Charles Philippe Leblond
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2008-05-20T20:35:09Z
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{{Infobox_Scientist
| name = Charles Philippe Leblond
| image = Cpl003 small.jpg
| caption = Charles Philippe Leblond, Canadian [[biologist]]
| birth_date = [[February 5]], [[1910]]
| birth_place = [[Lille]], [[France]]
| death_date = {{death date and age|2007|4|10|1910|2|5}}
| death_place = [[Montreal]], [[Quebec]], [[Canada]]
| residence = {{flag|Canada}}
| nationality = {{flagicon|Canada}} [[Canada|Canadian]]
| field = [[Biology]]
| work_institution = [[McGill University]]
| alma_mater = [[University of Paris]]</br>[[University of Montreal]]</br>[[Sorbonne]]
| doctoral_advisor =
| doctoral_students =
| known_for = [[Autoradiography]]</br>[[stem cell|Stem cell discovery]]
| prizes = [[Flavelle Medal]] (1961) </br>[[Gairdner Foundation International Award|Gairdner Foundation Award]] (1965)
| footnotes =
}}
'''Charles Philippe Leblond''', [[Order of Canada|C.C.]], [[Doctor of Medicine|M.D.]], [[Doctor of Philosophy|Ph.D]], [[Doctor of Science|D.Sc]], [[F.R.S.C.]], [[F.R.S.]], ([[February 5]], [[1910]] – [[April 10]], [[2007]]) was a pioneer of [[cell biology]] and [[stem cell]] research and a former Canadian professor of anatomy. Leblond is notable for developing [[autoradiography]] and his work showing how cells continuously renew themselves, regardless of age.
==Main research interests==
In 1946, Leblond found that, when he poured liquid [[photographic emulsion]] on a [[histological]] section containing a radio element, the emulsion was eventually activated by the radio-element; and if thereafter routine photographic development and [[Fixation (histology)|fixation]] were applied to the emulsion-covered section, black silver grains appeared in the emulsion wherever it overlay sites containing a radio-element. This liquid emulsion approach has been used to develop a new High Resolution Autoradiography procedure<ref> Bélanger, L.F. and C.P. Leblond. A method for locating radioactive elements in tissues by covering histological sections with a photographic emulsion. ''Endocrinology''. 1946, 39, 386-400.</ref> characterized by close contact between emulsion and section. Such close contact makes it possible to localize the radio-elements in the section at high [[Image resolution|resolution]], so that radio-elements can be localized at high magnification in the [[light microscope]].<ref> Gross, J., R. Bogoroch, N.J. Nadler and C.P. Leblond. The theory and methods of the radioautographic localization of radio elements in tissues. ''Amer. J. Roentgenoi.'' 1951, 65, 420-468.</ref><ref> Kopriwa, B. and C.P. Leblond. Improvements in the coating technique of radioautography. ''J. Histochem. Cytochem.'' 1962, 10, 269-284.</ref>
This procedure has been utilized to examine some of the dynamic features of body components, with the main findings as follows:
# The existence of stem cells in adult organs, as shown by autoradiography with labeled [[thymidine]].
# The continuity of [[Protein biosynthesis|protein synthesis]] in living cells, as shown by autoradiography with labeled [[amino acids]].
# The key role of the [[Golgi apparatus]] in protein [[glycosylation]], as shown by autoradiography with labeled [[hexose]]s.
His results threw doubt on the validity of three traditional concepts dear to biologists in the earlier part of the twentieth century: "stability" of the cell, in which the cell and its components are unchanging, permanent structures; "specificity" of cell function, in which each cell type has a distinct, unique function; and "activity-rest alternation" of cell function, in which each period of cellular activity is followed by a period during which the activity ceases.
He has proposed replacing cell "specificity" by "multipotentiality", "activity-rest alternation" by "continuity" and "stability" of cell components by "renewal". These various results have provided the foundation not only for modern stem cell research, but also for modern cell biology.
As Nobel laureate [[George Palade]] noted on the occasion of the 1992 [[Prix Marie-Victorin]] to Leblond, Charles Leblond’s discoveries are so fundamental that they are taught in schools and colleges throughout the world.<ref>{{cite web|url=http://www.prixduquebec.gouv.qc.ca/recherche/desclaureat.asp?noLaureat=159|title=Les Prix du Québec: CP Leblond}}</ref>
==Biography==
===Early days===
CP Leblond was born in [[Lille]], [[France]], in 1910, the son of a building contractor who died when Leblond was only 10 years old, leaving his mother to raise four boys on her own. A brilliant student, Leblond debated becoming a film producer, an architect or a scientist. In the end, he decided on science, and enrolled in Medicine at the [[University of Paris]]. He was fascinated by his first course in [[histology]] and decided to pursue this field as a career.
Leblond obtained his M.D. degree from the [[University of Paris]] in [[1934]]. His doctoral thesis described the histochemical localizion of [[ascorbic acid]], which he found to predominate in steroid-secreting cells.<ref>A. Giroud et C.P. Leblond. Etude histochimique de la vitamine C dans la glande surrénale. ''Arch. Anat. microse.'' 1934, 30, 105 129.</ref> This study led him, with a [[Rockefeller Foundation|Rockefeller postdoctoral fellowship]] in hand, to the [[endocrinology]]-orientated Department of [[Anatomy]] at [[Yale University]] in 1935, where he carried out studies on factors influencing maternal behavior.<ref>.C.P. Leblond. Extra hormonal factors in maternal behavior. ''Proc. Soc. Exp. Biol. Med.'' 1938, 38, 66 70.</ref> It was here that he met his wife Gertrude Sternschus, to whom he was married for 64 years.
In 1937, Leblond joined the Laboratoire de Synthese Atomique in Paris which was involved in preparing [[radioactive isotopes]] for use in investigating the fate of various molecules in biological processes. Under the guidance of Antoine Lacassagne, Leblond injected [[iodine|radioiodine-128]], into a [[rat]] and found that the label promptly accumulated in the [[thyroid gland]], presumably incorporated into the thyroid hormone precursor [[thyroglobulin]].<ref>C.P. Leblond et P. Sue. Passage de l'iode radioactif (1128) dans la thyroide stimulée par l'hormone thyréotrope de l'hypophyse. ''C.R. Soc. Biol.'' 1940, 133, 543.</ref> To localize this label more precisely within the thyroid tissue, Leblond attempted to use the novel technique of [[autoradiography]].
Unfortunately, Leblond’s first attempt to use autoradiography failed, the reason being that the radioiodine-128 isotope, with its extremely short [[half-life]] (25 minutes), disintegrated so quickly that too little [[radioactivity]] remained to be detected by the photographic emulsion.
===Development of autoradiography===
In 1941, Leblond moved to [[McGill University]] as a lecturer in histology, and quickly rose to assistant (1943), associate (1946), and then full professor of anatomy (1948). He served as the chair of the Department of Anatomy from 1957–1974.
At McGill, Leblond used the newly-available [[iodine-131|radioiodine-131]] with a half-life of 8 days, to repeat his autoradiographic experiment on thyroid tissue. With this method, the resolving power was less than 100 µm, but nonetheless he was able to localize the radioactivity to specific thyroid [follicles].<ref>C.P. Leblond. Extra hormonal factors in maternal behavior. ''Proc. Soc. Exp. Biol. Med.'' 1938, 38, 66 70.</ref>
Leblond’s early career at McGill was interrupted by [[World War II]], during which he served in the [[Free French Forces]]. He was dispatched first to [[Rio de Janeiro]], then to [[London]], where he conducted medical exams of would-be soldiers.
"In 1946, after returning to Montreal from service with the Free French Forces, it was clear to me that the crude technique previously used for radioautography had to be improved".<ref>Leblond CP. The time dimension in cell biology. ''FASEB J.'' 1995 Sep;9(12): 1234-8.</ref> In collaboration with Leonard Bélanger, Leblond worked on increasing the [[Optical resolution|resolution]] of the autoradiographic technique. They were advised by physicist Pierre Demers to melt the emulsion from [[Eastman Kodak]] lantern slides, paint it directly on the sections, and then develop the emulsion while it was still attached to the histologic sections. This resulted in a tenfold improvement in resolution.<ref>F. Bélanger and C.P. Leblond. A method for locating radioactive elements in tissues by covering histological sections with a photographic emulsion. ''Endocrinology'' 1946, 39, 386 400.</ref> Subsequently, Leblond and his colleagues developed a technique in which the histologic slides were dipped directly into liquid emulsion.<ref>C.P. Leblond and J. Gross. Thyroglobulin formation in the thyroid follicle visualized by the "coated autograph" technique. ''Endocrinology'' 1948, 43, 306 324.</ref> The use of thinner sections and emulsion coats led to further advances in resolution, and the introduction of tritium was a technical milestone.
High Resolution Autoradiography procedure continues to be used today by molecular biologists to detect [[RNA]] molecules in situ, and to study the localization of [[genes]] and [[DNA]] sequences.
===Studies on the turnover of cells===
Leblond used autoradiography to introduce radioactive precursors of DNA, and then examine the renewal and fate of cells of several basic tissue types. He demonstrated for the first time that most cells and tissues in the adult body undergo continued renewal. Using mathematical models and modern methods of quantitation, Leblond and his colleagues estimated with remarkable accuracy the turnover and [[mitotic]] rates of numerous cell types. He and his colleagues made fascinating discoveries that resulted in the introduction of "time dimension" to cells and tissues, opening the doors to the understanding of the cell cycle and to the identification of stem cells.
===Identification of Stem Cells in Adult Organs===
In the male [[seminiferous]] epithelium, studies by Leblond and Yves Clermont in the early 1950’s had deciphered how [[spermatogonia]] gave rise to [[spermatocytes]], which then differentiated into mature sperm cells in a specific cycle.<ref>C.P. Leblond and Y. Clermont. Definition of the stages of the cycle of the seminiferous epithelium of the rat. ''Ann. NY Acad.'' Sci. 1952, 55, 548 573.</ref><ref name="schiff">Y. Clermont and C.P. Leblond. Spermiogenesis of man, monkey, ram and other mammals as shown by the "periodic acid Schiff" technique. ''Am. J. Anat.'' 1955, 96, 229 250.</ref>
To maintain the population of spermatogonia, the seminiferous epithelium was shown to contain a population of stem cells which divided to produce differentiated cells as well as to maintain their own number. As noted in a seminal publication by Leblond, "the reappearance at each cycle of a new dormant cell which acts as the stem cell of [[spermatocyte]]s is described as the 'Stem Cell Renewal Theory'" . This article is the first one in which nests of cells dividing in an adult organ are designated as “stem cells”.<ref>Y. Clermont and C.P. Leblond. Renewal of spermatogonia in the rat testis. ''Am. J. Anat.'' 1953, 93, 475 502.</ref>
Leblond and his colleagues also found evidence for the presence of occasional adult stem cells even in tissues which are composed almost entirely of non-dividing cells. In [[skeletal muscle]], the muscle fibers had been shown to exhibit an age-related increase in the number of [[Cell nucleus|nuclei]].<ref>M. Enesco and C.P. Leblond. Increase in cell number as a factor in the growth of the organs and tissues of the young male rat. ''J. Embryol. Exp. Morphol.'' 1962, 20, 530 562.</ref> His studies showed that muscle satellite cells could be considered to be adult stem cells in muscle fibers.
From the studies of Leblond and his colleagues, it was concluded that the body has three types of cell populations:
#"Static cell populations", which are composed of non-dividing cells and include no adult stem cells. These populations have the "stability" formerly attributed to all cells
#"Expanding cell populations" in which small numbers of [[adult stem cell]]s exist, and give rise to skeletal fiber nuclei or [[glia]]l cells of the brain
#"Renewing cell populations" in which adult stem cells are an essential feature
To mark his 65th birthday in 1975, Leblond was honored at an international symposium on the existence of stem cells in adult tissues; the resulting book, Stem Cells of Renewing Cell Populations, was the first formal, comprehensive account on the subject.<ref>Cairnie A.B., Lala P.K. and D.G. Osmond. ''Stem Cells of renewing Cell Populations'' Academic Press. N.Y. 1976.</ref>
===Continuous Protein Synthesis in living Cells===
When Leblond and his colleagues used 14C-bicarbonate, and then 35S-labeled amino acids to investigate protein synthesis, they were astonished to find that virtually all cells in the body incorporated label.<ref>R.C. Greulich and C.P. Leblond. Radioautographic visualization of radio carbon in the organs and tissues of newborn rats following administration of C14 labeled bicarbonate. ''Anat. Rec.'' 1953, 115, 559 586.</ref><ref>C.P. Leblond, N.B. Everett and B. Simmons. Sites of protein synthesis as shown by radioautography after administration of S35 methionine. ''Am. J. Anat.'' 1957, 101, 225 271.</ref> This led them to the conclusion, considered heretical at the time, that all cells continually synthesized proteins. This was among the first evidence to replace the Specificity concept with the idea that most cells are [[multipotent]]ial in their functions.
It is of interest that autoradiographic studies by Leblond during this period also settled a controversy regarding the cellular site of synthesis of ribonucleic acid. Using radiolabeled cytidine in some forty cell types, he and his colleagues were the first to demonstrate decisively that RNA is continuously synthesized in the nucleus and then migrates to the cytoplasm.<ref>M. Amano and C.P. Leblond. Comparison of the specific activity time curves of ribonucleic acid in chromatin, nucleolus and cytoplasm. ''Exp. Cell Res.'' 1960, 20, 250 253.</ref><ref>M. Amano, C.P. Leblond and N.J. Nadler. Radioautographic analysis of nuclear RNA in mouse cells revealing three pools with different turnover times. ''Exp. Cell Res.'' 1965, 38, 314 340.</ref>
===Role of the Golgi Apparatus in Protein Glycosylation===
A majority of body proteins is [[glycosylated]], although the proportion of [[carbohydrates]] in [[proteins]] is quite variable. Leblond had shown in earlier studies that the [[Golgi|golgi apparatus]] region in most cell types was dramatically stained by the periodic acid-Schiff staining technique, which is specially directed to carbohydrate-rich proteins carrying 1,2 glycols.<ref name =schiff/><ref>C.P. Leblond. Distribution of periodic acid reactive carbohydrates in the adult rat. ''Am. J. Anat.'' 1950, 86, 1.</ref> In the electron microscope, using the periodic acid silver technique, there was a gradient of staining intensity from the [[cis]] to the [[trans]] side of the Golgi apparatus, suggesting that carbohydrate residues were added to proteins at this site.<ref>A. Rambourg, W. Hernandez and C.P. Leblond. Detection of periodic acidreactive carbohydrate in Colgi saccules. ''J. Cell Biol.'' 1969, 40, 395 414.</ref>
To test this hypothesis, light and then EM autoradiographic studies were carried out by Leblond and Neutra in 1966 after injecting rats with 3H-glucose or 3H-galactose.<ref>M. Neutra and C.P. Leblond. Synthesis of the carbohydrate of mucus in the [[Golgi complex]], as shown by electron microscope radioautography of goblet cells from rats injected with 3H-glucose. ''J. Cell Biol.'' 1966, 30, 119 136.</ref><ref>M. Neutra and C.P. Leblond. Radioautographic comparison of the uptake of 3H-galactose and 3H glucose in the Golgi region of various cells secreting glycoproteins or mucopolysaccarides. ''J. Cell Biol.'' 1966, 30, 137 150.</ref> Within ten minutes, the label was dramatically localized to the Golgi apparatus of intestinal goblet cells, indicating that this was the cellular site of addition of sugar residues in the synthesis of the carbohydrate side chains of mucous glycoproteins.
This discovery had a tremendous impact on the scientific community, being the first evidence for a functional role of the Golgi apparatus in the synthetic process.
===Other studies===
Other classics include: identification of how skeletal bones grow through osteoblast deposition and osteoclast remodeling,<ref>Leblond CP, Wilkinson GW, Belanger LF, Robichon J. Radio-autographic visualization of bone formation in the rat. ''Am J Anat''. 1950 Mar;86(2):289-341.</ref> early discovery of the biogenesis and metabolism of thyroxine<ref>Gross J, Leblond CP. Metabolism of the thyroid hormone in the rat as shown by physiological doses of labeled thyroxine. ''J Biol Chem.'' 1950 Jun;184(2):489-500.</ref> and detection of triiodothyronine,<ref>Gross J, Leblond CP. The presence of free iodinated compounds in the thyroid and their passage into the circulation. ''Endocrinology.'' 1951 Jun;48(6):714-25.</ref> early prediction of DNA semiconservative replication<ref>Stevens CE, Daoust R, Leblond CP. Rate of synthesis of deoxyribonucleic acid and mitotic rate in liver and intestine. ''J Biol Chem.'' 1953 May;202(1):177-86.</ref> published days after the Watson and Crick Nature article,<ref>J. D. Watson and F. H. C. Crick. A Structure for Deoxyribose Nucleic Acid. April 25, 1953, ''Nature'', 171, 737-738</ref> the discovery of axonal transport,<ref>Droz B, Leblond CP. Migration of proteins along the axons of the sciatic nerve. ''Science.'' 1962 Sep 28;137:1047-8.</ref> the Warshawsky et al<ref>Warshawsky H, Leblond CP, Droz B. Synthesis and migration of proteins in the cells of the exocrine pancreas as revealed by specific activity determination from radioautographs. ''J Cell Biol.'' 1963 Jan;16:1-24.</ref> finding that nascent proteins are processed from the rough endoplasmic reticulum through the Golgi apparatus into pancreatic zymogen granules (made in hot competition with the Palade lab at Rockefeller University), the first realization that the Golgi apparatus is the site of terminal glycosylation,<ref>Peterson M, Leblond CP. Synthesis of Complex Carboydrates in the Golgi region, as shown by radioautography after injection of labeled glucose. ''J Cell Biol.'' 1964 Apr;21:143-8.</ref> the discovery of the cell coat,<ref>Rambourg A, Neutra M, Leblond CP. Presence of a "cell coat" rich in carbohydrate at the surface of cells in the rat. ''Anat Rec.'' 1966 Jan;154(1):41-71.</ref> the cellular biogenesis of collagen,<ref>Weinstock M, Leblond CP.Synthesis, migration, and release of precursor collagen by odontoblasts as visualized by radioautography after (3H)proline administration. ''J Cell Biol.'' 1974 Jan;60(1):92-127.</ref> and new insights into the ultrastructure of basement membrane.<ref name=polymer>Inoue S, Leblond CP, Laurie GW. Ultrastructure of Reichert's membrane, a multilayered basement membrane in the parietal wall of the rat yolk sac. ''J Cell Biol.'' 1983 Nov;97(5 Pt 1):1524-37.</ref>
==="Retirement" and later days===
At 65, instead of retiring, Leblond continued his research with an [[NIH]] Fogarty Scholarship at the [[National Institute of Dental Research]], where he learned about [[immunohistochemistry]]. This launched a twenty-year molecular exploration culminating in the concept of the basement membrane as an integrated [[polymer]],<ref name=polymer/> rather than as layers of separated [[macromolecules]] initially favored by others.
Leblond continued to attend all weekly departmental seminars well into his 90’s and continued to publish in peer-reviewed journals into the new millennium. He learned to use a computer at age 90, starting a presentation at an international conference back in 2004, by noting: "A month ago, I thought [[Power Point]] was a tool for sharpening pencils."
His total contributions resulted in the publication of 430 scientific papers, many of them still frequently cited. In late September 2006, he published his final article—about detecting the [[MMP9]] [[cysteine]] activation switch for the first time in remodeling [[cartilage]].<ref>Lee ER, Lamplugh L, Kluczyk B, Mort JS, Leblond CP. Protease analysis by neoepitope approach reveals the activation of MMP-9 is achieved proteolytically in a test tissue cartilage model involved in bone formation. ''J Histochem Cytochem.'' 2006 Sep;54(9):965-80. Epub 2006 May 18.</ref>
Leblond was preceded in death by his wife of 64 years, Gertrude Sternschuss, who died in 2000. After Gertrude died, Leblond married a childhood friend, Odette Lengrand, in 2001; they were both 91. Odette died in 2004.
==Honors==
===Honorary Degrees of Doctor of Sciences===
* [[Acadia University]], 1972
* McGill University, 1982
* [[University of Montreal]], 1985
* [[York University]], 1986
* [[Sherbrooke University]], 1988
===Prizes===
* Prix Saintour, French Academy, 1935
* [[Gairdner Foundation International Award]], 1965
* Isaac Schour Award, [[International Association for Dental Research]], 1974
* Henry Grey Award, American Association of Anatomists, 1978
* J.C.B. Grant Award, Canadian Association of Anatomists, 1979
* E-B Wilson Award, [[American Society for Cell Biology]], 1982
* Duncan Graham Award, Royal College of Physicians and Surgeons of Canada, 1986
* Centennial Award, American Association of Anatomists, 1979
* [[Prix Marie-Victorin]], [[Quebec]] Province, 1992
===Medals===
* [[Flavelle Medal]], [[Royal Society of Canada]], 1961
* Medal Léo-Pariseau, «Assoc. Canadienne Française pour l’Avancement des Sciences», 1962
* McLaughlin Medal, Royal Society of Canada, 1983
* George Gomori Medal, Histochemical Society, 1988
===Other honours===
* 1951 – Fellow of the [[Royal Society of Canada]]
* 1965 – Fellow of the [[Royal Society]], London, UK
* 1970 - [[American Academy of Arts and Sciences]]
* 1988 - [[List of organisations in the United Kingdom with a royal charter|Royal Microscopical Society]], UK, 1988
* 1995 – [[Canadian Medical Hall of Fame]]
* 2000 – Companion, [[Order of Canada]]
* 2001 – Grand Officer, [[National Order of Quebec]]
==Trivia==
Periodic acid Schiff stain whose lively [[purple]]-[[magenta]] coloring of carbohydrate 1-2-glycol groups20,21 inspired Leblond’s Golgi and cell coat discoveries also elegantly inspired the Leblond wardrobe, automobile and even home interior and country home name – Val Mauve.
One of Leblond’s continuing interest was his endeavour to ensure the usage of the correct name for the technique which he had spent a lifetime developing. In a review chapter written in 1987, entitled: “Radioautography: The role played by anatomists in the development and application of the technique.”,<ref>C.P. Leblond. Radioautography: the role played by anatomists in the development and application of the technique. In: The American Association of Anatomists, 1888-1987. Essays on the History of Anatomy in America and a Report on the Membership – past and present. John E. Pauly, ed. Williams and Wilkins, Baltimore, 1987, pp 89-103</ref> he writes: The reasons why the term “radioautography” is preferred to “autoradiography” for the detection of radioactive elements by photographic emulsion are as follows. The term “autoradiography” is a compound word including the term “radiography”. This term is defined as a picture produced by an x-ray beam that has passed through an object. Since this object, for instance a bone examined after a fracture, is located between the source of radiation and the emulsion, it appears white in the emulsion; that is, it is seen as a negative image. In contrast, when radioactive elements are seen in sections, the object under study is itself the source of the radiation that influences the emulsion. The black image thus produced is a photographic positive. It may be referred to as an autograph, that is “the reproduction of form or outline of anything by an impression from the thing itself” (Oxford English Dictionary, 1975). Hence, the author called it initially a “radioactive autograph”. Later, on the advice of an editor, he condensed these two words into “radioautograph”. The procedure is often called “autoradiography” but “radioautography” is the correct term.
Leblond had 4 children for which, in a true [[Cartesian]] manner, he chose names starting with the letter "P": Philippe, Paul, Pierre and (Marie)-Pascale. He also had 7 grandchildren.
==References==
{{reflist}}
==External links==
* [http://www.thecanadianencyclopedia.com/index.cfm?PgNm=TCE&Params=A1ARTA0004600 Charles Philippe Leblond] at [[The Canadian Encyclopedia]]
* [http://www.cdnmedhall.org/laureates/?laur_id=21 Charles Philippe Leblond] at The Canadian Medical Hall of Fame
* [http://www.gg.ca/honours/search-recherche/honours-desc.asp?lang=e&TypeID=orc&id=974 Order of Canada]
* [http://www.prixduquebec.gouv.qc.ca/recherche/desclaureat.asp?noLaureat=159 Prix Marie-Victorin Ceremony] (in French)
* [http://www.ordre-national.gouv.qc.ca/recherche_details.asp?id=421 National Order of Quebec] (in French)
{{DEFAULTSORT:Leblond, Charles}}
[[Category:1910 births]]
[[Category:2007 deaths]]
[[Category:People from Lille]]
[[Category:Canadian Medical Hall of Fame]]
[[Category:Canadian scientists]]
[[Category:Canadian medical researchers]]
[[Category:Canadian biologists]]
[[Category:Companions of the Order of Canada]]
[[Category:Fellows of the Royal Society of Canada]]
[[Category:Fellows of the Royal Society]]
[[Category:Foreign Members of the Royal Society]]
[[Category:Grand Officers of the National Order of Quebec]]
[[Category:McGill University faculty]]
[[Category:University of Paris alumni]]
[[fr:Charles Philippe Leblond]]