n Memoriam

Tributes published in the Journal to honor Andrologists ho have

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Journal of Andrology, Vol. 24, No. 4, July/August 2003Copyright q American Society of Andrology

Memorial

On December 19, 2002, the American Society of An-drology and the Polish Andrology Society lost one oftheir ardent proponents and advocates at the age of 56.Grzegorz A. Szymczynski, MD, was for many years anactive member of ASA and played a pivotal role in es-tablishing the Andrology Society in his native country.Being trained as a gynecologist/urologist, he was devotedto popularizing andrology among the urologists, gyne-cologists, psychiatrists, and other medical specialists.Despite serious shortages in financial support, and due

to sheer perseverance, Gzegorz succeeded to initiate pub-lication of Progress in Andrology, a widely respectedjournal in Europe. At the time of his sudden and untimelydeath, volume 4 of the journal was coming off the press.It contained the proceedings of an international confer-ence, Andrology Today, which he organized in Byd-

goszcz, Poland, in June 2000. When we participated inthis conference, little did we realize that this would bethe last time we would see Grzegorz alive and bubblingwith enthusiasm.Gzregorz was born and educated in Poland; however,

his interests in andrology and his insatiable quest forknowledge took him well beyond the borders of his nativecountry. He spent a considerable amount of time in theUnited States working with such esteemed andrologistsas Sherman Silber, Arnold Belker, and Emil Steinberger.Many will cherish Gzregorz’s dedication and numerous

contributions to the field of andrology. Those who wereprivileged to know him personally will miss his enthu-siasm, loyalty, and good-natured humor. Andrology willmiss an ardent supporter.

Emil and Anna Steinberger

Journal of Andrology, Vol. 26, No. 6, November/December 2005Copyright ! American Society of Andrology

Memorial

Geoff Malcolm Hasting Waites (1928–2005)

Geoff Waites’ premature death robs the Andrology com-munity of an exceptional leader. Geoff was a kindly, wisepatron, a valued colleague, and loved friend of so manyof us. At 76, he had a full life as a scientist and manager,yet his restless energies never slowed and he still hadenough ideas, enthusiasms, and projects to fill anotherdecade. So, we are greatly diminished by his loss.Geoff completed his PhD 50 years ago. His research

produced fundamental concepts, such as the spermaticcord as a countercurrent heat exchanger maintaining tes-ticular temperature, the blood-testis barrier, the role of theepididymis in maturation of sperm function; and he dis-covered the first prototype posttesticular male contracep-tive. As a past master, Geoff deeply respected good sci-ence but even more loved the hard-working scientists whocreated it. Above all, he reveled in the opportunity toharness the best science for enhancing people’s lives.During his 11 years as Manager of WHO’s Male TaskForce, with Ebo Nieschlag as his Chairman, he directedits golden age securing proof of principle for hormonalmale contraception—a monumental undertaking—as wellas organizing multicenter clinical trials in China and In-donesia, creating the WHO Semen Manual, and runningpractical Andrology Workshops all over the world. Hislast paper was a wide-ranging history of the WHO MaleTask Force. When male contraception comes to its over-due fruition, it will owe a great debt to his quiet persis-tence and effective guidance. Kind and gentle in all hisdealings, Geoff was nevertheless impatient for progress.To the world of science, he brought organizational insight,while to the world of scientific project management, hebrought the intelligence and creativity of an outstandingscientist. Both were immeasurably enriched and in bothhe placed people at the heart of matters. Whatever he did,measuring his achievement by what went before and after,he was truly great.Geoff had a life-long commitment to his students and

colleagues, always staying in touch and being proud oftheir progress. He understood the value of tact and timelyencouragement, virtues he practiced rather than preached.He led by example, with faith in the soft power of per-suasion. Above all, Geoff’s scientific and personal integ-rity and dignified humility shone like a beacon, infusedwith wit and wisdom. Now we will miss him, his wryand priceless humor, his love of puncturing pomposity,

DOI: 10.2164/jandrol.05150

and his valuing of fairness and progress. Never again willwe get to see that wonderful sight of an angular figure,in a crumpled camel-colored suit, loping toward you witha cheeky grin that barely held back some funny story,eager to seize you by the arm for his latest enthusiasm,and to sweep you away with his love of life, his joy inscience, and his abiding mission to leave the world a bet-ter place, especially for those whose lot in life was unfair,unjust or just plain unlucky.Geoff Waites was a pioneer of Andrology and com-

mitted internationalist. He traveled a lot, not just becausehe delighted in meeting different people and unfamiliarcultures, but because he loved bringing Andrology ex-pertise to all corners of the globe. Although shunning thetributes, he would have liked that the ISA honors himwith a talk on the science of Andrology in a global frame-work, for that is his legacy.

690 Journal of Andrology · November/December 2005

Personal RemembrancesFrom Trevor G. Cooper and Ching-Hei YeungChing-Hei and I met Geoff at Reading University, Eng-land, where he was our PhD supervisor.Trained as a physiologist, Geoff’s early work was on

nerves and vasomotor responses, and he retained this in-terest in vascular physiology in his future research on themale reproductive tract. Working in Australia with theproblems of summer infertility in rams, he showed theimportance of apocrine sweat glands of the scrotum (aNature paper) and that countercurrent heat exchange inthe spermatic cord acts to cool incoming arterial blood tothe testis. He followed this up by demonstrating that thedamaging effects of scrotal heating on semen quality weredue to unequal changes in testicular blood flow and me-tabolism. His work with Brian Setchell reflected theircommon interest in measurement of blood flow and vas-cular permeability, and they were the first to demonstratesignificant variations in different regions of the epididy-mis (another Nature paper) before the morphological dif-ferences in endothelia were known. His expertise at can-nulation led to most important discoveries on the com-position of intratubular testicular fluid and the study ofimmature spermatozoa just shed from the germinal epi-thelium (yet another Nature paper) first in rams and sub-sequently in rats and monkeys by Geoff and in other spe-cies by others. This ability to collect, in this way, uncon-taminated samples of testicular spermatozoa, sometimesover days from a conscious animal, enabled the nature ofsperm maturation to be defined by comparing these sper-matozoa metabolically and biochemically with those fromthe cauda epididymidis. Studies on the effects of heat ontesticular fluid and sperm production and metabolism fol-lowed in a series of classic papers. When combined withcannulation of testicular lymphatics, the way was open todemonstrate that a blood-testis barrier existed limiting theingress of circulating compounds, again initially shownin rams but subsequently demonstrated in rats (my(T.G.C.) PhD thesis) and later in other species. Ultrastruc-tural techniques were soon employed in other labs to lo-cate the anatomical sites of the barrier. Testicular fluidproved to be similar in many species but far differentfrom fluid leaving the epididymis, and the consequencesof this for the maturing spermatozoa are still being stud-ied. One aspect, osmolality, is our current research inter-est.Geoff’s lab was a crossroads for international scientists,

many from the Indian subcontinent, who worked on re-search on male contraceptives (Gossypol, Trypterigiumwilfordii). The action of testicular toxicants fired Geoff’sinterest in the entry of these compounds into the testisnow that both testicular fluid and sperm could be col-

lected. The inhibition of sperm glycolysis by alpha-chlo-rohydrin prompted the synthesis of other chlorinated com-pounds at Reading, which were tested for their action onsperm metabolism and male fertility. The requirement forpotential posttesticular contraceptives to access epididy-mal spermatozoa brought Ching-Hei to Geoff’s lab fromHong Kong, where she had developed, with PatrickWong, an epididymal cannulation procedure. Ching-Hei’sPhD thesis and my postdoctoral work with Geoff utilizedthis technique to characterize the blood-epididymis barrierin rats. Geoff’s introducing us to ‘‘functional sterility’’proved an important concept for us both, as it has con-sumed a major part of our research lives since leaving hislab.When asked to mention Geoff’s role as a teacher, both

Ching-Hei and I had to stop and think about it. The dif-ficulty lay in his teaching method: knowledge was im-parted subliminally, not imposed, as we sat together atthe bench pulling catheters and cannulating any orificeavailable. There was always a cheerful rapport in the lab,and we both remember carefree days lived in true ca-maraderie, delighting in the same ridiculous sense of hu-mor. Much admiration has been aired about Geoff’s in-tegrity, wisdom, and sense of humor, but what we remem-ber him for mostly is his warmth and kindness, his will-ingness to listen and encourage, and his readiness to helpyoung scientists, particularly those in developing coun-tries. Outside academia, this interest was extended to chil-dren from the developing world, and nowhere was thismore evident than in his work with Concern Ethiopia andthe building of schools (the first bearing his wife Doreen’sname) in that country.We shall miss him a lot because he was more than a

scientific adviser, but also a close personal friend, inwhose lab we met, and in whose home we could relaxwhen times got tough.

From Brian P. SetchellThe death of Geoffrey Waites in May 2005 has deprivedthe world of one of a most able and experienced scientificandrologist. Geoffrey did his first degree, a BSc, at Bir-mingham, then did a Part II in Physiology at Cambridge.As he had a Cambridge degree, he was permitted to dohis Cambridge PhD at the Institute of Animal Physiologyat Babraham just outside Cambridge, with the first Direc-tor Ivan de Burgh Daly, on the nerve supply to the heartof the sheep. Geoffrey came to Australia in 1958 to theCSIRO Division of Animal Physiology, Prospect, a sub-urb of Sydney, to work with George Moule, a veterinarianwho had noticed that rams in Queensland became infertileduring the summer heat. Geoff and George collaboratedfor several years and published several papers together,two of which (Waites and Moule, 1960, JRF 1, 223; and1961, JRF 2, 213) were important in establishing that the

691Memorial

spermatic cord in the ram acted as a countercurrent heatexchanger and also eliminated the pulse from the arterialblood. I did my PhD at the Veterinary School at Cam-bridge at about the same time as Geoff, and although Ivisited Babraham several times, as far as we could deter-mine, Geoff and I never actually met during that time.When I returned to Australia at the end of 1957, a

mutual friend from Babraham, Alan Pierce, later Chief ofCSIRO Animal Health, suggested that we meet whenGeoff came to Sydney. Our long-lasting collaboration onthe physiology of the testis and epididymis began undersomewhat unusual circumstances. At the time we met inSydney, I was interested in the possibility that the sheephad a different catecholamine response to hypoglycemia,which might have influenced their clinical response.When I read a paper that claimed that the temperature ofthe denervated ears of rabbits could be used as a bioassaysystem for adrenaline, and as I found that Geoff had de-scribed a surgical method for removing the superior cer-vical ganglion of sheep (1957, J Physiol 135, 52), westarted a project on the catecholamine response of sheepto hypoglycemia, the results of which were written up inour first joint paper (1962, J Physiol 164, 200).In this study, we also denervated the hearts of two

sheep, again using Geoff’s knowledge of the relevantanatomy obtained during his PhD (1957, J Physiol 139,417, and 135, 58). During this time, we discovered thatwe shared an interest in cricket, and arranged severalmatches between the Prospect team and a team from thelaboratory where I was based, which included a NSWstate player and his younger brother, who later played forAustralia. Geoff was a very fine batsman and also a goodfast bowler, but he had some uneasy moments facing thisvery tall 17 year old, who, a few years later, was bowlingfor Australia in Test matches.Then in 1962, I applied for and got a research job at

CSIRO Prospect and Geoff and I began a collaborationto develop techniques for measuring blood flow and me-tabolism in the testis and epididymis (1964, J Physiol171, 411; 1964, Nature 203, 317). As well as demonstrat-ing important regional variation in the epididymis, wefound interesting effects in the testis, during changes intemperature (1964, JRF 8, 339) and nutritional status(1965 JRF 9, 149). During these blood-flow studies, wenoticed that the values obtained with two different indi-cators (iodoantipyrine and rubidium) were quite differentin the testis and brain, but similar in all other tissues weexamined. This observation led to the concept of theblood-testis barrier, analogous to the blood-brain barrier,which was further developed when Geoff and I, with SeppVoglmayr, established a technique for chronic collectionof fluid from the rete testis of rams (1966, Nature 210,861) and found that this fluid was quite different in com-

position from blood plasma or testicular lymph (1967,JRF 14, 87).The situation at Prospect when I joined the staff there

and for the first few years was a scientific paradise. Fa-cilities were excellent and money seemed to be no prob-lem. In fact, Geoff and I were invited to apply for thefunds for an automatic gamma counter by a representativeof the Ford Foundation, so that we could avoid sitting upto 2 or 3 AM counting the isotope levels in pieces of testisand other tissues, using a manual counter, but the Chiefof the Division refused us permission to do so, sayingthat if the project was worth doing, CSIRO would buythe necessary equipment, and they did.In 1965, Geoff had the opportunity to work at Jouy-

en-Josas near Paris with Robert Ortavant, which devel-oped his love of France and things French, and when hereturned to Australia, he was appointed Associate Profes-sor of Physiology at Sydney University. At Geoff’s sug-gestion, I spent 1967 at Babraham working with Jim Lin-zell, who had been a close friend of his during his Ba-braham days. My collaboration with Geoff in Sydneycontinued until 1969, when both of us independently, andwithout the other knowing, accepted positions in the Unit-ed Kingdom, Geoff as Professor at Reading and me as aResearch Officer at Babraham. We left Sydney withindays of one another, Geoff and his family going east andus going west, by boat. We continued our collaborationin the United Kingdom for some years, some experimen-tal, but mostly writing together, and one of Geoff’s PhDstudents (Steven Main) worked with me for several yearsat Babraham as a post doc funded by NIH.Geoff was a very fine sportsman and a wonderful ra-

conteur. His stories were mostly based on his own ex-periences. For example, he told a story of a young re-search student, when the current technique for measuringblood pressure involved a fluid catheter connected to areservoir topped by a fine rubber diaphragm. The bestrubber for this purpose was the rubber from a condomand, on one occasion, the young, rather naive student wassent by a senior colleague to purchase a condom. Whenasked whether he wanted plain or teat-ended, he repliedthat it did not matter as they cut off the end before theyused them, somewhat to the surprise of the assistant inthe pharmacy.The other story I remember well involved two cleaners

at Prospect, who came in after office hours, by which timemost of the scientific staff had usually gone home. Geoffwas sitting at his desk behind the door on this occasionwriting up the results of the day’s long experiment, whichhad produced a certain amount of mess in his lab. As thetwo cleaners came level with his open door, he heard oneof them exclaim ‘‘Oh Jesus Christ.’’ The other said:‘‘Shussh, he’s behind the door,’’ which Geoff claimedgave him divine status.

692 Journal of Andrology · November/December 2005

When he was at Jouy, he was invited to join CharlesThibaut, the then director, for lunch with a rather crassAustralian who was visiting the lab and who shall remainnameless. They were served a lovely bottle of Loire Rosewine. Thibaut had perfect English, but when the Austra-lian visitor asked if we in Australia made rose wine bymixing equal parts of red and white wine, all Thibautcould say was ‘‘jamais’’ (never), the only time Geoff hadknown Thibaut’s English ever to desert him.I had less contact with Geoff during his later work at

WHO, but he certainly made, during that time, an im-portant contribution to international andrology. His con-nections with China helped bring andrology in that coun-try into the mainstream of the discipline, and he fulfilledan important role as Chairman of the Advisory Board ofthe Asian Journal of Andrology up to the time of hisdeath.Geoff was the best colleague and most entertaining

companion I have ever known, and I will miss his friend-ship more than I can ever say.

In Memoriam: Dr Charles Philippe Leblond, MD, PhD,1910–2007

On April 10, 2007, Dr Charles Philippe Leblond passedaway peacefully at his residence in Montreal surroundedby his family. Dr Leblond was the most prominentmember of the Department of Anatomy and CellBiology of McGill University.

He began his scientific career in France studying thedistribution of vitamin C in various tissues of laboratoryanimals. He also learned to use newly discoveredradioactive isotopes, in particular radioiodine, whichhe found concentrated in the thyroid gland. After a briefstay in Canada in 1941, he went back to France toreturn to the Department of Anatomy of McGillUniversity in 1946. In collaboration with Dr Bélanger,he developed the technique of radioautography, whichpermits the exact localization of radioactive molecules intissues and cells. Radioautography was extensively usedby Dr Leblond in his laboratory at McGill and byresearchers around the world to investigate and clarifya variety of biological processes. This procedurecontinues to be used today by molecular biologists todetect RNA molecules in situ and to study thelocalization of genes and DNA sequences.

Dr Leblond used radioautography to introduceradioactive precursors of DNA and thus to study therenewal and fate of cells of several basic tissues. Hedemonstrated for the first time that most cells andtissues in the adult body undergo continued renewal.Using mathematical models and modern methods ofquantitation, Dr Leblond and his colleagues estimatedwith remarkable accuracy the turnover and mitotic ratesof numerous cell types. He and his colleagues madefascinating discoveries that resulted in the introductionof ‘‘time dimension’’ to cells and tissues, opening thedoors to the understanding of the cell cycle and to theidentification of stem cells. In fact, Dr Leblond and hiscolleagues devised several methods for isotopic labelingof the nuclei of cells in the process of division. Thelabeled cells could then be identified for as long as theysurvived in the living tissues. Exploiting this approach,Dr Leblond and his colleagues made the first measure-ments of cell renewal in most tissues of the body. Hefound that the rate of renewal was remarkably high in

the epithelial lining of the stomach and intestine, wherethe cells completed their life cycle within a few days.

In collaboration with Dr Yves Clermont, he alsostudied the complex changes that occur in the Golgiapparatus and acrosomic system during the transfor-mation of spermatids into spermatozoa in the rat,mouse, hamster, and guinea pig. These changes wereused to divide the cycle of the seminiferous epithelium ofthe rat into 14 stages whose duration was later estimatedto be 12 days for the whole cycle. This classificationserved as a classic model for the study of the cycle of theseminiferous epithelium in other mammalian species,including the human. The 2 original papers by Leblondand Clermont (Leblond CP, Clermont Y. Spermiogen-esis of rat, mouse, hamster and guinea pig as revealed bythe ‘‘periodic acid-fuchsin sulphurous acid technique.’’Am J Anat. 1952;90:167–216; Leblond CP, Clermont Y.Definition of the stages of the cycle of the seminiferous

Received for publication April 27, 2007; accepted for publicationApril 27, 2007.

DOI: 10.2164/jandrol.107.003202

Journal of Andrology, Vol. 28, No. 5, September/October 2007Copyright E American Society of Andrology

epithelium of the rat. Ann N Y Acad Sci. 1952;55:548–573) constitute milestones in andrology, and perhapsthey are the most frequently cited papers in the field ofmale reproduction.

During the mid 1960s, Dr Leblond and his colleaguesachieved the refinement of radioautography and itsapplication to electron microscopy to exploit the highresolution of this technique. Dr Leblond was able toanalyze intracellular pathways followed by radiolabeledamino acids and carbohydrates. Again, he was one ofthe first to identify the role of different compartmentsand subcompartments of the cell involved in thebiosynthesis and secretion of glycoproteins.

Throughout his illustrious career, Dr Leblond initi-ated numerous other projects involving immunocyto-chemistry both at the light and electron microscopelevels, which led to elegant results and importantdiscoveries. These contributions resulted in the publica-tion of 430 scientific papers, many of them stillfrequently cited. In addition to his passion as a re-searcher, Dr Leblond served as the head of theDepartment of Anatomy of McGill University for morethan 25 years. During that time he developed one of thebest international research centers in cell biology, and hewas the mentor of many prominent colleagues.

Throughout his long and distinguished career, DrLeblond’s contributions to modern medical science havebeen outstanding and recognized by numerous distinc-

tions and awards. Dr Leblond was a Fellow from theRoyal Society of London, the Royal Society of Canada,and the American Academy of Arts and Sciences. Hereceived honorary doctorates from Acadia University(1972), McGill University (1982), l’Université de Mon-treal (1985), York University (1986), and l’Université deSherbrooke (1988). He was also an officer of the Orderof Canada and Officier de L’Ordre National du Québec.

Dr Leblond was a distinguished colleague in everysense of the word. He was one of those individuals whocould not pass unnoticed, becoming an emblematic iconfor McGill University. Visitors and colleagues fromaround the globe always asked if he was still coming tothe department and if they could pay a visit to him.Until recently, before his health started to decline, hewas attending the departmental seminar series and oftenasked insightful questions. Dr Leblond was a person ofgenuine finesse and originality and a role model formany graduate students. Dr Leblond has touched thelife of many colleagues, former students, and post-doctoral fellows and will be forever remembered withgratitude, admiration, and respect.

Carlos R. Morales, DVM, PhDDepartment of Anatomy and Cell Biology

McGill UniversityMontreal, Quebec

Canada

Morales N In Memoriam: Dr Charles Philippe Leblond, MD, PhD 637

In Memoriam: Matthew P. Hardy, PhD, 1957–2007

With great warmth and profound sadness we rememberDr Matthew P. Hardy, who died unexpectedly onSunday, November 4, after completing the New YorkMarathon. In Matt’s quiet, unassuming way, he hadbecome one of the preeminent reproductive biologists inthe country, one who deservedly enjoyed a national andinternational reputation. He was also a longtime andloyal member of the American Society of Andrology,where he contributed mightily of his time and talent.

Matt received his PhD in Biology at the University ofVirginia in 1985. He then went on to conduct post-doctoral work with Larry Ewing at Hopkins’ School ofHygiene and Public Health, from 1985 to 1990. WhenMatt completed his postdoctoral, he left Baltimore forNew York City for the position of Staff Scientist at thePopulation Council. He was promoted rapidly through

the ranks at the Population Council, becoming a Scien-tist (equivalent to Associate Professor) in 1997, and thenSenior Scientist (equivalent to Professor) about 3 yearslater. He also held joint appointments at The Rock-efeller University and at Weill Cornell Medical College.

Matt became internationally recognized for his in-novative, groundbreaking studies on the origin anddevelopment of Leydig cells in the mammalian testis. Ina series of what have become classic papers, he conductedanalyses of the development of the adult population ofLeydig cells from Leydig cell precursors, demonstratingthe sequence of events that occur to form the adultpopulation, and the mechanisms by which the sequence isregulated. Along the way, he made the counterintuitiveobservation that androgen itself is involved in thedifferentiation of Leydig cell precursors, a critical andnow well-accepted observation. He also elucidated indetail the role of lutenizing hormone (LH) in both theshort-term and long-term regulation of Leydig cells. Overtime, the studies involved morphological, physiological,biochemical, and molecular methodologies, a hallmark ofMatt’s scientific style; he always did whatever wasnecessary to answer questions, not restricting himself towhat was comfortable.

While continuing to work on Leydig cell development,and most recently on Leydig stem cells, Matt studieddevelopmental changes in LH and androgen receptorexpression, the differential expression and regulation ofsteroidogenic enzymes, Leydig cell proliferation and itsregulation, the role of paracrine factors (primarily growthfactors) in Leydig function, aging and stress effects onLeydig cells, mechanisms through which reproductivetoxicants impact testicular function and puberty, and therole of neonatal hypothyroidism on Leydig cell numbersand function. In all, Matt authored over 70 peer-reviewedpublications, coauthored a very well-known and highlycited book (The Leydig Cell), and published about 35chapters. His creativity and high level of productivitycontributed to his being named the 2000 YoungAndrologist by the American Society of Andrology.

Matt exhibited a unique ability to understand theclinical implications of many of his observations andplayed a pivotal role in the development of futureclinician-scientists. He routinely contributed to medicalstudies at Cornell, documenting that even men withsmall testes maintained their quantitative complementof Leydig cells, compared with normal men.

Photo credit: Karen Tweedy-Holmes, Population Council.

DOI: 10.2164/jandrol.107.004564

Journal of Andrology, Vol. 29, No. 1, January/February 2008Copyright E American Society of Andrology

Matt lived his belief in contributing to science in itsbroadest sense, including service to scientific societies. Hechaired ASA’s Bylaws Committee (1995–1997), waselected to the ASA Executive Council (1998–2001), andcochaired the Membership Committee (2001–2002). Hisquiet, capable leadership as Chair of the Testis WorkshopExecutive Committee (2005–2007) has facilitated therewarding relationship between the Testis Workshop andASA. Having served on our editorial board (1998–2001),Matt was selected to become Co-Editor-In-Chief of theJournal of Andrology (2002–2007). Together with PeterSchlegel, he significantly enhanced the reputation andscope of our journal. Throughout all these activities, hebrought a clarity of vision and a sense of ethics rarely seenin our community. It was fitting, then, that he was to berecognized with the ASA Distinguished Service Award atthe upcoming 2008 ASA Annual Meeting.

Matt’s service activities extended beyond the ASA.He was Chair of the Bylaws Committee of SSR (1998–2000) and a member of the editorial boards of Archivesof Andrology, Biology of Reproduction, Endocrinology,and Molecular and Cellular Endocrinology. At the timeof his death, he was serving as a member of the Cellular,Molecular and Integrative Reproduction Study Section(CMIR), among his other study section activities.

On a personal level, virtually anyone would describeMatt in the following way: If one did not like andrespect Matt, then they must not like people or respectanyone! Matt was a wonderful human being, one whowas kind, helpful to anyone (he never recognizeda ‘‘pecking order’’), willing to give without expectinganything in return. His decency and civility were evidentregardless of the circumstances. He possessed the rarestof qualities: He was a person who did not wait to beasked when there was need, but rather volunteered. Hewas a wonderful, patient teacher and fine mentor. Hewas highly principled, strong, consistent, and everthoughtful of others. He served without making noise,without calling attention to himself. He had become anoutstanding scientific leader who knew how to makehard decisions and to enlist others. He shared his scienceand his dreams with his lifelong partner, Dianne. He willbe sorely missed by his father, brother, sister, wife, andhis many friends.

Barry R. ZirkinPeter N. SchlegelBernard Robaire

Sally D. Perreault (Darney)

2 Journal of Andrology N January !February 2008

Memorial: Emil Steinberger, MD, FACE, 1928–2008

It is with great sadness that I inform you that Dr EmilSteinberger died October 12, 2008. Dr Steinberger wasthe first president of the American Society of Andrology(ASA) and played an important role in the initiation anddevelopment of the ASA. Our field has lost a trueRenaissance man, talented in many areas beyond thoseof reproductive biology and medicine.

Dr Steinberger attended medical school in Germanyand then came to the United States, where he graduatedwith an MD from the State University of Iowa Collegeof Medicine in 1955. In fact, he would have been the firstperson to be awarded an MD, PhD degree from theUniversity of Iowa, but the Dean insisted that he couldnot receive 2 degrees at once! He refused to stay for anadditional semester, although he completed all require-ments for a PhD degree. Nevertheless, this basic sciencetraining greatly contributed to his depth of understand-ing and appreciation of the complexity of malereproductive biology. Dr Steinberger then moved toWayne State University Medical School for specialtytraining, followed by a position as a senior medicalresearch officer for the National Naval Medical Center.He served as chairman of the Department of Endocri-nology and Human Reproduction at Albert EinsteinMedical Center from 1965 to 1971. In 1971 he moved toHouston, Texas, to assume the position of chairman ofthe Department of Reproductive Medicine and Repro-ductive Biology at the newly created University of TexasMedical School at Houston. This department was aremarkable blending of the talents of clinical endocri-nologists, steroid chemists, biochemists, and cell biolo-gists, all with research interests focused in malereproductive biology. Together with his associates KeithSmith, MD, and Louis Rodriguez-Rigau, MD, DrSteinberger clinically evaluated each of the infertilepartners, because these andrologists recognized that thecomplexities of successful reproduction reflected thereproductive potential of the couple together rather thanjust the problems of the female partner. In 1983, DrSteinberger was honored with the Ashbel SmithProfessorship. Together with Drs Smith and Rodriguez,he then established the Texas Institute of ReproductiveMedicine and Endocrinology in 1983.

Much of his career focused on the hormonal controlof spermatogenesis. His studies of spermatogenesis

began in the mid-1950s when he worked together withthe famed Dr Warren O. Nelson. His early research setthe stage for work that continues in many andrologylaboratories today, and included studies of the gonado-toxicity of alkylating agents; in vitro penetration ofcervical mucus by sperm; human sperm cryopreserva-tion; effects of heat, ischemia, and cryptorchidism onspermatogenesis; and the controls of the hypothalamic-pituitary-gonadal axis. In the mid-1970s he published apaper on the frequency distribution of sperm counts offertile and infertile males, noting that sperm counts,except at the extreme low end of the scale (in the absenceof other deficiencies), were not major factors in acouple’s infertility, and thereby set the stage for thedevelopment of sperm function testing. Other clinicalwork focused on the endocrine manipulation ofhyperandrogenic women and those with other causesof ovulatory dysfunction. With his wife, Dr AnnaSteinberger, testicular organ culture and in vitrospermatogenesis were established. Together they report-ed on the presence of a follicle-stimulating hormone(FSH)–inhibiting protein secreted by Sertoli cells thatacted on the pituitary. Today we know that this proteinis inhibin. Work in his department focused on androgenand FSH regulation of spermatogenesis and Sertoli cellfunction, Leydig cell steroidogenesis, sexual behavior,and other aspects of reproductive biology. Dr Steinber-ger was honored for these achievements by the ASA asthe recipient of the Distinguished Andrologist Award in1987.

In his later years, Dr Steinberger became an authorand published 2 books on his life before, during, andDOI: 10.2164/jandrol.109.007575

Journal of Andrology, Vol. 30, No. 3, May/June 2009Copyright E American Society of Andrology

after the Holocaust. The first, Between the Devil andthe Deep Blue Sea, tells of his experiences as a youngboy growing up before, during, and after the SecondWorld War. It is an amazing story that chronicles hisfamily’s flight from Poland, escape from extermina-tion camps, and imprisonment in a Soviet labor camp.After the war, he attended medical school prior tomoving to the United States. His second book, ThePromised Land: Woes of an Immigrant, relates hisexperiences beginning with his arrival in the UnitedStates with just $20 in his pocket. There is much to belearned from his story of perseverance to overcomegreat obstacles in life, a man who ultimately became aleader in our field—a leader who will be missed by allwho knew and loved him.

Emil Steinberger was passionate about andrology andmedicine, hard-working, demanding, innovative in hisresearch, challenging to his students, and devoted to hisfamily. However, he also lived life to the fullest—enjoying his family and friends, sailing, travel, and writing.

He enthusiastically embraced life and showed unflinchingloyalty and support to his family and trainees. Evenafter learning of his illness, the Steinbergers traveled tothe far reaches of the world—retracing their flightthroughout Eastern Europe and Central Asia duringWorld War II; traveling to South America, Asia, andAustralia; and visiting with their many friends andcolleagues. Every moment of life was precious andenjoyed. His loving wife Anna, his 2 daughters andgrandson, his sister, his family and friends, and hiscolleagues and students will all miss him. His contribu-tions to the field of andrology are legendary. He willremain a role model of the ultimate physician-scientist/andrologist and the first president of the ASA. He leavesus his memory to cherish with the implicit reminder thatwith integrity, love, loyalty, determination, and drive wecan all accomplish the impossible if we, like him, pursueour dreams of excellence.

Dolores J. Lamb

350 Journal of Andrology N May !June 2009

Memorial

Dr C. Alvin Paulsen, a founding member of theAmerican Society of Andrology (ASA) and an EmeritusProfessor of Medicine at the University of Washington,died on December 18, 2008, at age 84. Dr Paulsen wasinternationally well known and admired for his work inclinical, teaching, and investigative male reproductiveendocrinology and biology. Dr Paulsen was a dedicatedphysician, an inspiring teacher, an innovative clinicalscientist, a lifelong friend and mentor to many, and aloving husband, father, and grandfather.

Dr Paulsen was born and raised in Portland, Oregon.He attended the University of Oregon as an undergrad-uate and for medical school, interrupted by time in theUS Navy in World War II. At Oregon, he did work inmale reproductive endocrinology with Dr Carl Heller,his early career mentor, including a presentation as astudent to the Endocrine Society and a publication. Hewent on to a busy internship and residency at DetroitReceiving Hospital (where he worked with anotherrecently deceased Andrology pioneer, Dr Emil Steinber-ger) and continued at Wayne State as an EndocrinologyFellow with Dr William Maddock. In 1958, he moved toSeattle, where Dr Heller then worked, and Dr Paulsenstarted his lifelong academic career at the University ofWashington, rising to Professor in 1970. Dr Paulsen wasthe long-term Chief of Endocrinology at the US PublicHealth Service Hospital in Seattle, which served as atraining mecca for approximately 44 EndocrinologyFellows over the years.

Dr Paulsen was an active physician throughout hiscareer, maintaining a group of patients who consideredhim to be their primary physician. This fascinating andloyal patient group was one of the first things thatattracted trainees to his program. Dr Paulsen’s wife,Wanda, was an active partner in this regard, contribut-ing her nursing and administrative work to his Saturdayclinics, as well as to his research work. He was also afirst-rate teacher both at the bedside, modeling excellentpatient care skills, as well as in his lectures, wellillustrated with carefully prepared patient and labora-tory pictures and data.

As an investigator, Dr Paulsen was courageous andoriginal, elucidating areas that were understudied, suchas Klinefelter Syndrome, a very common condition thatwas poorly understood. His work in Kallmann Syn-

drome was an excellent example of using the assessmentand treatment of patients as a means of understandingkey aspects of genetics, physiology, and pathophysiol-ogy. He also studied male infertility and male contra-ception, 2 inadequately addressed areas of wide societalinterest and importance. His work was careful, meth-odologically accurate, clearly reported, and repeatable.He was a very early adapter of many technologicaladvances for the study of human beings, includingchromosomal analysis, radioimmunoassays of hor-mones, and improved techniques for biopsies andseminal fluid analysis. He selected outstanding technicalstaff who were very loyal to him, in some casesdedicating their entire careers to working with him.DOI: 10.2164/jandrol.110.011569

Dr C. Alvin and Wanda Paulsen

Journal of Andrology, Vol. 32, No. 2, March/April 2011Copyright E American Society of Andrology

Dr Paulsen was an international leader in hisdiscipline. He published widely (over 150 contributions)in the scientific literature and provided the excellentchapter on the testes for the original and widelyinfluential Textbook of Endocrinology, edited by RobertH. Williams, the first Chairman of the Department ofMedicine at the University of Washington. He served oneditorial boards, including the Journal of Andrology andthe Journal of Clinical Endocrinology and Metabolism,and advisory groups for the National Institutes ofHealth, the National Pituitary Agency, the US Foodand Drug Administration, the United Nations FamilyPlanning Program, and the US Agency for InternationalDevelopment. He played a particularly influential rolein the World Health Organization programs in malefertility and its control. Dr Paulsen was a foundingmember of ASA and his service was recognized by hiselection to the Presidency in 1979 and his receipt of theDistinguished Andrologist Award in 1989. He was alsothe Society’s first Serono Award Lecturer in 1980 and itsfirst Distinguished Service Awardee in 1994. He also

served as President of the Fertility Society (1980), nowthe American Society for Reproductive Medicine. DrPaulsen ‘‘retired’’ in 1992 and as an Emeritus Professorremained active in academic programs until limited byhis health in recent years.

Dr Paulsen was preceded in death by Wanda, his dearwife of nearly 60 years, who passed away only a fewmonths before he did. He is survived by his 5 children,Chuck and Peter Paulsen, Sydney Bersante, Judy Hayes,and Linda Jones and their families, including 8grandchildren and 1 great-grandchild.

It is said that the great teacher never dies but lives on inhis students. The hundreds of students and residents, andparticularly the 44 fellows who trained under Dr Paulsen,are testimony to this truth. The fellows have gone on totrain hundreds more physicians and scientists, generatinga living legacy of Dr Paulsen’s very fruitful life. All of us,as well as the countless patients, families, colleagues,friends, and others who interacted with him are gratefuland recognize with respect and admiration the continuingeffect he exerts on their lives.

124 Journal of Andrology N March !April 2011

Richard J. Sherins, MD

Memorial

How many scientists can say that their work hasimpacted our planet in a positive and measurable way?Dr JoGayle Howard could. She was instrumental inhelping save some of the world’s most charismaticspecies through basic and applied research and trainingyoung professionals. It is with great sadness that wereport that Dr JoGayle Howard died March 5, 2011,after a prolonged battle with cancer. Dr Howard servedas the theriogenologist at the Smithsonian ConservationBiology Institute, National Zoological Park, in Wash-ington, DC. Dr Howard was a leader and inspiration forscientists within the American Society for Andrologyand the zoo and wildlife community. She was an activemember of the American Society of Andrology andserved as a member of the Executive Council (1993–1996); Vice-President, Women in Andrology Committee(1999–2000); and President, Women in AndrologyCommittee (2000–2001). Dr Howard also delivered theBuckeye Lecture in 1997.

Dr Howard received her DVM from Texas A&MUniversity in 1980 and her PhD from the University ofMaryland in 1989. Dr Howard’s career at the NationalZoo started in 1980, when, as a freshly graduatedveterinarian, she did a postgraduate internship incomparative reproduction under the supervision ofDrs David Wildt and Mitchell Bush. During those earlyyears, Dr Howard developed approaches and conductedfertility examinations on a host of wildlife speciesranging from bats to elephants while traveling the worldextensively, working both in the field in Africa and inzoos and breeding centers.

Dr Howard’s basic research resulted in the publica-tion of more than 100 peer-reviewed papers, 20 bookchapters, and numerous reports. She was a recipient of aprestigious Special Emphasis Research Career Awardfrom the National Institutes of Health. In recognition ofher many accomplishments, she was recognized forResearch Achievement (US Fish & Wildlife Service), asa Featured Conservation Scientist (British AirwaysExhibit at Millennium Dome, London), and with theUlysses S. Seal Conservation Award, the DistinguishedResearch and Scientist Award (American Association ofZoo Veterinarians), and the Recovery Champion Awardfor her leadership in helping save the black-footed ferret(US Fish & Wildlife Service). Dr Howard was active in

the Conservation Breeding Specialist Group of theIUCN–World Conservation Union, the Felid TaxonAdvisory Group, and numerous Species Survival Plansof the Association of Zoos and Aquariums. She alsoheld adjunct appointments at both the University ofMaryland and George Mason University. She exten-sively lectured to university, conference, and layaudiences worldwide.

Dr Howard’s research focused on fertility, infertility,and the role of reproductive technologies for promotingreproduction in wildlife. She was instrumental indeveloping novel approaches for propagating geneticallyvaluable animals, especially using artificial insemina-DOI: 10.2164/jandrol.111.013680

Dr JoGayle HowardPhoto credit: Nashville Zoo, Tennessee.

Journal of Andrology, Vol. 32, No. 4, July/August 2011Copyright E American Society of Andrology

tion. Her basic research interests largely focused onunderstanding reproductive mechanisms in never-be-fore-studied species and the phenomenon of teratosper-mia. Her specific contributions to the field of andrologyare summarized as:

N Developing technologies for semen and spermanalysis in diverse species, including the Africanelephant, rhinoceros, antelopes, felids, and the black-footed ferret both in captivity and in the wild.

N Demonstrating a link between a loss in geneticdiversity and an increased incidence of abnormalsperm, especially in felids and specifically in thecheetah, Florida panther, and clouded leopard.

N Characterizing cellular and molecular mechanismsinvolved in spermatogenesis, spermiogenesis, andsperm structure and function in felids.

N Demonstrating the negative impact of poor spermmorphology on low cryo-survival.

N Establishing the link between poor nutrition and theincidence of oligospermia or teratospermia in felids.

N Developing field-friendly sperm cryopreservationtechniques for numerous species, including thecheetah, clouded leopard, elephant, antelope, black-footed ferret, and giant panda.

N Demonstrating the transmissibility of feline immu-nodeficiency virus via semen (applicable to assistedreproductive techniques) and also developing pro-cessing techniques to minimize viral load.

Dr Howard was a champion for numerous species onthe brink of extinction. Her contributions to wildlifeconservation are summarized as:

N Recovering species that were on the edge of being lostforever. Especially relevant were her contributions tosaving the black-footed ferret that at one time wasdeclared to be extinct, until a small remnantpopulation was found in Montana in the early1980s. Zoos were called to assist, and, under DrHoward’s oversight, the National Zoo’s ferret breed-ing program flourished with the birth of more than500 kits, including 150 via artificial insemination.Most recently, she produced black-footed ferret kitsfrom sperm frozen more than 20 years ago. She alsoinitiated a multidisciplinary program to conduct abiomedical survey of wild black-footed ferrets toassess health, disease, reproduction, immunology,and genetics of reintroduced populations in theNorth American grasslands ecosystem. This programresulted in the successful reintroduction of the speciesin 6 US western states, Canada, and Mexico.

N Understanding the reproductive biology of species thatwere dramatically affected by small population size andlost gene diversity. The population of Florida

panthers collapsed to fewer than 50 individuals inthe 1980s because of habitat fragmentation in southFlorida. Dr Howard tromped through the swampswith her colleagues to assess the reproductive statusof Florida panthers, discovering an extraordinaryhigh incidence (.90%) of morphologically abnormalspermatozoa in this critically endangered subspecies,as well as other physiological defects, includingcryptorchidism. These findings helped the US Fish& Wildlife Service to implement a radical interbreed-ing strategy using the genetically distinct pumas fromTexas to restore genetic vigor.

N Applying reproductive technologies to enhance zoopopulations. Dr Howard was part of a team that, forthe first time, collected, cryopreserved, and importedsperm from free-ranging cheetahs in Africa (Namib-ia) for producing offspring in the United States byartificial insemination. The overall benefits of thiswork included vastly expanded knowledge of cheetahbiology and genetics, and improved the ability tomaintain a reproductively viable population througha combination of science and reproductive manage-ment.

N Enhancing reproductive efficiency in captive popula-tions of an iconic species. Dr Howard was a keymember of a multi-institutional team that workedside by side with Chinese colleagues to understandthe biology of giant pandas for the purposes ofunlocking the mysteries of poor reproduction. WithChinese colleagues, she developed new protocols forsperm cryopreservation and artificial insemination,often by training young Chinese professionals in therange country or her laboratory at the National Zoo.Dr Howard also is credited with successfully insem-inating the female giant panda Mei Xiang at theNational Zoo, producing the famous cub Tai Shan in2005.

N Championing for species conservation in range coun-tries. The clouded leopard is a beautiful yet secretivecat living in southeastern Asia. When the breedingprogram in US zoos was failing, Dr Howardestablished the Thailand Clouded Leopard Consor-tium that was devoted to breeding this endangeredspecies in the range country of Thailand. She and herteam have produced more than 50 cubs whilesimultaneously raising funds to support conservationprojects both in the field and in captivity. A subset of6 of these cubs has been imported into the UnitedStates to rejuvenate the American zoo breedingprogram, with another 8 offspring already produced.

The final component of Dr Howard’s passion forandrology, reproductive science, and conservationinvolved training a next generation of biologists. She

346 Journal of Andrology N July !August 2011

was an outstanding mentor with high ethical standardsand relentless focus on ensuring highest-quality scien-tific output. She was a constant source of inspiration fornumerous young students and professionals. Forty-sixundergraduates, 7 graduate students, 10 postdoctoralfellows, and 12 visiting scientists (from Russia, Thai-land, South Africa, Namibia, and China) benefited fromher mentorship and now carry on her legacy. DrHoward also conducted numerous training courses inwildlife reproduction, male fertility assessment, gametecryopreservation, nutrition, and veterinary medicine,including in Malaysia, Thailand, and China.

Dr JoGayle Howard was unique in having not onlytouched the lives of hundreds of people, but actuallysaving a species and contributing to the recovery ofmany others. We all have lost not only a superb scientistand advocate for our profession but also a champion forwildlife conservation. Dr Howard will forever beremembered with respect, love, and gratitude.

Budhan Pukazhenthi, written on behalf ofher family at the Smithsonian Conservation Biology Institute

National Zoological ParkFront Royal, Virginia

Pukazhenthi N Memorial 347

Memorial

On Monday, May 14, 2012, Dr Claude Gagnon passedaway after a long fight against Parkinson disease.Claude was one of the outstanding and most respectedreproductive biologists worldwide. His fundamentalcontributions to our understanding of sperm biologywill withstand the test of time, and his service to thescientific community will serve as a beacon for cominggenerations.

Claude earned his PhD degree in biochemistry atthe Université de Montréal in 1974 at a remarkablyyoung age. He first went to Basel University, Switzer-land, to do postdoctoral studies with Dr Hans Thoenenfrom 1974 to 1976, and then to the Laboratory of Cli-nical Sciences, National Institute of Mental Health,Bethesda, Maryland, with Dr Julius Axelrod, from1976 to 1978. He was appointed as an assistantprofessor in the Département de Pharmacologie,Faculté de Médecine, Université Laval, in QuébecCity in 1978 and was promoted to associate professorin 1983.

In 1984 Claude was recruited as an associate pro-fessor by the Urology Division of the SurgeryDepartment of McGill University to build the UrologyResearch Laboratory at the Royal Victoria Hospital,and in 1993, under his leadership, this lab was extendedto the Montreal General Hospital; both hospitals arepart of the McGill University Health Centre. Duringthe 20 years he served as director of these laboratories,Claude made numerous world-class contributions indifferent areas of reproductive sciences (andrology,surgery, and prostate cancer) in close collaborationwith clinical urologists.

For his outstanding contributions, Claude receivednumerous honors, including Scientist Awards in 1983and 1988 from the Medical Research Council ofCanada, the Young Investigator Award from the Clubde Recherches Cliniques du Québec in 1987 (given to anindependent investigator within the first 10 years of hisacademic career), Chercheur-Boursier de mérite excep-tionnel (Exceptional Merit Scholarship) from the Fondsde la Recherche en Santé du Québec (1988–1993), andthe Award of Excellence from the Canadian Fertilityand Andrology Society in 2003.

As a biochemist, Claude first made significantcontributions to our understanding of the role ofprotein carboxymethylation in secretion, but duringthe past 3 decades his passion for research focused onsperm physiology: he wanted to understand themolecular mechanisms involved in sperm motility,flagellar movements (using human spermatozoa andChlamydomonas models), and sperm fertilizing ability.In the 1990s, his lab pioneered the concept thatreactive oxygen species (ROS) play a physiologicalrole in sperm activation. This was a groundbreakingachievement because, at that time, ROS action wasconsidered to be strictly detrimental to sperm func-tion. Claude and his team dug deeper and deeper intothe signaling pathways that enable the spermatozoonDOI: 10.2164/jandrol.112.017426

Dr Claude Gagnon

Journal of Andrology, Vol. 33, No. 6, November/December 2012Copyright E American Society of Andrology

to fertilize an egg; his goal was to understand thecauses of male infertility in order to find a treatmentfor that condition.

Claude contributed to the operation of many societiesand played a leadership role in the organization ofmany congresses. He was the president of the CanadianFertility and Andrology Society (1996) and the orga-nizer of the International Symposium on Spermatology(1998). For that particular event, he demonstratedremarkable talents as an organizer and fund-raiser toassure the great success of the scientific meeting. Claudealso rendered major service to the American Society ofAndrology, both as an elected member of ExecutiveCouncil and chair of the Industrial Liaison Committee,and was a loyal member of the Society for the Study ofReproduction.

Claude had a gift for finding excellent collabora-tors and keeping lasting relationships with them. Heestablished fruitful joint research projects with scientistsall over the world. He could always find the right peopleand obtain grants to support these activities. Numerousgraduate students, postdoctoral fellows, and clinicianswere trained under his supervision; many of them havegone on to establish themselves as independent inves-tigators in the area of reproductive biology and fertilityresearch. All of his trainees warmly remember Claudefor his guidance and his ability to motivate theirprogress. Claude exemplified for them everyday lifeas a scientist and a researcher. Uppermost in the mindof those who worked with him are Claude’s sharpintelligence, his moments of major enthusiasm, and his

ability to analyze and discuss results, as well as hisingenuity in proposing ‘‘way-out’’ hypotheses. One ofClaude’s common sayings was, ‘‘Speculation is theonly inexpensive thing in research.’’ There were alsothe extreme moments associated with grant applica-tions and the euphoria of getting them—his lab wascontinuously funded from the time he opened his labin 1978 until 2011.

Claude had a love for life and a superb sense ofhumor; his friends and trainees share wonderfulmemories of going on fishing trips and of numerousparties (birthdays, Christmas, and any good news ofaccepted articles, prizes, and even sunny days).

Claude was an outstanding scientist and worldwideleader in his field; he had a talent for visualizing thingsthat others would not have considered important. Hewas a dedicated teacher and mentor. In addition to hismother, wife, sons, grandchildren, and family members,he will be sorely missed by everyone who had theprivilege to work, train, or simply discuss science withhim, because he had the ability to make people feel likefriends.

Cristian O’FlahertyEve de Lamirande

Pierre LeclercLinda Lefievre

Bernard Robaire

Published simultaneously in both Biology of Reproductionand Journal of Andrology

1126 Journal of Andrology N November !December 2012

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