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Home , Albert Claude, David Baltimore, Francisc Rainer, George Emil Palade, Journal of Cell Biology, Jules Bordet

REVIEW ARTICLES REVIEW –A LIFE FOR , REMEMBER

Mirela E. CADAR* and Mircea GABOREANU

Faculty of Animal and , University of Agricultural and Veterinary , 3-5 Mănăştur Street, 400372 Cluj-Napoca, *Corresponding author, email: [email protected]

Bulletin UASVM Animal Science and Biotechnologies 71(2) / 2014, Print ISSN 1843-5262; Electronic ISSN 1843-536X DOI: 10.15835/buasvmcn-asb:10484

Abstract Four decades ago, the Nobel Prize for Medicine and honored and rewarded the activity in Cellular and domain of three researchers, (1899-1983), (1917-2013) and George Emil Palade (1912-2008). Viewing the life and activity history of GE Palade comes out his respect for a certain research ethics and his demeanor for his very numerous collaborators and disciples.

Keywords: cellular biology, molecular biology, Nobel Prize, GE Palade

INTRODUCTION and . His PhD thesis was of Four decades ago, the Nobel Prize for Medicine anatomy domain with the theme “The nephron of and Physiology honored and rewarded the activity cetacean Delphinus delphi”. in Cellular and Molecular Biology domain of three After university graduation, GE Palade wor- researchers, two Belgians –Albert Claude (1899- ked for a short time as assistant in the Clinic 1983) and Christian De Duve (1917-2013) and of Internal Diseases, and than he returned at an American of Romanian origin –George Emil Anatomy Discipline, where in few years became Palade (1912-2008). The motivation of Nobel assistant, lecturer and associated professor. Committee was “For discoveries concerning the The chief of Anatomy Discipline was Professor functional organization of the that were seminal Grigore T. Popa, which in 1930 during a research events in the development of modern ”. stage at Cambridge University has discovered and described with Unna Fielding the hypophysial 1912, Iaşi Romania portal vessel system. GE Palade was born in , 1912 in GE Palade married in 1944 Irina Malaxa, Iaşi, Romania, in a family in which both parents physician and daughter of Nicolae Malaxa –the (father Ioan Emil Palade, 1885-1961 and mother magnate of heavy Romanian industry. In 1945 was Constanţa Cantemir Palade, 1886-1976) and born Georgia, the irst child, and the second one also grandparents had intellectual professions Philip, will be born in 1957 in USA. (professors, schoolteachers, reverends). GE Palade had two sisters, Adriana who became Geography 1946, New York USA teacher and Constanţa, pediatric physician. In 1946, GE Palade with his wife and daughter emigrated in USA, with the wish to continue their 1940, Romania studies. Having a letter of introduction from After graduation of high school in 1929, GE professor Grigore T. Popa to Robert Chambers Palade was admitted at Carol Davila School of from , Department of Biology, Medicine from Bucharest, Romania and received GE Palade was accepted as “Visiting Investigator”. his MD in 1940. During the irst university Here, during few months, he continued his years, GE Palade revealed a special interest for morpho-functional study of kidney, for this time 78 CADAR et al on in vitro cultures of renal tissue from chicken Sanford Palay, the botanist Ruth Sager and a long- embryos, and the results guiding to his irst distance collaborator (from 1954 until 1961) the published article in USA in 1947. In the same biochemist , all together forming year, being invited by Albert Claude, GE Palade “the second Rockefeller group”. goes to Rockefeller Institute for Medical Research, During the activity of this group, GE Palade where he will stay 25 years. There, Albert Claude described for the irst time the structure of had also as collaborators the naturalist Keith with all its aspects from different Porter, the physician George Hogeboom and the cellular type, named cristae mitochondriales biochemist Walter Schneider. The preoccupations the prolongations inside them and demonstrated of this researchers’ group, which GE Palade named that mitochondria are permanent in “the irst Rockefeller group”, were focused mainly cytoplasm of all eukaryotic cells. in two directions: cell fractionation for When he published these data, GE Palade be- separation and their biochemical analysis and bio- came a naturalized American citizen. logical sample examination in electronic micro- Together with Keith Porter, GE Palade continued scope. the study of , for this time Albert Claude initiated cell fractionation and on ultraine slides, describing its two aspects: ultracentrifugation of cellular homogenate since rough or granular endoplasmic reticulum and 1940, and he succeeded to isolate only small smooth or agranular endoplasmic reticulum. vesicular structures, named microsomes. They studied the same organelle in muscular GE Palade improved this technique perfor- ibers and named it sarcoplasmic reticulum. mance conceiving the procedure known as GE Palade with Philip Siekevitz had demons- “ultracentrifugation in sucrose density gradient”. trated that cytoplasmic subcellular structures Albert Claude with Keith Porter used for the described by Keith Porter and named microsomes, irst time in 1947 an electronic microscope of are in reality vesicular fragments derived from industrial type for cell study. Examining a single- endoplasmic reticulum during cells’ fractioning. layer cell culture they described in cells’ very thin But, one of the most important discoveries cytoplasm a structure with lace aspect, vaguely of GE Palade was that of cytoplasmic structures, contoured, which was named endoplasmic which he named “small granular components of reticulum. cytoplasm” being either free in cytosol, isolated In Rockefeller Institute the irst electronic or grouped or attached to endoplasmic reticulum microscope was made in function in 1948. It use surface. These structures were subsequently in biological research imposed ultraine sections, named granules of Palade, ribonucleoproteic thick of only 50-150 nm, and a more correct particles, and nowadays are named . chemical ixation of subcellular structures, which The study of endoplasmic reticulum and allows obtaining optimal resolution on images. ribosomes was continued as biochemical one, in These dificulties were surpassed when was collaboration with Philip Siekevitz. Were developed invented the ultramicrotome by Keith Porter and experiments with tracer atom aminoacids marked Johan Blum and the ixation method of biological with radioactive isotopes concluding that these samples with a osmium tetraoxyde buffered with cytoplasmic structures have as main function the sodium veronal at 7.0-7.5 pH, conceived by GE protein synthesis. Palade and known as “ixative of Palade”. In same period GE Palade, having as labora- With these means, “the irst Rockefeller group” tory chief the neurophysiologist Herbert Gasser achieved the isolation of mitochondria from (Nobel Prize 1944) targeted his researches hepatocytes (1947), their cytochemistry and also on electrono-microscopic study of nervous demonstration of presence in tissue. For the irst time there were marked out somatic cells (1949). the nervous fibers of C type leading the pain In 1949, the researchers’ group from sensation, and was named the mesaxon as an Rockefeller Institute diminished by departure annexed structure of them. The collaboration with of Albert Claude, Walter Schneider and George Sanford Palay, also neurophysiologist, gave to GE Hogeboom, but with Keith Porter and GE Palade Palade the opportunity to demonstrate the fact came the neurophysiologists Herbert Gasser and that Nissl bodies or granulations from optical

Bulletin UASVM Animal Science and Biotechnologies 71(2) / 2014 Review –A life for Nobel Prize, remember George Emil Palade 79 microscopy are in electron microscope dense Blobel. Starting 1978, there came also 14 post- packs of lattened sacks of granular endoplasmic doctoral physicians, geneticists and biochemists reticulum. GE Palade was also those one who irst from Romania. In next 12 years of activity, the described the inter-neuronal and neuromuscular main preoccupations of GE Palade with mentioned synapse ultrastructure, with the two portions, collaborators were: the intracellular aspects of pre- and post-synaptic elements, the presence of protein process, the ultrastructure of synaptic vesicles and existence of synaptic cleft, of capillaries, the mechanisms thus demonstrating the contiguity relation and of capillaries’ permeability and cytomembrane not continuity one between neurons. biogenesis. Collaboration with the botanist Ruth Sager During 1962-1964, GE Palade in collaboration leaded to study the ultrastructure with Lars Ernster, Lewis Joel Green, Johan Kirsch, from green algae, thus demonstrating that beside Colvin Redman, Gustav Dallner, Any Leskes and them, in vegetal cells, is also present the endoplas- more often with Lucien Caro and James Jamieson mic reticulum. effected majority of experiments through of were In 1957, GE Palade in collaboration with proved the protein secretion phases in glandular the anatomist Benet Russel conceived the me- cells of exocrine pancreas, both in vivo and in thod with potassium telurit to make evident vitro. In this study unless electron microscopic dehydrogenases’ activity in electron microscope, examinations were done complementary techni- fact that allows for them to ascertain the activity ques as: cellular fractionation, separation of intensity of these enzymes inside mitochondria. cellular components (endoplasmic reticulum, In same period, GE Palade made with the Golgi vesicles, mitochondria, secretion vesicles biochemist Philip Siekevitz a biochemical study and granules etc.) by ultracentrifugation in of endoplasmic reticulum and of ribosomes sucrose density gradient. Also, were used as from cells of Guinea pig exocrine pancreas. tracer atoms of some aminoacids marked with Pursuant to intercessions of Keith Porter and radioactive isotopes as [3H[-leucine, by pulse- GE Palade starting of 1955 appeared the Journal chase analysis followed by autoradiographic of Biochemical and Biophysical Cytology. In examination in electron microscope. In this way further years, GE Palade was member in Editorial can be characterized the synthesis phase of Board of Journal of Molecular Biology, Journal of polypeptides at level, the segregation Membrane Biology, and phase of them inside cisternal endoplasmic Annual Review of Cell Biology. reticulum, the intracellular transport phase by In 1961, “the second Rockefeller group” chan_ transitional elements of endoplasmic reticulum ged its composition because of departure of Keith and peripheral Golgi vesicles, the concentration Porter and Philip Siekevitz. The laboratory activity phase in condensing vesicles, the intracellular was further coordinated by GE Palade, to who storage phase in zymogen granules and, inally, joined in time a large number of researchers from the discharge phase with membrane fusion USA, Canada, , Italy, Germany, Hungary, and ission interactions, which take place in inal Israel, Great Britain and Romania. Among these, phase. some are professors with approved scientiic During 1963-1964, GE Palade in collaboration notoriety, as the anatomist Ewald Weibel, the with the biochemists of team had done cytoche- pathologist Guido Majno, the and mistry researches on exocrine pancreatic cells pathologist Marilyn Gist Farquhar, the physician from Guinea pig, rat, pigeon and bovines specifying Nicolae Simionescu and his wife biologist Maya the biochemical of condensing vacuoles, of Simionescu, the biochemists Lars Ernster, Gustav zymogen granules and those ones of secretion. In Dallner, Tsuneo Omura, Itzhak Ohad and Johan an interview for a journal of New Haven USA, GE Kirsch, and the biophysicist Lucien Caro. Other Palade summarized this activity saying “All of us we ones, scholars of post-doctoral studies, as Steve known that in an organism with evolved structure Wissing, Francesco Clementi, Rudolph Burns, and almost all cells product proteins. I identiied this many young medical under-graduated as James process”. Jamieson, Colvin Redman, Yutaka Tashiro, Lewis Also in 1964, GE Palade with James Jamieson Joel Green, Any Leskes, Alan Tartakoff and Günter were the irst ones which discovered and described

Bulletin UASVM Animal Science and Biotechnologies 71(2) / 2014 80 CADAR et al in electron microscope the presence in atrial cardiac in the open speech of Symposium “Junctional muscle of some secretion granules, which were Complexes of Epithelial Cells” in London 1987, GE named specific atrial granules. Subsequently in Palade and were named “grand 1981, De Bold and his collaborators established parents of intercellular jonctional systems”. that these granules contained a hormone, which In a morphometric study of pulmonary was named atrial natriuretic factor (ANF). arterial vascularization made in 1964 by GE During 1962-1967, GE Palade started the Palade and Ewald R. Weibel –anatomy professor in study of cellular membrane biogenesis with and the president James Jamieson, Philip Siekevitz, Gustav Dallner, of International Society of Stereology- marked Any Leskes, Itzhak Ohad, Tsuneo Omura, Jacopo out for the irst time in electron microscope the Maldonesi and Lars Ernster. Succeeding the existence of some “new components from arterial isolation of endoplasmic reticulum membranes endothelial cells” further named Weibel-Palade without ribosomes and their analysis on rat bodies. In present, it is known that these ones hepatocytes, can be estimated the membrane contain one of numerous blood clotting factors, proteins’ turnover. For the membranes of tilakoid named . bodies were used cells of green algae. Thus, can The researches of GE Palade concerning be demonstrated that in cells are not produced permeability of different blood capillary types new membranes (“de novo”), but the structural were started almost in 1961 when, in collaboration proteins and synthesized and segregated enzymes with Marylin Farquhar and Steve Wissing had are incorporated in pre-existent membranes’ tested the renal glomerular capillary permeability structure. From this phenomenon results the using as tracer atoms molecules of feritine. The continuity in time, the organization and experiments were continued in collaboration with functional continuity of cellular membranes. Rudoph Bursn on capillaries from muscles, with Another research theme that was developed Francesco Clementi on capillaries from intestine, great part concomitantly with previous ones was and with Guido Majno had studied the effect o f the study of blood capillaries’ endothelium and its histamine and serotonine on capillary permeability role in permeability process. GE Palade started this in case of inlammatory process. During these study from 1960 in collaboration with Rudolph experiments, GE Palade observed and described in Burns, Steve Wissing, Ewald R. Weibel, Francesco electron microscope an unprecedented transport Clementi and Guido Majno. But, the most articles modality of some molecules by plasmalema were published in collaboration with Marilyn G. invagination forming carrier vesicles, which were Farquhar, specialist in zoology and experimental named plasmalemal vesicles. The phenomenon , who came at Rockefeller Institute in was named transcytosis. 1961. She married GE Palade in 1970, after one The study of capillary endothelium permeabi- year of irst wife Irina death. lity became almost the unique preoccupation The irst researches targeted the renal with arrival in 1970 at Rockefeller Institute of the glomerular capillaries specifying the ultrastruc- physician Nicolae Simionescu and his wife, the ture of urinary filtration barrier and marking biologist Maya Simionescu, invited by GE Palade in out for the irst time the presence of a third cell 1970 when he visited Romania. The collaboration type, with phagocytary role, in connective matrix in this “famous Romanian trio”, as was named in of stalk region of the capillary tuft, which were specialty groups, subsisted 10 years. The test of named mesangial cells. endothelium permeability was done with anionic Also, on this capillary type, but in rats with and cationic molecular tracer atoms having nephritic syndrome, they had described for the different size molecules as: feritine, peroxydase, irst time a special type of occlusive intercellular myoglobine, dextran, glycogen and N-peptids. Came junctions, which were named occluding zonules. out that beside the plasmalemal vesicle pathway Extending the study of intercellular junctions the endothelium crossing is possible also by non- also on other epithelia types, GE Palade and Marilyn permanent transendothelial channels, which are Farquhar described other two junctions’ types, formed by certain vesicles’ junction. Also, came out which were named zonula adherens and macula that these transport ways depends on electrical adherens or desmosomes. For these discoveries, charge of transported molecules. Moreover, it

Bulletin UASVM Animal Science and Biotechnologies 71(2) / 2014 Review –A life for Nobel Prize, remember George Emil Palade 81 intervenes also the chemical interaction between (Nobel Prize 1975) wrote transportable molecules and certain receptors in his book “Molecular Cell Biology” (1986): from endothelial cell membrane, process that was “Many of the irst identiications of cellular named transcytosis mediated by receptors. All structures in electron microscopy as well some these observations constituted the start basis to biochemical characterizations of cellular functions study the arteriosclerosis ethiopathogeny further we do to researches of Albert Claude and his three done by Nicolae and Maya Simionescu at Institute collaborators: Christian De Duve, George Emil of Cellular Biology and Pathology from Bucharest, Palade and Keith Porter. The sophisticated image Romania. Given to the experience in this domain that we have about cell structure and function is, in of the two Romanian disciples of GE Palade, is not large part, the fruit of their research. They were the surprising the fact that Leon Weiss –the editor architects of modern Cellular Biology”. of well-known book “Cell and Tissue Biology” The title of this review limits the subject only (6th edition, 1988)- consigned the chapter “The for the life and work of GE Palade, without losing Cardiovascular System” to Simionescu family, and track of the fact that researches and discoveries of the chapter “The Exocrine Pancreas” to James the other two Nobel Prize laureates –Albert Claude Jamieson, which had the longest collaboration and Christian De Duve- had same importance for with GE Palade for protein synthesis study in modern Cellular Biology development and with pancreatic cells. practical consequences in cellular pathology. In 1973, the new manager of Rockefeller In , GE Palade had activated Institute, the physicist Frederick Steitz, do not between 1974 and 1990 as professor and chairman agree the existence of some large research team of Cellular Biology Division as well as Senior and the multi-disciplinary approaching of research Research Scientist of Cellular Biology Department. themes, fact that constituted a permanent cause His wife, Marilyn activated as of disagreements with GE Palade. Moreover, in of Cellular Biology. In this period, GE Palade and Institute were organized in addition 5 research his collaborators had continued the study of laboratories, all in Cellular and Molecular transcytotic transport system of large molecules Biology domain. This situation motivated GE through endothelial barrier, started in 1972. Palade to accept the invitation to establish and lead a Molecular Medicine Center and to teach 1990, San Diego , USA Cellular Biology for students in Yale University, In 1990, GE Palade moved at California San School of Medicine New Haven, Connecticut USA. Diego University, School of Medicine , The transfer was done in 1974, with GE Palade USA, where he activated as Professor of Medicine departing his wife Marilyn, James Jamieson, of Residence and as Dean of Scientiic Affairs in Nicolae and Maya Simionescu, David Castle and Cellular and Molecular Medicine Department, and Ann Hubbard. his wife Marilyn as professor of Pathology. In San Diego, GE Palade continued the 1974, New Haven, Connecticut USA researches for vectorial transport of intracellular The , New York, that in proteins with a numerous research team, among his history had four Nobel Prize laureates (Alexis who were the Romanian physicians Lucian Carrel-1912, -1930, Herbert Săucan and Dan Predescu. But, now instead of Gasser–1944 and Payton Rous–1966) had failed ultracentrifugation in sucrose density gradient, their continuation list with another three ones. considered out of date, was used the immune- And this was because in October 1974, were isolation procedure to separate the cytoplasm communicated the names of the three Nobel Prize components. Thus, can be followed the route of laureates for Physiology and Medicine: Albert synthesized proteins at ribosome level, in their Claude –manager of Institute from way to approximate 25 possible destinations Bruxelles, , Christian De Duve – professor (endoplasmic reticulum, Golgi complex, lysoso- of biochemistry in Catholic University of Louvain, mes, membranes etc.). It can be established Belgium, and George Emil Palade – professor in that each protein type possesses an encoded Yale University, New Haven USA. signal, formed by few aminoacids, which will be recognized by a speciic receptor from the

Bulletin UASVM Animal Science and Biotechnologies 71(2) / 2014 82 CADAR et al membrane of destination cellular component. Was fate, which we do not control, and which made our even succeeded the identiication and isolation of achievements possible”. such signals and receptors. Then, it is his demeanor given to very nume- Concomitantly, GE Palade was involved in rous colleagues and disciples, about whom he said promotion of Cellular and Molecular Biomedical that “they were devoted, skilled and less recognized, Problems activating as advisor in National without whom our advances would have been less Scientiic Foundation and in National Institute impressive”. of Health. In his Banquet Speech, in Stockholm, At the end of Nobel Lecture conference, with December 10, 1974, GE Palade declared that: “We title “Intracellular aspects of the process of protein feel that the new distinction acknowledges beyond secretion”, GE Palade marked the names of 16 us, beyond personal achievements the rebirth of our collaborators in this theme research naming them ield of knowledge and the vistas it has opened for “Good colleagues and comrades to whom I dedicate the medicine of tomorrow”. this work with affection and recognition for the And indeed, in further period, new knowledge accomplished work”. from Cellular and Molecular Biology lead to His collaborators, young in majority, came not identiication of numerous disease causes as, only from USA but also from many other countries, for example, those ones of mitochondrial and GE Palade having the conviction that “At their best lysosomial origin, which form today the object of the human minds belong to mankind, above nations Cellular Pathology. and above any frontier” and that “The science has an international character and do not develop in 2008, the last gong isolation”. In the last period of his life, GE Palade was In the numerous published articles, we often affected by a severe form of visual retina degene- remark that GE Palade put, as recognition sign, as ration, very probably as consequence of countless irst author his young collaborators. Many of them, hours of visual effort on the small luminescent at the specialization period end had developed screen of the electron microscope. in their countries Cellular Biology Centers, thus GE Palade passed away in October 7, 2008, at becoming specialists of international notoriety. age of 96 years in his home from Del Mar California. Such, for example, it was the case of Philip His ashes, as he wanted, was bring in Romania by Siekevitz, James Jamieson, Yutaka Tashiro, Tsuneo Philip and Georgia (son and daughter) and was Omura, Nicolae and Maya Simionescu etc. let go in the wind from “Vârful cu Dor” (“Nostalgia James Jamieson, past student of GE Palade, Peak”) – Meridional Carpathians. became professor in Cellular Biology Department From his family, Marilyn Farquhar Palade of Yale University. He organized the “GE Palade EM remained in San Diego, his daughter Georgia Slide Collection” where are kept and can be studied married Van Dussen lives in USA, as well the son the electron microscopic images made by GE Philip, professor of neurophysiology in Galverstone Palade during 1948-1990, images which are free University, Texas USA. Also, he had three nephews: accessible in entire world via Internet. the physician and bacteriologist Anca Israil and In case of Nicolae and Maya Simionescu, the surgeon Radu Palade, from his sister Adriana the intersession of GE Palade at Romanian state marriage with the jurist Mihai Dumitri Israil, and authorities and his contribution with equipments Irina Alexandra Matiu, physician in Huston, Texas and funds lead to establish in 1973 in Bucharest USA, from his sister Constanţa marriage with the the Institute of Cellular Biology and Pathology, physician Ionescu Matiu. which had as manager until 1995 the professor Nicolae Simionescu, and after that his wife Maya. George Emil Palade and research Starting with 1978, a large number of medical ethics undergraduate students from this Institute were Viewing the life and work history of GE accepted as scholars for post-doctoral study in Palade, it keeps attention his respect for a certain laboratories of GE Palade from Yale University. ethics of research. A prime aspect is that we can As concerns the research managing, GE deduce when he received the Nobel Prize his irst Palade considered that “Researchers must develop appreciation words were “for the workings of free their ideas, but with the quality control of

Bulletin UASVM Animal Science and Biotechnologies 71(2) / 2014 Review –A life for Nobel Prize, remember George Emil Palade 83 projects and results”. He believed also that “It must and after Nobel Prize with cu Brown-Hasen 1983, be avoided the effort concentration to resolve a Henry Gray 1986 and president Ronald Reagan certain problem concentrating massive work forces conferred him the National Medal of Science in and resources. It needs accumulation of data and 1986. GE Palade was also invited in Pontiical facts to reach a critical mass of basis knowledge. Academy of Rome, where Pope Giovanni Paolo the In this moment, we can formulate an applied second gave him a necklace in appreciation. When research project and could be proposed a target. the Pope gave him the necklace, GE Palade said: But until then, it is only lost time, wasted resources “Holly Father, but I am orthodox, you must know!” and disappointed expectations”. But, GE Palade and the Pope replied “No matter dear, for how considered that “We must habituate with the short many things you do for humanity you get over any life of discoveries, because the ideas for eternity are denomination borders”. few”. He was also elected “Doctor Honoris Causa” by As concerns the scientiic discovery priority universities from USA (Yale, Chicago, Philadelphia, conferred as a rule to those one who is the irst Michigan, Ohio, New York and Columbia) and that publish his results, GE Palade considered that from abroad (Bern, Bristol, Siena, Uppsala). Since could be equally associated with “those one who 1961, GE Palade became member of “US National bring the most illustrative and solid arguments for Academy of Science”. the research theme”. Persons who known GE Palade, for example Early his student period he was inluenced Keith Porter, James Jamieson, Alan Tartakoff, by his professors, the anatomists David Sabatini and Yutaka Tashiro, Günter Blobel and Grigore T. Popa and by the biochemist and Ewald Weibel in different occasions have André Boivin, GE Palade considered in the spirit published articles about his role in development of functional structure conception that “To of Cellular Biology. The most ample biographical discover a certain structure is not a big thing if presentation was done by professor of Medicine you don’t succeed to make out also what function History Radu Iftimovici who, in June 1991 in La accomplish” as well that ”The signiicance of vital Jolla, met GE Palade and their long and repeated phenomenon can be understood only descending discussions were recorded and formed the content to molecular level, practicing a concrete, deepened of volumes in Romanian language “George Emil and multidisciplinary experiment”. Palade, the irst Romanian laureate of Nobel Prize” In connection with this concept, we can tell a published in 1993 in Bucharest Romania and story during a conference of GE Palade with the “Confession of a winner” published in 2006. subject “Structure and function of endoplasmic reticulum” in 1964 at Chemistry Institute from “There is no Prophet who is despised except in Cluj Romania. During the dissertation, when his country…” he used images with endoplasmic reticulum (Matthew 13:57) abundant in exocrine pancreas cells, he mentioned that yet couldn’t afirm which is the energy source for chemical effort of protein synthesis from As concerns the relations of GE Palade with polyribosome level. In the conference end, during Romanian oficials, it is known that before Nobel discussion time, a person from hall afirmed that Prize (1974) he was considered apostate and the energy source for protein synthesis is doubtless his name maintained in anonymity. His visits in inside mitochondria, which are in intimate contact Romania were limited only to meet his mother and with granular endoplasmic reticululm. The answer sisters. After 1975 the situation is changed, so GE of GE Palade was “We also have thought to that, but Palade was elected honoriic member of Romanian the fact must be demonstrated”. Academy, was decorated with Romania Star and in Finally, it is necessary to be mentioned that 2007 with Romania Star as Necklace Distinction, even before he became Nobel Prize laureate GE and in few city his name was gave to some colleges Palade was honored for his scientiic activity with and streets. He made speeches only in the Institute other important prizes as Warren Triennial 1962, of Cellular Biology and Pathology from Bucharest Passano 1964, Lasker 1966, Ducket Jones 1966, and in the Chemistry Institute from Cluj, but not in Louisa Gross Horowitz 1970 and Dickson 1971, one of the six Medicine Universities from Romania.

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The GE Palade destiny was not unique. Nu- 12. Motta PM (2001). George Emil Palade and Don Wayne merous other intellectuals of Romania elite Fawcett and the development of modern anatomy, histology and contemporary cell biology.Italian journal do not found in own country work conditions, of anatomy and [Archivio italiano di comparable with those from developed and anatomia ed embriologia] (Italy) 106 (2 Suppl 1): XXI– with old tradition countries, to be favorable XXXVIII. ISSN 1122-6714.PMID 11730003. for their vocation and competence validation, 13. Muscan Catinca (1982). Despre Această etică a cunoaşterii. so they emigrated in countries, which could Almanahul revistei Steaua, Cluj. provide these vital conditions. In this case, it was 14. NEWS ALERT: A murit Emil Palade realitatea.net, accesat 8 conirmed what the American biochemist Vincent octombrie 2008 du Vigneaud (Nobel Prize 1955) said: “Nothing 15. Palade George E. (2008) Autobiography. Nobelprize.org. 2008-10-07. Retrieved 2011-04-03. stop more the science progress than appropriate 16. Palade George Emil (1974). Banquet Speech. The Nobel idea at unseasonable moment”. It was the case of Foundation, Stockholm. musicians George Enescu, Constantin Silvestri 17. Palade George Emil (1974). Intracellular aspects of the and Sergiu Celibidache, the philosopher Emil process of protein secretion. Nobel Lecture, December 2, Cioran, the dramatist Eugen Ionesco, the sculptor 1974, Nobel Foundation, Stockholm. Constantin Brâncuşi, the historian Mircea Eliade, 18. Porter KR (1983). An informal tribute to George E. Palade. J. the engineer Henri Coandă and the writers Elie Cell Biol. () 97 (1): D3–7. ISSN 0021- Wiesel (Nobel Prize for Peace, 1986) and Herta 9525. PMID 6345553. Müller (Nobel Prize for Literature 2009). But even 19. Raju TN (1999). The Nobel chronicles. 1974: Albert Claude (1899-1983), George Emil Palade (b 1912), and Christian so, from abroad, always afirming their origin Réne de Duve (b 1917). Lancet (England) 354 (9185): without reserves, they had promoted the prestige 1219. doi:10.1016/S0140-6736(05)75433-7.ISSN 0140- of their motherland country… 6736. PMID 10513750. 20. Sabatini DD (1999). George E. Palade: charting the REFERENCES secretory pathway. Trends Cell Biol. 9(10): 413-417. 1. Banca Națională a României: Emisiune numismatică - 21. Simionescu N and Maya Simionescu (1988). The monedă din argint dedicată aniversării a 100 de ani de la cardiovascular system, in “Cell and Tissue Biology”. Ed. nașterea lui George Emil Palade Leon Weiss, Urban and Schwarzenberg, Baltimore. 2. Darnel J, Lodish H and Baltimore D (1986). Molecular Cell 22. Singer, Manfred V (2003). Legacy of a distinguished scientist: Biologz. W.H. Freeman and Crup, New York. George E. Palade. Pancreatology (Switzerland) 3 (6): 518–9. doi:10.1159/000076328. ISSN 1424-3903. PMID 14730177. 3. Doherty Judi (1980). Un rol pentru un laureat al Premiului Nobel. The New Haven Register. doi:10.1172/ 23. Tartakoff, AM (2002). George Emil Palade charismatic JCI37749 Obituary virtuoso of Cell Biology. Nat. Rev. Moll. Cell. Biol. 3(11): 871-876. 4. Farquhar MG, Wissig SL, Palade GE (1999). Glomerular permeability I. Ferritin transfer across the normal glomerular 24. Tashiro Y (1975). Accomplishment of Drs. Albert Calude and capillary wall. 1961. J. Am. Soc. Nephrol. (United George E. Palade and the birth of cell biology. Tanpakushitsu States) 10 (12): 2645–62. ISSN 1046-6673.PMID 10589706. Kakusan Koso (Japan) 20(1): 74–6. ISSN 0039- 9450. PMID 1094498. 5. Hăulică I (2002). Professor George Emil Palade at 90 years of age. Revista medico-chirurgicală a Societăţii de Medici 25. The Independent (2008). Prof. George Palade: Nobel şi Naturalişti din Iaşi, 107(2): 223-225. prize-winner whose work laid the foundations for modern molecular cell biology. 22 October 2008. Retrieved 2011-02- 6. Iftimovici Radu (1977). Creaţia românească în biologia 09. Archived universală, Ed. Albatros, Bucureşti. 26. The preceding Romanian translation of the quotation from Dr. 7. Iftimovici Radu (1994). Istoria medicinei. Ed. ALL. George E. Palade’s “Autobiography” is Copyright ©2008- Bucureşti. 2011 by Dr. I. C. Baianu, (in USA) 8. Iftimovici Radu (1993). George Emil Palade, primul 27. US NSF - The President’s National Medal of Science: român laureat al Premiului Nobel. Ed. Viitorul Românesc, Recipient Details Bucureşti. 28. Weibel ER, Palade GE. (1964). New cytoplasmic components 9. Jamieson DJ (1988). The exocrine pancreas, in “Cell in arterial endothelia. J. Cell. Biol. 1964, 23: 101-112).http:// and Tissue Biology”. Ed. Leon Weiss, Urban and www.jcb.org/cgi/content/abstract/23/1/101 Schwarzenberg, Baltimore. 29. http://wikipedia , the free encyclopedia: George Emil 10. Jamieson JD (2008). A tribute to George E. Palade. Palade, Biography. November 8, 2008 30. http://cushing.med.yale.edu/gsdl/cgi-bin/ 11. Jamieson JD (2009). George E. Palade EM Slide Collection, library?c=palade&p=about Henry Cushing and Hay Whitney Medical Library, Yale, USA. http://cushing.med.yale.edu/gsdl/collect/palade 31. http://library.medicine.yale.edu/

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32. http://nobelprize.org/nobel_prizes/medicine/laureates/1974/ 37. http://www.nobelprize.org/nobel_prizes/medicine/ palade-autobio.html laureates/1974/duve.html 33. http://nobelprize.org/nobel_prizes/medicine/laureates/1974/ 38. http://www.nobelprize.org/nobel_prizes/medicine/ palade-lecture.pdf The Nobel Prize Lecture of George E. laureates/1974/palade-facts.html Palade is in Text Format Pdf 3.78 MB, Copyright ©1974 by 39. http://www.nobelprize.org/nobel_prizes/medicine/ The Nobel Foundation, ISBN 981-02-0791-3 laureates/1974/palade-lecture.pdf 34. http://nobelprize.org/nobelfoundation/publications/lectures/ 40. http://www.nobelprize.org/nobel_prizes/medicine/ WSC/physio-71-80.html laureates/1974/palade.html 35. http://www.nobelprize.org/nobel_organizations/ 41. http://www.rockefeller.edu/nobel.html nobelfoundation/publications/lesprix.html 42. https://www-er.ucsd.edu/givetoucsd/secure/paymenttran/ 36. http://www.nobelprize.org/nobel_prizes/medicine/ onlinegiving.asp?sk=224 laureates/1974/claude.html

Bulletin UASVM Animal Science and Biotechnologies 71(2) / 2014