Lessons learned in Andrology: Yves Clermont, an interview by Lonnie D. Russell

Dr. Yves Clermont (Fig. 1) is famous for putting order into the life history of mammalian germ cells as they evolve from spermatogonia to spermatozoa. He accomplished this by defining specific cellular associations in cross-sections of seminiferous tubules referred to as stages of the cycle of the seminiferous epithelium in many species, including humans. He and Dr. C. P. Leblond were credited with the very first description of stem cells (Clermont & Leblond, 1953) and actually coining the term for the first time, cells that are today instrumental in many facets of modern day cell and molecular biology. Dr. Clermont has also touched on many different facets of germ cell differentiation (spermatogenesis), such as the flagellar cytoskeleton and Sertoli cell participation. He is also famous for his description of the germ cell Golgi apparatus in 3-dimensions and that of many other cell types in other organs and coined the term Golgi ribbon. He had an extraordinary love for teaching that reached thousands of undergraduate and medical, dental and graduate students and was awarded the prestigious William Osler Teaching Award. As a skilled artist (Fig. 2), he rendered blackboard drawings during his histology lectures, which were indeed spectacular. His style and passion for his work were complemented by a remarkable sense of good humor. He was born on August 14, 1926 and passed away on October 10, 2014. Lonnie Russell, who had completed a postdoctoral fellowship under the supervision of Dr. Clermont from 1973 to 1975, interviewed his mentor shortly after a symposium held in his honor at the Spermatology Symposium held in Montréal, Québec, Canada in September of 1998. Dr. Russell passed away unexpectantly in 2001 and the original tape recording was discovered in his office. The following is an edited transcript of this interview. The recorded interview will be made available on the McGill University website. I was a student at the University of Montreal in biology. I was having a general training in biology and taking courses in a wide range of disciplines such as bacteriology, biochemistry, and zoology. At the time, the number of openings in graduate studies was minimal, and they were in areas that didn't excite me too much. While I was completing my BSc in biology, I was concerned about what I would do afterward. I was asked by one of the professors in biology at the University of Montreal to go and work on a summer project. It happened that Leblond was involved in that project. The goal of the project was to try to explain whether salmon molting was caused by the influence of the thyroid gland. So, I went to work in the Laurentians (mountains north of Montreal) the year before I graduated. The only job I was doing was to take care of the fish, to clean the basins and to inject the fish with thyroid hormone. So, it was a very mundane type of activity. The person in charge of the project worked directly under the supervision of Leblond. So I didn't have direct contact with Leblond. But at the end of the summer, the end of August 1948, he said in passing, casually, that ‘if you are interested to do graduate studies in histology you may come to see me.’ That was it. After that it was absolutely evident to me that I had no choice. I had to come to work with him at McGill. That was appealing to me, I had done a little bit of histology at the university and had found it absolutely dull, presumably because the teacher was not stimulating. But, I don't know why, I liked to look through the microscope at organisms of all kinds, including histological sections. It rang a bell so to speak and I said now I know what I will do next. That was summer 1948. Then, September arrived, I completed my BSc degree in May 1949. Instead of planning on a summer vacation, I decided to go see Dr. Leblond. At the time, I didn't know a word of English. I knew how to read English, but had never had an opportunity to speak it. I remember that I came to the Anatomy Department at McGill University and I had to write my introductory sentence. ‘May I see Dr. Leblond, please?’ because the secretaries were only English speaking. He accepted me, but contrary with what he said at the meeting the previous summer, he wasn't sure of keeping me as a graduate student. So, he took me because I was coming from the University of Montreal and he wasn't sure of my background. He said, ‘I will take you conditionally.’ I still have the letters from the graduate faculty saying that I was accepted – first, if I had my degree and second, if I was satisfying the requirements of Dr. Leblond. As soon as I graduated in May, I came to work - I didn't wait until September. He very quickly showed me these slides that he had on periodic acid Schiff stained sections showing carbohydrates. So, I worked during that summer, and it is then that I worked out the classification of steps of spermiogenesis and of the stages of the cycle of spermatogenesis. When Leblond saw that, he felt that I was probably good enough to pursue my graduate studies and changed my status from conditional to permanent. Then, things were moving very well and very quickly. We were accumulating observations of all kinds. He felt that I should not spend time writing a Master's thesis, which was at the time a requirement. I moved straight to the PhD. In the meantime, instead of writing a thesis, we wrote our first article dated 1952, but we took 2 years to write it. So the time that we spent writing the first manuscript was enormous. But, the final product was there (Leblond & Clermont, 1952a). Because I went straight for the PhD, I finished it in just 4 years, in 1953. Oh yes, before I finished my PhD I had five or six publications (Leblond & Clermont, 1952a,b; Clermont & Leblond, 1953; Clermont, 1954; Clermont & Benoit, 1955; Clermont & Haguenau, 1955; Clermont et al., 1955a,b). One written in French on the hamster (Clermont, 1954). We worked well together all along. He was very demanding, because he was telling us all the time that ‘perfection is the enemy of the good’. But, he was, himself, a real perfectionist in writing these papers. Most of his publications are very well thought through – every line, every word is perfectly in place. This is something that I learned from him. You have to realize that this classification of spermatogenesis that we devised was performedvery simply for one reason. It was to find a tool, an instrument to investigate the renewal of the spermatogonial population. I thought of this classification after what, a maximum of 6 months of work or less. This orientation to study the spermatogonial population encouraged me to go in that direction - that was my first objective. It was a very exciting period because I could see data accumulating on the spermatogonial population at various stages of the cycle. An exciting story that was far more precise and accurate than those that had been circulating at the time, We had no idea of the duration of the process at the beginning, but we had some landmarks. From quantitating spermatogonial population we could see, for example, exactly when the spermatogonia were dividing, and whether the divisions were synchronized or at random. We clarified that right from the beginning. Then we injected colchicine to block cells in mitosis and we immediately saw these peaks of mitosis. That was quite exciting. My thesis was directed at understanding the behavior of the spermatogonial population in the rat, in the monkey and in one or two other species. The staging of spermatogenesis was a fallout of something I needed for my thesis; it has been useful for my own work and for the work of others. The idea of the existence of the cycle, was well known by everybody who was working on the testis; it had been well-described by Regaud at the beginning of the century (Regaud, 1901), in particular, and many others. My objective was to find a reproducible method of identifying the stages of the cycle of spermatogenesis - that is all. Interestingly enough, this classification has become better known that the renewal of spermatogonia. Having worked on whole-mount of seminiferous tubules with Bustos-Obregon (Clermont & Bustos-Obregon, 1968), having accumulated a mass of quantitative data with Louis Hermo (Clermont & Hermo, 1975), Martin Dym (Dym & Clermont, 1970) and other students, and using radioautography, I developed the conviction that there are two classes of stem cells - and this has not changed. Whether others agree with that, I now couldn't care less. I still believe that what we have done was well-documented, it was evident, it does not mean that it will not change, but if you ask my opinion now, I am as convinced about the various modalities of cell proliferation and renewal than I was 25 years ago. Now, understand that I have stopped working on these things. You cannot spend all your life on one thing, and I was not trained to use other techniques such as the isolation of spermatogonia. I did not have the tools or the expertise to use these methods. But the evidence I had, indicated to me that I was on the right track. Maybe I am prejudiced, but I was never convinced that my opponents, and I have at least two or three, were right. Because, it was not evident what they were telling me. It never overcame the quality of the mapping and what this mapping was saying in normal or in irradiated animals. I still have tons of data that I have never published – I did not have time – on X-irradiation on the spermatogonial population. I still have that at home, and it simply confirms our views, my views and the views of my colleagues. So you see, this is my attitude. Now, if you tell me that in certain conditions, the spermatogonial renewal takes place differently, if they have the facts to support this, I will have to accept it. Don't forget that my career is practically over and I don't care if somebody shows something different – on the contrary, I would be happy to see somebody move ahead with some solid facts. For 10 years, I was practically alone working on these topics, so I was not prejudiced one way or the other. And I was alone with my students or colleagues. I had accumulated a mass of facts that was supporting my theory; no, it is not a theory, it is a suggestion, So that is my attitude, even though I was not sure that my ‘suggestion’ was correct, when I was seeing other studies, I wanted to be absolutely convinced that they had the facts to support their view. Yes, of course. I think this is very interesting. It is very positive too, because as soon as I finished my PhD, I stayed here for a year and wrote a few articles on the renewal of spermatogonia, the very first one on that topic. I started to work on the wave of the seminiferous epithelium. I did all sorts of studies on the development of the testis, with Bernard Perey, this work I did as a graduate student and published only later. There were so many interesting things. I was doing pretty much all of this by myself. Then I went for a year of post-doctoral training and in those days post-doctoral training was very nice, especially when you were going to Europe. I enjoyed my year tremendously, but I used it to learn the techniques of electron microscopy. That was in 1954–55. The first Porter-Blum microtomes were developed, the first RCA microscopes were being produced, so it was a field that was opening up. I went in the lab at the Cancer Institute in Villejuif (Paris, France), and worked with Wilhelm Bernhard and learned the methodology of electron microscopy. Then I came back to the Department and in my mind, I was not thinking of going anywhere else, because when I left for my post-doc, Dr. Leblond told me ‘If you want to come back, you may come back.’ I said, ‘I am coming back.’ So it is nothing complicated. And this is the interesting thing. I continued to work, more or less in contact with Dr. Leblond. We published a number of articles on the spermatogonial population of the monkey. We worked on the isolation of the carbohydrates from the acrosome of the guinea pig – we did all sorts of things, quite interesting. There is a point that should be underlined, which is exceptional in a sense. Considering that a man of the caliber of Dr. Leblond, a very strong, dynamic person, there were two possibilities for our relationship. Either he gave me my full independence, or I would go under his thumb and therefore, would not have stayed here very long. Dr. Leblond in the most elegant way, told me ‘now, you are on your own.’ … That was quite interesting, because, as you know in the European style the ‘patron is the patron’ and for life. But, from then on he was always interested in what I was doing; he was always making suggestions and corrections of the manuscripts, but never imposed himself on my research orientation. This is really close to a miracle, when you consider the character of the man. I should like to mention that my relationship with Dr. Leblond is exceptional…. He would have liked to have considered me as a son. But no I said ‘no, I am not your son, I am your friend,’ which is a different story. And we continued, in this manner throughout these 50 years. Most memorable student; well I never thought along these lines. There are some who I appreciated because they were very efficient, original, but I appreciated practically everybody that worked with me. There are some who gave me a lot of headaches at times. I had a lot of experiences, imagine, I probably had 30 students and some I had to tell them that they had to leave. So that was difficult. And they had to leave, why? Because, I was convinced that I could not train them properly. They could train themselves, so after their Master's, I was asking them to leave. In general, I always had good relations with my students. Some of them were not easy, but this is life. If you were to ask me who was the most interesting collaborator, I would tell you immediately who that is: Dr. Rambourg - and this for his vast variety of qualities. This fellow is exceptional, very peculiar character, but for some reason we worked together in a most efficient manner. This fellow works by himself, in France. He is a person who has a vast spectrum of interests, and not only in science. What a character. He used to come here twice a year for 3 weeks and we would work together. We were fighting each other like dogs at times, but we always managed to work well, very well together, in fact,. For a period of 25 years, we published possibly something like 30–35 articles together – always difficult to write. He is a fascinating person. He has interest in languages, he has interest in philosophy, he has interest in music. It has been a very enriching experience each time he came here. You know that I am retired, and he will be retiring soon. We will keep in touch, but it means that the relationship will change. So, I am helping him, he is helping me. He is really a genius, but not easy, you know. I don't know how, but I manage to work well with him. We were complementary to each other. I was SLOW, he was FAST.

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