) 1910. La fixation chez les infusoires cili6s. Bull. Sci.
7.
France Belg. 44,27-50.
(24) 1951. Rcologie des protistes littoraux. Ann. Biol. 27,
(59) 1912. etudes cytologiques sur quelques infusoires des
437-47.
marais salants du Croisic. Arch. Anat. Micros. 13, 401-79.
( 2 5 ) 1952. La diversification structurale des ciliCs. Bull. SOC.
(60) 1912. Points controversCs de la spermatogenhe de IAscaris
Zool. France 77, 274-81.
megalocephala. C . R . SOC.Biol., Paris 73, 271-3.
(26) 1953. Lhypothgse de la senescence et les cycles de re(61) 1912. Un appareil de Golgi dans Ioeuf de Idscaris
organisation nuclCaire chez les ciliCs. Rev. Suisse 2001.60, 426- megalocephala. Bull. Soc. 2001.France 27, 135-8.
38.
(62) 1912. Etude des foraminifBres de la mission fransaise
(27) 1953. Morphology of protozoa. Ann. Rev. Microbiol. 7,
antarctique. Bull. Mus. Paris, year 1911, pp. 76-9.
1-18.
(63) 1913. Les foraminiferes de la seconde mission antarctique.
(28) 1954. Les problPmes de la diffkrenciation chez les proDeuxieme campagne du Pourquoi-Pas? Bull. Soc, Zool. France
tistes. Bull. SOC.Zool. France 79, 31 1-29.
(29) 1954. Notice nCcrologique (RenC Legendre). Bull. SOC. 38, 260-71.
(64) 1913. Le cycle germinatif chez IAscaris megalocephala.
Zool. France 79, 70-3.
( 3 0 ) 1956. Finer morphology of microorganismes. Ann. Rev. Arch. Anat. Micros. 15, 435-757.
(65 j 1914. Deux infusoires planktoniques: Tontonia appendiMicrobiol. 11, 1-6.
(31) 1958. Organismes, cellules, molecules. Le cas des in- culariformis (nov. gen. nov. sp.) et Climacostomum diedrum
(nov. sp.). Arch. Protistenk. 34, 95-107.
fusoires ciliks. Biol. Jaarb. 24, 47-55.
( 3 2 ) 1958. Structure et ultrastructure des protistes. Rev. Path.
(66) 1922. Le cycle de croissance des colonies de vorticellides.
Bull, Biol. France Belg. 56, 427-53.
G i n . Physiol. Clin. 58, 265-79.
~~

E. FAURfS-FREMIET (1883-1971)
(67 ) 1922. Mkthodes chimiques et microchimiques applicables

B 1Ctude cytologique des oeufs. C . R . Assoc. Anat., year 1922, p.

149.
(68) 1923. Le premier cycle de croissance du tetard de Rana
temporaria. Arch. Znt. Physiol. 21, 403-37 (with J. Dragoiu).
(69) 1924. Loeuf de Sabellaria alueolata. Arch. Anat. Micros.
20, 211-342.
(70) 1927. Les amibocytes des invertCbrCs B 1Ctat quiescent et
B ICtat actif. Arch. Anat. Micros. 23, 99-173.
(71) 1928. La culture des tissus in vitro. Ann. Anat. Path.
Anat. Norm. 5, 157-80 (with B. Ephrussi).
( 72) 1929. Caract6res physico-chimiques des choanoleucocytes
de quelques invertCbrCs. Protoplasma 6, 52 1-609.
(73) 1930. Growth and differentiation of the colonies of
Zoothamnium alterans (Clap. and Lachm.). Biol. Bull. 58, 28-51.
(74) 1931. Quelques rbsultats obtenus avec la mCthode des
lames immergbes. Bull. Soc. Zool. France 56, 479-82.
(75) 1932. Division et morphogensse chez Folliculina ampulla
0. F. Miiller. Bull. Biol. France Belg. 66, 77-1 10.
(76) 1932. Morphogenhe exPCrimentale (reconstitution) chez
Ficulina ficus. Arch. Anat. Micros. 28, 1-80.
( 7 7 ) 1932. Involution exPCrimentale et tension de structure
dans les cellules de Ficulina ficus. Arch. Anat. Micros. 28, 121-57.
(78) 1935. La famille des Philasteridae Kahl (Ciliata Holotricha). Bull. Soc. Zool. France 40, 127-43.
(79) 1936. La famille des Folliculinidae (Infusoria Heterotricha). M i , . Mus. Roy. Hist. N u t . Belg., SCr. 2, 3, 1129-75.
(80) 1936. La structure des fibres delastoidine. Arch. Anat.
Micros. 32, 249-70.
(81) 1938. Structure de la capsule ovulaire chez quelques
sClaciens. Arch. Anat. Micros. 34, 23-51.
(82) 1938. Etude roentgCnographique de quelques collaghes.
J . Chim. Phys. 35, 223-32 (with G. Champetier).
(83) 1938. Etude roentgenographique des kCratines secrCtCes.
C . R . Acad. Sci., Paris 207, 1133-5 (with G. Champetier).
(84) 1941. La nature chimique du pkdoncule des vorticellides.
Bull. Soc. Zool. France 66, 277-87.
(85) 1942. Composition chimique du tegument chez Coleps
hirtus. Bull. Biol. France Belg. 78, 136-42 (with M. Hamard).
(86) 1943. Commensalisme et adaptation chez un acinCtien:
Erastophrya chattoni, n. gen., n. sp. Bull. SOL. Zool. France 68,
145-7.
(87) 1943. Commensalisme et adaptation chez une vorticellide:
Epistylis lwoffi, n. sp. Bull. SOL.Zool. France 68, 154-7.
(88) 1943. Etude biometrique de quelques trichodines. Bull.
SOC.Zool. France 68, 158-69.
(89) 1944. ProtCines de structure et cytosquelette chez les
urcColaires. Bull. Biol. France Belg. 78, 143-56 (with J. Thaureaux) .
(90) 1944. Polymorphisme de 1Enchelys mutans (Mermod).
Bull. Soc. Zool. France 69, 212-9.
(91) 1945. SymCtrie et polarit6 chez les ciliCs bi- ou multicomposites. Bull. Biol. France Belg. 79, 106-50.
(92) 1945. Polymorphisme du Monodinium uorax nov. sp.
Bull. SOC.Zool. France 70, 69-79.
(93) 1946. Sur un infusoire holotriche histiophage Deltopylum
rhabdoides n. gen., n. sp. Bull. SOC.Zool. France 71, 161-4 (with
H. Mugard).
(94) 1947. Les affinitCs des infusoires acinCtiens et la ciliature de leurs formes vagiles. Bull. SOC. Zool. France 72, 12-6
(with Y . Guilcher).
(95) 1947. Trochilioides filans n. sp. infusoire holotriche de
la famille des Dysteriidae. Bull. SOC. Zool. France 72, 106-12
(with Y. Guilcher).
(96) 1948. Le rythme de marbe du Strombidium oculatum
Gruber. Bull. Biol. France Belg. 82, 3-23.
(97) 1948. Croissance et morphogenhse des colonies de Carchesium limneticum Svec. Ann. Acad. Bras. Cien. 20, 103-15.
(98) 1948. DCveloppement allomktrique des constituants du
pCdoncule chez Epistyle capitulum n. sp. et Opercularia cupulata
n. sp. Ann. Acad. Bras. Cien. 20, 95-102.
(99) 1948. Un cas dassociation entre ciliC et tentaculifhre
Cpizoiques. Ann. Acad. Bras. Cien. 20, 117-23.
( 100) 1948. Doublets homopolaires et rCgulation morphoqCnCtique chez le ciliC Leucophrys patula. Arch. Anat. Micros.
37, 183-203.
(101 ) 1948. ktude expCrimentale de la calcification t6gumentaire chez un infusoire riliC, Coleps hirtus. Biochim. Biophys. Acta
2, 668-73 (with J. Stolkowski & J. Ducornet).

399

(102) 1949. The ecology of some infusorian communities of

intertidal pools. J . Anim. Ecol. 17, 127-30.
(103) 1949. Le dimorphisme de Espejoia mucicola Penard.
Hydrobiologia 1, 379-93 (with H. Mugard).
(104) 1949. Action du lithium sur la stomatogen&se chez les
ciliCs. Be1g.-Nederl. Cyto-embryol. Dagen., Gent, pp. 100-2.
( 105) 1950. Rcologie des ciliks psammophiles littoraux. Bull.
Biol. France Belg. 84, 35-75.
( 106) 1950. Rythme de m a r k dune Chromulina psammophile.
Bull. Biol. France Belg. 84, 207-14.
(107) 1950. Morphologie comparCe et systkmatique des ciliCs.
Bull. SOC.Zool. France 75, 109-22. (Now classical paper revolutionizing the approach to overall systematics of ciliate Protozoa.)
( 108) 1950. MCcanismes de la morphogenhse chez quelques
ciliks gymnostomes hypostomiens. Arch. Anat. Micros. 39, 1-14.
( 109) 1950. CaulobactCries Cplzoiques associCes aux Centrophorella (ciliCs holotriches). Bull. SOC.Zoo!. France 75, 134-7.
( 1 10) 1951. Associations infusorienne a Beggiatoa. Hydrobiologia 3, 65-71.
( 111) 1951. The marine sand-dwelling ciliates of Cape Cod.
Biol. Bull. 100, 59-70.
( 1 1 2 ) 1951. The tidal rhythm of the diatom Hantzschia
amphioxys. Biol. Bull. 100, 173-7.
(113) 1951. Symbiontes bactCriens des ciliCs du genre Euplotes.
C . R . Acad. Sci., Paris 135, 402-3.
(113a) 1953. Feeding in vegetative and carnivorous gymnostome ciliates (Motion picture). Proc. SOC. Protozool., 4, 4 (with
technical assistance of J. Dragesco and commentary by J. 0.
Corliss).
( 114) 1953. La bipartition Cnantiotrope chez les ciliCs oligotriches. Arch. Anat. Micros. 42, 209-25.
( 115 ) 1954. Reorganisation du type endomixique chez les
Loxodidae ciliks et chez les Centrophorella. J . Protozool. 1, 20-7.
( 116) 1954. Morphogen&se de bipartition chez Urocentrunz
turbo (ciliC holotriche). 1. Embryol. Exp. M o r p h . 2, 227-38.
( 11 7 ) 1954. Les processus de Ienkystement chez Euplotes
muscicola Kahl. Bull. Biol. Zool. France 88, 154-67 (with M.
Gauchery & M. Tuffrau) .
(118) 1954. Les inclusions protkiques de Ioocyte de Parascaris equorum. Exp. Cell Res. 7, 153-68 (with J.-P. Ebel & J.
Colas).
( 119) 1954. La position systCmatique du genre Balantidium.
J . Protozoal. 2 , 54-8.
(120) 1955. Sonderia labiata, n. sp. ciliC trichostome psammobie. Hydrobiologia 7,2 10-8 (with M. Tuffrau).
( 121) 1956. La dorganisation macronuclCaire chez les Euplotes. Exp. Cell Res. 12, 145-50.
(122) 1956. Structure et origine du pCdoncule chez Chilodochona. J . Protozool. 3, 188-93 (with C. Rouiller & M. Gauchery). (One of FaurC-Fremiets earliest full papers using electron
microscopy.)
( 123) 1956. Lappareil squelettique et myoide des urcColaires;
Ctude au microscope Clectronique. Bull. Soc. Zool. France 81,
77-85 (with C. Rouiller & M. Gauchery).
(124) 1956. Les tentacules dEphelota. Etude au microscope
Clectronique. 1. Protozool. 3, 194-200 (with Ch. Rouiller & M.
Gauchery) .
(125) 1956. Sur les pigments du cilik Stentor niger. C . R .
Acad. Sci., Paris 62, 2182-4 (with M. Barbier & E. Lederer).
( 126) 1956. Origine ciliaire des fibrilles sclCroprotCiques pCdonculaires chez les ciliCs pCritriches. etude au microscope Clectronique. Exp. Cell Res. 11, 527-41.
( 1 2 7 ) 1956. Les structures myoides chez les ciliCs; dtude au
microscope Clectronique. Arch. Anat. Micros. 43, 139-61 (with
C. Rouiller & M. Gauchery).
(128) 1957. Le macronuclCus hCtCromhre de quelques ciliCs.
J . Protozoal. 4 , 7-17.
( 129) 1957. ConcrCtions minCrales intracytoplasmiques chez les
ciliCs. J . Protorool. 4, 96-109 (with M. Gauchery).
(130) 1957. Trichopus lachmanni n. sp.; structure et morphogenhse. J . Protozool. 4, 145-50.
(131 ) 1957. Ultrastructure rCticulCe dune fibre squelettique
chez un ciliC. J . Ultrastruct. Res. 1, 1-13.
( 132) 1957. Structure fine dun flagell6 chrysomonadien:
Chromulina psammobia. E x p . Cell Res. 14, 47-67 (with C.
Rouiller) .
(133) 1957. Etude au microscope Clectronique dune bactCrie
sulfureuse Thiouulum majus Hinze. Exp. Cell Res. 14, 29-46
(with C. Rouiller).

400

E. FAURE-FREMIET (1883-1971)

(134) 1958. Le ciliC Condylostoma tenuis n. sp. et son algue

symbiote. Hybrobiologia 10, 43-8.
( 135) 1958. Lultrastructure des trichocystes fusiformes chez
Frontonia atra. Bull. Micros. Appl. 7, 135-9 (with C. Rouiller).
(136) 1958. The origin of the Metazoa and the stigma of the
phytoflagellates. Quart. J . Micros. Sci. 99, 123-9.
(137) 1958. Ultrastructure des cinCtosomes B ICtat de repos
et B ICtat cilifkre chez un ciliC pbritriche. J . Ultrastruct. Res. 1,
289-94 (with C. Rouiller).
(138) 1958. Myonkmes et cinktodesmes chez les ciliCs du genre
Stentor. Bull. Micros. Appl. 8, 117-9 (with C. Rouiller).
(139) 1958. MatCriel pour la rCcolte du microplankton dans
les mares et Ctangs. Hydrobiologia 12, 142-8.
(140) 1959. Le cortex de la vacuole contractile et son ultrastructure chez les ciliCs. J . Protozool. 6, 29-37 (with C. Rouiller).
(141) 1959. La famille des Nassulidae (Ciliata Gymnostomatida) et le genre Nassulopsis n. gen. C. R. Acad. Sci., Paris 249,
1429-33.
( 142) 1961. Quelques consid6rations sur les ciliCs mCsopsammiques B propos dun rCcent travail de J. Dragesco. Cah. Biol.
Mar. 2, 177-86.
( 143) 1961. Remarques sur la morphologie comparCe et la
systhatique des Ciliata Hypotrichida. C. R. Acad. Sci., Paris
252, 3515-9.
(144) 1961. Les ciliCs Cyrtophorina et leur diversification
morphologique. C. R. Acad. Sci., Paris 252, 3912-6.
( 145) 1961. Le cytoplasrne stomo-pharyngien des ciliCs cyrtophores. C. R. Acad. Sci., Paris 253, 357-62.
(146) 1 9 6 1 . Documents et observations Ccologiques et pratiques sur la culture des infusoires cilibs. Hydrobiologia 18, 300-20.
( 147) 196 1. Pleurocoptes hydractiniae (Ciliata Pleuronematina) et son adaptation structurale B la vie epizoique. Cah. Biol.
Mar. 2. 447-52.
(148) 1962. Le genre Paranassula Kahl (Ciliata Cyrtophorina).
Cah. Biol. Mar. 3, 61-77.
(149) 1962. Le aenre Trimvema Lackey et les caracteristiaues
structurales des cili& Trichostomatida. J. Protorool. 9, 146-9. *
(150) 1962. etude au microscope Clectronique des ultrastructures dEpistylis anastatica (cili6 pCritriche) J. Micros. 1, 2873 12 (with P. Favard & N. Carasso).
(151 ) 1962. Ultrastructure de Iappareil excrCteur chez quelques
ciliCs pCritriches. J . Micros. 1, 455-68 (with N. Carasso and P.
Favard) .
( 152) 1963. Images Clectroniques dune microbiocenose marine.
Cah. Biol. Mar. 4, 61-4 (with N. Carasso & P. Favard).
(153) 1963. Conchostoma longissimum n. g., n. sp., ciliC
trichostome psammobie. Cah. Biol. Mar. 4, 193-9.
(154) 1963. Ultrastructure de Iappareil adhCsif des urcColaires
(ciliks pCritriches). J . Micros. 2, 337-68 (with P. Favard and N.
Carasso).
(155) 1963. Remarques sur la morphologie comparCe des
cilibs Trichostomatida. C . R. Acad. Sci., Paris 257, 3089-94.

BOOK R V / W . .

(156) 1964. Les ciliCs hypotriches rCtrocursifs. Arch. 2001.

Exp. Gkn. 104, 65-74.
(157) 1965. Plesiotrichopus natans n. g., n. sp. (Ciliata Cyrtophorina). Arch. Zool. Exp. Ckn. 105, 355-60.
( 158) 1965. Lorganisation du ciliC gymnostome Plagiocampa
ovata Gelei. Arch. 2001.Exp. G6n. 105, 361-8 (with J. Andrb).
(159) 1965. Etude au microscope Clectronique de Tillina
praestans Penard (ciliC Trichostomatida). Arch. 2001.Exp. Gkn.
105, 345-53 (with J. AndrC).
(160) 1965. Morphologie des Dysteriidae (Ciliata Cyrtophorina). C. R. Acad. Sci., Paris 260, 6679-84.
(161) 1965. Morphologie cornparbe des cili6s Peritrichida. Excerpta Medica, Int. Cong. Ser. No. 91, 2nd Int. Conf. Protozool.,
London, July-Aug. 1965, Progress in Protozoology, pp. 13-4.
(162) 1966. Polycycla (Urceolaria) synaptae CuCnot. Cah.
Biol. Mar. 7, 61-8 (with A. Czapik).
(163) 1967. La frange ciliaire des Nassulidae (Ciliata Cyrtophorina) et ses possibilitks Cvolutives. C. R. Acad. Sci., Paris 264,
68-72.
(164) 1967. Formation et structure des concrCtions calcaires
chez Prorodon morgani Kahl. 1.Micros. 6, 391-8 (with J. AndrC).
(165) 1967. etude au microscope Clectronique du ciliC Pseudomicrothorax dubius Maupas. J . Protozool. 14, 464-73 (with J.
And&).
(166) 1967. Le genre Cyclogramma Perty 1852. 1. Protozool.
14, 456-64.
(167) 1967. LCsions cytoplasmiques provoqubes chez un ciliC
par un tentaculifsre parasite. Protistologica 3, 121-6 (with J.
AndrC) .
( 168) 1967. Technique de rassemblement de rnicroorganismes;
prCinclusion dans un caillot de fibrine. J . Micros. 6, 1063-6 (with
R. Charret).
(169) 1968. Les genres Dexiotricha Stokes et Loxocephalus
Eberhard dans leur relations auxomorphiques. Protistologica 4,
115-25.
(170) 1968. Structure fine de 1Euplotes eurystomus (Wrz.).
Arch. Anat. Micros. 57, 53-78 (with J. AndrC).
( 1 7 1 ) 1968. Structure fine de Iorganite f i a t e u r des Dysteriidae. C. R . Acad. Sci., Paris 267, 954-7 (with J. Andre &
M.-C. Ganier).
( 1 7 2 ) 1968. Structure corticale dune amibe Cdaphique. Protistologica 4, 195-208 (with J. AndrC).
(173) 1968. Calcification tkgumentaire chez les ciliCs du genre
Coleps Nitzsch. J . Micros. 7, 693-704 (with J. AndrC & M.-C.
Ganier)
(174) 1969. Remarques sur la systkmatique des ciliCs Oligotrichida. Protistologica 5, 345-52.
(175) 1969. Morphologie et structure fine du ciliC Chaenea
vorax Quenn. Protistologica 5, 353-61 (with M.-C. Ganier).
(176) 1970. Structure fine du Strombidium sulcatum C1. et
L. (Ciliata Oligotrichida) . Protistologica 6, 207-23 (with M.-C.
Ganier)

Smith, C. E. G., ed. 1972. Research in Diseases of the Tropics.

Brit. M e d . Bull., published by T h e British Council, 97 and 99
Park St., London W1Y 4HQ, England. 98 pp. $6.50.
For the first time since at least 1955, the British Medical
Bulletin, a review journal, discusses some protozoan diseases.
There are 17 papers on tropical diseases in this issue, of which
5 deal with protozoan diseases. These are Immunology of malarial infection and its possible consequences by I. A. McGregor
( 6 pages), Advances in malariology relating t o control and
eradication by W. Peters ( 6 pages), Trypanosomiasis by W. H.
R. Lumsden (5 pages), Leishmaniasis in the O l d Worl d by R.
S. Bray (5 pages), and Leishmaniasis of the N e w W o r l d : Taxonomic problems by R. Lainson and J. J. Shaw ( 5 pages). Anyone who deals with Plasmodium, Trypanosoma, or Leishmania
should read these reviews.
The paper by McGregor is considerably shorter than his
review of the same subject in the International Review of
Tropical Medicine, but gives a few more recent references; it

deals exclusively with Plasmodium falciparum. Peters deals

primarily with chemical agents both against the vectors and
the protozoa; propoxur and carbaryl (both carbamates) are
promising against the former and diformoldapsone against the
latter, although we still have no single compound to recommend
against drug-resistant P. falciparum. Lumsdens review cites only
key references, but mentions a number of reviews and abstracts
which give more. Brays review, among other things, brings out
the recent delineation of diffuse cutaneous leishmaniasis in
Ethiopia and advances in our knowledge of the immunology of
cutaneous leishmaniasis. Lainson and Shaw introduce a new
classification of the vexing New World leishmanias: the L.
mexicana complex, including L . m. mexicana, L. m . amazonensis
n. ssp., L. m . pifanoi and L . enriettii; and the L . braziliensis
complex, including L . b. braziliensis, L. b. guyanensis, L. b.
panamensis n. ssp. and L . peruviana; details about these subspecies and species can be obtained from their paper.-NORMAN D. LEVINE, College of Veterinary Medicine, University
of Illinois, Urbana, Ill. 61801.