1

Francisco Alberto Sez

First Cytogeneticist of Latin America

LIFE AND RESEARCH WORK

(1898 1976)

By

Mximo E. Drets
2014

MEMORIAL
The author of this contribution dedicates it to the memory of the Chemist Gloria Martnez de
Drets, the researcher who introduced the first biochemical investigations on nitrogen fixing
microorganisms in Uruguay. Her research activity allowed her to meet Prof. Francisco Sez at
our institution, with whom she maintained a close friendship and reciprocal admiration over the
decades. The almost half a century of close bond that the author, and his unforgettable wife and
intellectual companion, maintained with such eminent researcher, this book attempts to reflect the
deep mutual affection that permanently we cultivated and shared with Prof. Sez.
M.E. Drets, Montevideo, Uruguay.June30, 2012

Prologue
To the Master,affectionately
Dear Professor:
For a long time I wanted to write you but, you know, one is sometimes too indolent. This did not mean at
all that I had forgotten you. On the contrary, every now and then, some of your work reappeared reminding the
good times we had together. I began thus to read again all your work and the huge amount of documents and
letters that we exchanged and that I carefullykept in my library. I confess toyou that all these readings, some of
thempublished long ago, were always very inspiring life experiences reminding unforgettable scientific teachings.
There was a moment when I remembered your repeated an insistent request that I should write an obituary
about you but, I must admit, that I did not consider your request very seriously so I never satisfied you. Perhaps
your request produced in me a deep sadness rejecting it in my mind and in my heart.
But, dear Professor, astime passed since I started studying all your papers again an intense nostalgia
arose in my mind because they were not only the expression of your scientific work but also the long-lastinghuman
relationship we cultivated.
Thus, with this short note I decided herewith to write a long story particularly gratifying for me because
it's almost like a kaleidoscope of life that has brought me so many shared experiences. Not only
willemotionsreappear, memories, and my own words, joined so other valuable mutual friends, all of which have
had the virtue of recallingbeautiful times gone but not forgotten, always deeply persisting in my heart.
I hope that this work really represented what you wanted me to do. If so, I will have achieved a goal that
surely will join us closer as colleagues and as friends.
Also, I sincerely hope that one intelligent young student reading this story of life, could fall in love with
chromosomes as it happened with both of us!
Your disciple, always, with all my affection and faithful admiration,beyond the times.
"Maximn

The Origins
Biographical shades
Sez was born on March10th, 1898, in the city of Montevideo, Uruguay, in an old house that was
purchased from to Dr. Perez Scremini. He wasthe youngest of six children of Francisco Leoncio Sez Lpez(18491916) and Luisa Snchez Represa(1857-1939) who had married on April 19, 1873, in the city of Mercedes
(Department of Soriano, Uruguay). His parents were originally from Mercedes, and married in that city. His father
was ascrivener of the Government Treasury of Uruguay under Presidents John Idiarte Borda and Lindolfo Cuestas.
Idiarte Borda, from Mercedes, was also member of the family through his wife Luisa Snchez.
His mother, a woman of great character, was a widelyappreciated lady. She devoted her life to her family
and to charity. She was a member of the Society of St. Francis of Paul, where she practiced charity, like other
ladies from Montevideo.
The well-off family was noted for theirhigh cultural level, not very common in those days, being one of
their brothers, Carlos Federico Sez, (1878-1901), one of the greatest painters of Uruguay, who died at the age of
22 years. Other brothers were Francisco Arturo, also died early, Eduardo Mario, outstanding ocean captain, Sara
Silvia, and Maria Luisa, opera singer, all of them born in Mercedes.
Francico Alberto Sez had five children. The older he was very young, Carlos Alberto, from his union with
Marina Rossi. Then, he married Matilde Dolgopol, first Paleontologist of Argentina, being his sons Federico and
Jorge Carlos Alberto, both medical doctors. Finally, from his marriage with Rosa Gershanik, had two children,
Graciela Luisa,who becameDirector of the Historical Museum of Morn, Argentina, and Eduardo Francisco
Guillermo, an art photographer. His childhood was spent amid deep solicitude and affection by his family. His
surprising arrival to the world, 16 years after his last brother, represented a happy event for all of them. However,
the joy did not last long because of the deathof his ill-fated brother Carlos Federico that occurred in 1901 in the
gardened house ofPiedras Blancas. This home, years later, was the residence to the end of his life, of Don Jos
Batlle y Ordez, who later became President of the Republic.
Francisco Alberto, who was nicknamed "Papico", spent his early years in the home of Misiones Street, a
residence that had large marble staircases and spacious rooms adorned with artwork and exquisite decorations,
many of them brought by Carlos Federico from his trips to Rome,when he had been awarded a scholarship the
Government of Uruguay to study painting. A "guardian" gave way to the visitors, especially during the days when
Doa Luisa entertained their friends who wereattended by seven servants.
His family later moved (1902) to a house in Trouville (Pocitos,Montevideo) that was very close to the
beach, being a very happy time for Sez, who enjoyed friends and sports. The sea was very important in his life,
and always lived in Montevideo very closeto it.
When Papico was only seven years old, during ahorseracing excursion with his familythe animal dropped
one of his legs in a vizcachera(a hole in the soil made by a rodent called vizcacha) causing the child to fall. This
accident caused a fracture of his right femur that obliged him tospentseated more than a yearwith his brokenleg

depriving itself of the free expansion of the childhood years. He remembered being in a wheelchair watching the
enjoying games of his friends.
Striking enough, after so many years, the eminent Uruguayan pediatricians Profs. Drs. Luis Morquio and
Francisco Soca prescribed this long period for fixing his legs.The famous doctors suspected that, in reality, the
bone lesion was not a simple break but probably an underlying infectious process that prolonged curingthe
wound.
As at that time there was no available means of treating microbial diseases so doctors wereforced to
prescribe such prolonged treatments. It is plausible to speculate that the infected bone fracture must have
compromised the growth cartilage of the femur which resulted in one of his legsbeing shorterthan the other.
Besides, at that time, it was very frequent to have familiarcontamination of tuberculosis.Therefore,
perhaps the child was infected by his younger brother Carlos Federico who was unfortunately suffering from the
terminal phase of this disease. Thisclinical interpretation is possible because Prof. Saez personally commented to
the author, during our stay in Mexico City in 1974, that he also had the impression that this medical interpretation
could be the real cause of their prolonged child suffering and shortening of his leg.
That period of his childhood was undoubtedly of great suffering and he could not play like other children,
however, this difficult situation of long convalescence did not affect his future characterwho, however, was
always a very cheerful person characterized by sympathy, and a natural kindpersonality.
As he recovered, he started playing outdoorsfeeling very attracted by the sea, spending long hours on the
rocks of Pocitos contemplating the changing landscape. He knew the recesses of the coast and the places where
different animals of the local fauna inhabited.His healthand fitness allowed him to play sports since his teens. He
practiced rowing and swimming in the Nautilus Club of Montevideo, especially during summer time, until the
end of his life. He also devoted energy to boxing allowing him to become anamateur champion of boxing, at the
age of l5 years.Fighting repeatedly, Saez posted this hobby into an album that he made himself with photos and
newspaper clippings, which he titled "My Life in the Ring". Additionally, Sez performed public exhibitions of
boxing styles together with Mr. Alberto F. Suppici, who was the athletic Trainer of the Uruguayan football team
that won in 1930 the final game with Argentina when Uruguay became world champion for the third timeof its
football history
The discipline at school did not agree with the Sez character, so he had to be educated in three different
colleges namely; the College directed bythe Uruguayan Adolfo Berro Garca and Mr. Ducusseau (who died
during the war in France l914).During the Seminar time Sez always remembered that the priest repeated, "You
do not fight Sez, you do not fight" so he was not accepted there and the same happened atthe Barcel
College.Finally, Sez had to make his first letters with a private teacher who was attending daily his house, Ms.
Eloisa Sacarello Castellanos, remembering her as a sweet woman who patiently guided his rebellioushand using
the Vzquez Cores notebooks, very popular school items at the time.Saez started High School being a student
without any discipline. The course tests were taken freely according to his preferences. His father, probably not
very pleased with his sporting triumphs, employed him at the FrigorificoSwift slaughterhouse(1914) where he had
to get up at 6:30 am, every day,crossing the bay of Montevideo in a tugboatthat went from the piers of
Montevideo bay to go to the Cerro Maciel including backhazardous returning trips during evening storms.

His new employement lasted one year (1915) but, however, he continued studies. When he was 18 years
old he suffered the sudden loss of his father. He confessed that, at that time, he needed his intelligent direction
and advice. Sez remembered that they were both big fans of field shooting frequently going hunting in the
country. Also, one of his favorite walks with his father was invariably to visit the zoological garden of Villa
Dolores. There, his father, a connoisseur of animal life, gave interesting explanations to his son Sez who listened
him with great attention.
At that time,the Institute of Industrial Chemistry directed by Dr. Latham Clarke1 was established.This
renowned American Chemist was .aided byseveral friends to manage the Institute,such as Raphael Batlle
Pacheco, Silvio Moltedo, Horacio Terra Arocena, Ricardo Garese and other important people of the time.Sez
completed two years of coursespreferentially chemistry. He had installed a laboratory in his home, carrying out
experiments repeating or practicing the ones he developed at the Institute. A beautiful Spencer microscope, a gift
from his mother, completed his equipment. It was then when he began to explore the microscopic world,
especially the hydrofauna, which increased his enthusiasm about microscopy.
Sez mentioned that he read a lot and unsystematically, scientific books, especially
biology,philosophy,poetry, and literature.Graduallyhis inexperienced library was enriched by his kind mother
always justifying these expenses, sometimes excesive, made in the old bookstores Garca and Monteverde of
Montevideo, very popular in that time.
His enthusiasm for sports decreased while his interest in spiritual expressions increased. He regularly
attended the Vaz Ferreira conferences on philosophy and other cultural events. Books of science and philosophy
of science were his favorites. The fermentativepower that produced reading these books including Rules and
Counselings on Biological Research by Ramn y Cajal2 together with the book of Wilhelm Oswald 3 The great
men was very intense and he could not stop reading every night the wonderful and challenging works which
already influenced a lot on his spiritual background.
At that time, a group of young universitarian students and intellectuals, created the Center of Students
"Ariel". Sez became member of the Board of Directors for several years together with Carlos Quijano, Arturo
Acevedo, Adolfo Folle Joanic, Augustine Ruano Fournier, Carlos Rodriguez Pintos, Luis Giordano, Luis
Pieyro Chain, Carlos Benvenuto, Julio E. Barrios, Enrique Amorim, Arturo Quesada and other associates. This
center had a significant activity inside and outside the University and constituted, at that time, one of the most
advanced groups of members with reformist principles. As a result of these conferences, Sez wrote an essay on
The Personality in Science that has unfortunately remained so far in complete anonymity.
________________________________
1-. The Institute of Industrial Chemistry was created by law in 1912, having as one of its fundamental tasks the national
fuel production based on alcohol, promoting research, development and production as well as teaching and technical
training. Dr. Joaquin Enrique Zanetti was appointed as Director and Dr. Latham Clarke as Assistant Director (West
Kingston, USA, 1881 - Montevideo, Uruguay, 1962). In 1913, after the resignation of Zanetti few months after his
appointment, Latham Clarke assumed the position of Director he held until late 1926. When in 1931 the National
Administration of Fuels, Alcohol and Portland (ANCAP) was created, Latham Clarke was member of its directory
(Martnez 2007).

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2 - The book Rules and advices on biological research: tonics of the will was written by Santiago Ramn y Cajal
(Petilla of Aragon, 1852 - Madrid, Spain, 1934). Nobel Prize in Physiology or Medicine (1906); thisspeech was read
during his admission as a member of the Royal Academy of Exact, Physical and Natural Sciences of Spain in 1897.
3 - Friedrich Wilhelm Ostwald (Riga, Latvia, 1853 - Grossbothen, Germany, 1932). Chemist and philosopher who
received the Nobel Prize in Chemistry in 1909 for his work on catalysis and research on fundamental principles
governing chemical equilibria and reaction rates.

Atthat time, chemistry no longer polarized his attention and he wanted to leave the country to study
biological sciences.The arrival in Montevideo (1930) of the distinguished Spanish physiologist Augusto Pi Suer4
helped turn his irresistible attraction to biology. He not onlyattended his lectures but also read the magnificent
abstracts published in the press by a notable reviewer who he then knew was a young school teacher named
Clemente Estable. While this was happening Sez was looking for ways to travel abroad in search of new
horizonts for his biological studies. He was determined to go to Paris and enter in the Faculty (i.e. The Sorbone) to
continue there his career dreams.
Pi Suer dispelled doubts about the direction of his studies because, personally consulting this professor, he
counseled him that instead of medicine, to study sciences, which much comforted Sez because this was precisely
what he ardently desired to do. Sez mother sadness manifested herself at every moment at the prospect of his long
stay in Europe, expressing a well justified fear that the illness of his brother Carlos who was in Rome for seven
years returning ill to die soon afterfearing a repetition. This situation enforced him to cancel the trip to Europe and
to search for a closer place. Therefore, he went to the city of La Plata in February 1921. There, in theErudite
University he began his studies of a Doctor in Natural Sciences and Professor in Biological Sciences at the
Institute of the Museum of La Plata and the Faculty of Humanities respectively. Saez arrived there for only a few
years but remained 27!
In La Plata he had the privilege of having excellent teachers, mostly Europeans. Walter Schiller, geology;
Santiago Roth, paleontology; Robert Lehman-Nitch5, Anthropology; Augusto Scala, Botany; Jacob Cristofred,
neurobiology; Fritz Kopatchek, biochemistry; among others.
But the great figure among his teachers was the renowned zoologist Miguel Fernndez, discoverer of
polyembryony in the armadillo. who he was a disciple of the famous Swiss zoo1ogist Arnold Lang. Dr. Fernndez
was an exemplary scholar with whom Sez cultivated a fraternal intimacy in the daily and long hours in the
Laboratory Zoology of Museum of La Plata, where he worked for many years following his brilliant courses and
learning his rigorous scientific research methods.
Dr. Fernndez was responsible for teaching eight courses divided in zoology, general biology,
protozoology, embryology of invertebrates, vertebrate embryology, comparative anatomy, heredity and biometry.It
was Dr. Fernandez who, for the first time, started giving a course on Heredity in the laboratory. Sez devoted 40
hours per week of work in the Laboratory of Zoology of comparative anatomy of invertebrates and vertebrates
___________________________
4 - Augusto Pi Suer (Barcelona, Spain, 1879 - Mxico City, Mxico, 1965) was a prominent physician,
physiologist, writer and politician. His scientific research work began with his doctoral thesis on the anaerobic

8
life, by which he obtained his doctorate in Madrid in 1900. It was a great-nephew of Francisco Suer
Capdevila, the first Dean and Head of the Department of Physiology, Faculty of Medicine, Uruguay, founded
in 1875.

and microscopy. He also studied other courses in different faculties of the University, like medicine,
physiology, biochemistry and biophysics, veterinary, anatomy of domestic animals, histology and embryology,
general pathology, microbiology and hygiene, and in the Faculty of Humanities, pedagogics, philosophy and
literature.It was the time of the universitarian reform and Sez often actively participated in student protests in
defense of the ideological currents that would help to strengthen the new university.
In this quiet town of lime trees his youthful dreams crystallized achieving scientific and teaching activities
in compliance with all the steps required to become a professional student. His first official appointment (1927)
was produced in the most unexpected way. Sez finished taking the exam in anthropology after "with standing for
one hour the ravages of Prof. Lehmann-Nitsche5" who wanted to crush him in angry manner because their answers
about the origin and evolution of man were radically different from those expected by the famous
anthropologist.Finally, concluded the stormy exam and Sez walked away to wait the result taking for granted a
failure. At this point he saw that Dr. Luis Torres6, archaeologist and Director of the Museum of La Plata, turned
away from the group of members of the examining table. Beckoning he came near him telling: since today you
are appointed as Assistant. Completely confused Sez thanked him and asked "what about the examination? "You approved. The exam was great! "
After graduating with high scores (10 subjects with Distinction and 6 Distinguished) at the University of La
Plata (Sez was the first graduate in the area of Biological Sciences in the history of the University) He was
appointed in April 1927 by competitive examination test, Laboratory Assistant of the Section Animal and Plant
Cytology at the new Laboratory of Biological Sciences, created in Montevideo for a talented young researcher just
back from Spain, where he had worked for three years with Santiago Ramn y Cajal: Clemente Estable7.
_______________________________________
5 Robert Lehmann-Nitsche (Radomitz, Prussia, now Poland, 1872 - Berlin, Germany, 1938). Ethnologist,
anthropologist and physician, after a doctorate in natural sciences (Freiburg, 1894), anthropology (Munich, 1894) and
medicine (Berlin, 1897), travelledto and settled in Argentina at the age of 25 years. There he worked in the Research
Institute of the Museum of La Plata, where he was Director of the Department of Anthropology. He made a deep study
of the cultural characteristics of the country investigating the material culture, folklore, linguistics, ethnology and
mythology. In 1930, after his retirement, he returned to Germany to settle in Berlin until his death.
6 The Argentine naturalist and archaeologist Luis Mara Torres (Buenos Aires, Argentina, 1878-1937) was Director of
the Museo de la Plata between 1920 and 1932
7 Estable, Clemente (called Saint Lucia Margat - today, 1894 - Montevideo, Uruguay, 1976) was a renowned educator, scientist
and philosopher Uruguayan son of Italian immigrants. In 1922 he received a grant from the Spanish government to study at the
Institute for Histological Research in Madrid, at that time under the direction of Santiago Ramn y Cajal, Nobel Prize winner. He
remained there three years researching the nervous system and publishing his first scientific papers. Then, he continued studies in
biology and psychology at institutes in France, Germany, Austria, Monaco and Italy, working with renowned scientists. Returning to
the country in 1927 he was appointed Director of the Laboratory of Biological Sciences, created for research, where he began a
tireless scientific research activity, publications, conferences and teaching.

Figure 1 Genealogy of the Sez family.

I, 1 - Pedro Prez de Velasco; I, 2 Josefa Rodrguez Da Cunha, I, 3 - Juan Manuel Echavarry; I, 4 - Mara
Medina y Pavn. -- II, 1 - Francisco Sez; II,2 - Francisca Prez de Velasco, II, 3 - Jos Lpez; II, 4 - Francisca
Fernndez; II,5 - Jos Rodrguez; II, 6 - Mara Petrona Echavarry y Medina; II, 7 - Matas Snchez; II, 8 - Manuela
Gonzlez; II, 9 - Flix Martnez; II, 10 - Jacinta Velasco; II, 11 - Juan Manuel Represa; II, 12, unknown name- III,
1 - Francisco Sez; III, 2 - Clorinda Rodrguez; III, 3 - Juan Jos Lpez; III,4 - Mara Francisca Rodrguez y
Echavarry; III, 5 - Ildefonso Snchez; III, 6 - Manuela Martnez y Velasco; III,7 - Santiago Represa; III, 8 -J. Daz
- IV, 1 - Emiliano Sez y Rodrguez; IV, 2 - Mara Josefa Lpez y Rodrguez; IV, 3 - Gregorio Snchez y
Martnez; IV, 4 - Luisa Represa Daz; V, 1 Francisco Leoncio Sez Lpez; V, 2 - Luisa Snchez Represa -- (VI,
hermandad Sez-Snchez); VI, 1 - Francisco Arturo; VI, 2 - Carlos Federico; VI, 3 - Eduardo Mario; VI, 4
Malvina Urtubey; VI, 5 Sara Silvia; VI,6 Jos Len Ellauri; VI, 7 - Mara Luisa; VI, 8 - Francisco Alberto
Sez; VI, 9 - Marina Rossi; VI, 10 - Matilde Dolgopol; VI,11 - Rosa Gershanik - VII, 1 Carlos Alberto; VII, 2
Eduardo Mario VII, 3 - Ricardo Len; VII, 4 Mara Luisa; VII, 5 Carlos Alberto; VII, 6 Sara Reina; VII,
7 Carlos Alberto; VII, 8 Federico Carlos; VII, 9 Jorge Alberto; VII, 10 Graciela Luisa; VII, 11
Guillermo Francisco Eduardo.

10

Figure 2 The boyPapico.

Figure 3 -The student Sez working in the Laboratory of Zoology

of La Plata Museum, June, 1921.

11

Prof. Sez, Ergasto Cordero8and Jos Mara Martnez, a young assistant appointed to assist Estable, opened
the doors of the romantic laboratory of Milln Avenue (1927) (see Fig. 5a) which over time became the Institute of
Biological Sciences whose name was changed in 1976 in honor of the late Prof. Estable for "Institute for
Biological Research" Clemente Estable.
Economic and familyreasons very reluctantly forced Sez, after three years of work, to return to La Plata.
However, he was always close in spirit with his great friend Estable and during the summer holidays he used to
return to Montevideo at his workplace, the Laboratory of Biological Sciences.
His early and faithful bond as a researcher with the Institute induces one to consider Sez, as a scientific
co-founder of the institution initiated by Estable whose close relationship remained until his death.
During his stays in Montevideo, Sez, Estable, Varela Fuentes, the Rubino brothers and other partners,
founded the Society of Biology in Montevideo after the International Congress of Biology of 1930. Also with
Estable, Ergasto Cordero and Vogelsang Sez founded the Uruguayan Society of Natural Sciences, with the
cooperation of some botanists, zoologists and paleontology enthusiasts of the time.

______________________
8 - Ergasto Hctor Cordero (Montevideo, Uruguay, 1890-1951) was the first Uruguayan zoologist. From an early age
he showed interest in the natural sciences, and defined his vocation in zoology during his high school years. In 1911 he
joined the Faculty of Medicine, graduating in 1918 as Doctor of Medicine with a thesis entitled Studies on some
protozoan ciliates of Uruguayan freshwater. Between 1922 and 1924 he deepened his studies of zoology in Europe,
mainly in Germany, and back in South America travels to Brazil and Venezuela where he continued his teaching and
research career. In 1928 he entered by competition the Laboratory of Biological Sciences headed by Estable. In 1942 he
was appointed Director of the National Museum of Natural History, a position he held until the end of his life.

12

Figure 4 - Sez in the first Institute of Biological Sciences Research in 1928.

(Photography partially restored).

13

Sez and the Institutescientific diversity

Sez began working from the initial time of its foundation in the Instituto de Investigaciones of Ciencias
Biolgicas in August 1st, 1927, as did the young assistant of Estable, Jos Mara Martnez and the great biologist
1
Ergasto Hctor Cordero.
Sez recalls that unforgettable moment: We were the 4 musketeers of the emerging biology that, with the
passing oftime flourished in the present Institute. The Institute was set up initially in a small house in Milln
avenue (Fig.5a) and soon moved to a larger house, also in Millan Avenue where they remained until the early 50s
(Fig. 5b). After that, the Institute moved to a new present building constructed specifically as a research Institute
located in Italy Avenue, Montevideo.

Figure 5 Institute for Biological SciencesResearch.a: first site: Milln Ave 3872; b: second site: Milln
Ave. 4096.

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Figure 6 - First four investigators of the Institute.a: Prof Ergasto Hctor Cordero, (zoologist);b: Prof Sez
with his friend Prof. Julio Mara Sosa, (histologist), picture taken during his visit to the Andes University, Mrida,
Venezuela, (circa September, 1968);c: Wilhem Franz Herter, (botanist),in 1924.

15

The second site of the Institute was an old house (Fig. 5, b) located very close to a small beautiful stream,
the Miguelete brook. However,during the rainy seasonseveral precarious laboratories of the old house were always
flooded by uncontrollable inundationsdestroying experimental material under investigation and the modest
research installations established with great effort by Estable. In the middle of this terrible panorama appeared Dr.
Harry Milton Miller9, Associate Director of the Rockefeller Foundation, who was visiting a number of Latin
American research laboratories to estimate theirrequirements concerningnew instrumentation and reagents needed
for scientific development. It is easy to imagine the sad impression of Dr. Miller looking at the flooded modest
laboratories of the Institute. Facing this miserable situation of the Estable laboratory, Miller decided to support the
Institute financing the cost of instruments, reagents, and fellowships, covering travel and living expenses for one
year specifically dedicated to those researchers interested in developing scientific stays in the United States.
One of the first candidates was Sez who conducted investigations on Cytospectrophotometry (see page
68). At the end of his stay in the USA the Foundation bought all the parts needed by Sez to build a first
cytospectrophotometer to be installed in our Institute. The second fellow was Roberto Sotelo who spent one year
becoming familiar with the use and applications of Electron Microscopy. This program of the Rockefeller
Foundation financed the developmentof various research laboratories of the Institute and covered several years of
support (from 1948 to 1954 approximately). In addition, the Rockefeller Foundation donated additional funds to
finish the construction of a building for the conference and library facility.
______________________
9- Harry Milton Miller (1895- 1980) served the National Sciences Division of the Rockefeller Foundation as a
Fellowship Administrator (1932-1934), Assistant Director (19341946), and as Associate Director (1946-1950). The
Rockefeller Foundation is a philanthropic organization and private foundation based at 420 Fifth Avenue, New York
City. The famous institution was established by the six-generations of the Rockefeller family. It was founded by John
D. Rockefeller ("Senior"), along with his son John D. Rockefeller, Jr. ("Junior"), in New York State, May 14, 1913,
when its charter was formally accepted by the New York State Legislature. Its central historical mission is "to promote
the well-being of mankind throughout the world." Although it is no longer the largest foundation by assets, the
Rockefeller Foundation's pre-eminent legacy ranks it among the most influential National Government Organizations in
the world. By year end 2008 assets were tallied at $3.1 billion from $4.6 billion in 2007, with annual grants of $137
million..

16

Figure 7.a: Harry Miller, President of the Rockefeller Foundation, Francisco A. Sez and Clemente
Estable. b: Professor Estable becoming familiar with the controls of the Siemens Electron Microscope
obtainedthrough funds donated by the National Institutes of Health (USA) and Uruguayan Banksin the decade of
the seventies.
Dr. Miller also helped financially other Latin American Countries,particularly Brazil. When Miller visited
Sao Paulo he met Dr. Andrs Dreyfus who told him that they were more interested in developing the area of
Genetics in Brazil instead of receiving funds for establishing laboratories. Miller listened with great interest this
proposal and when returning to New York he immediately contacted Professor Theodosius Grygorovich
Dobzhansky, an eminent Russian geneticist, trying to convince him to go to Brazil to spend an undetermined time
there giving courses and developinggenetics research all financed by the Foundation. Dobzhansky accepted the
offer and went straight away to Sao Paulo. At the airport several geneticistswere waiting for his arrival and
discussing about thepotential difficulty of an English speaking professor giving courses to students speaking
mainly Portuguese. The great surprise to the waiting group occurred when Dobzhansky rapidly left the plane and
started telling them this historical phrase: Se os senhores falarem devagar eu posso entend-los.(translated
from Portuguese: If you speak slowly, I can understand you ). Dobzhansky had had the personal initiative to
study Portuguese before travelling to Brazil to prevent communication problems with his future young students
and laboratory collaborators!
The Dobhansky stay in Brazil was a complete success. Not only he could investigate with his colleagues of
Brazil many interesting cytogenetic problems in particular about the Brazilian native Drosophila, publishing many
papers on this topic, but he created a very productive climate for training a considerable number of young
scientists interested in genetics. From this original group of researchers who worked with Dobzhanskymention can
be made of Crodowaldo Pavan, Antonio Rodrigues Cordeiro, Oswaldo Frota Pessoa, Francisco Mauro Salzano,
Newton Freire Maia, A.G. Lagden Cavalcanti, Alcides Carvalho, Chana Malogolowkin, Cora M. Pedreira,

17

Friedrich Gustav Brieger, Giorgio Schreiber (see Fig. 39), P H. Saldanha, and Warwick Estevam Kerr, etc, who
played an important role in the intensive development of genetics coveringthe immense territory of the Brazilian
subcontinent!
The pioneer group of researchers facilitated the development of animal genetics throughout Brazil with
great success involving the states of Ro Grande do Sul, Rio de Janeiro, Paran and Bahia. In the following 15
years the Brazilian genetics grew so rapidly that became one of the 10 most developed all over the world.

Figure 8AGallery of selected Brazilian geneticists-. a: Andrs Dreyfus, pioneer of genetics development
in Brazil ; b: Theodosius Dobzhansky, Genetics Researcher who created a great universitarian genetics center in
Sao Paulo financed by the Rockefeller Foundation. The most important Brazilian geneticists that were trained
there appear illustrated in this figure; c: ProfessorCrodowaldo Pavan teaching genetics in Austin, Texas,
September, 1970; d: Antonio Rodrigues Cordeiro;investigating Drosophila willistoni, October, 1954; e:
Francisco Mauro Salzano; human geneticist, receiving the Prize Doctor Honoris Causa of the Costa Rica
University, May 11, 2011;

Figure8 -.BGallery of selected Brazilian geneticistsf: Newton Freire Maia, human geneticist; g: Osvaldo
Frota Pessoa human geneticist; h:Friedrich Gustav Brieger; botany geneticist; i:Alcides Carvalho, plant
geneticist; j:Warwick Estevam Kerr, bee geneticist. The picture shows a well protected beehive (including a stone
on top) to prevent escape of the aggressive African bees under investigation.

18

Due to the enormous creative vision of Estable not only admitted in his nascent Institute the
abovementioned researchers but their number was enlarged so that the Institute could cover other areas of biology.
Thus, already in the early stages of the Institute installed in 1952 on Italy Avenue had the following areas of
biology: Botany, (Wilhelm Franz Herter); Histology, (Julio Mara Sosa); Neurophysiology, (Alberto Vaz
Ferreira); Cytogenetics, (Francisco A. Sez); Biochemistry, (Maria Isabel Ardao); Microbiology, (Napoleon
Pradines-Brazil); ElectronMicroscopy, (Eduardo P. De Robertis); Neuroanatomy, (Jos Roberto Sotelo);
Biophysics, (Carlos Maria Franchi). Likewise, Estable also installed a well-equipped machine shop (Ramn
Rodriguez) and another one on electronics (Alberto Rivero) intended not only for the maintenance and repair of the
Institute instruments but also for the construction of new research instruments.

Figure9 -Buildings of the Institute (IIBCE). a:the IIBCE in 1956;(Photo M.E. Drets); b:the IIBCE in 1980.Present
site: Italy Avenue 3318.
The scientific structure outlined for the Institute wasprogressively transformed from its beginnings into an
exemplary institution for basic research and training of human resources whose number was steadily increasing,
turning it into the first basic research laboratory of Uruguay in the past century.At present, the Institute doubled the
laboratory area due to the fact that a new building was constructed and the old one was rebuilt to include modern
facilities. Comparison of Fig. 9 a, and b, gives an idea on the continuous Institute transformations.

19

Figure 10 - Two examplesof major research instrumentation available at the Institute. a: FACS B.D. cell sorting
and flow cytometer obtained through a grantof the Bank Interamerican of Development (BID) (1990) This
instrument is operated in the frame of a National Service of Uruguay.b: Transmission Electron Microscope JEOL
donated by the Japanese Minister of Cooperation in 1980.

20

Figure.11- Photograph showing part of the scientific members of the Institute working at the end of 1956. 1, Ral
Vaz Ferreira; 2, Oscar Vincent; 3, Eduardo Pagani; 4, Alvaro Daz Maynard; 5, Ada Pintos; 6, Amalia Laguardia; 7,
Margarita Prez; 8, Nadir Brum; 9, Jos Sas; 10, Omar Trujillo Cenz; 11, Br. Amatti (Italia); 12 Anbal Alvarez; 13,
Elsa Trinkle; 14, Gloria Martnez; 15, Adela Wittemberger; 16, Mercedes Rovira; 17, Nilsa Iriondo; 18, Hugo Freire;
19, Tomasa Rodrguez; 20, Jos Mara Martnez; 21, Walter Acosta Ferreira; 22, Hctor Franco Raffo; 23, Mximo
Eduardo Drets; 24, Nubia F. de San Germn; 25, Mara Isabel Ardao; 26, Clemente Estable; 27, Francisco Alberto
Sez; 28, Jos Roberto Sotelo.
Personnel circumstantially absent: Claudio Solari, Csar Galeano, Julio Mara Sosa, Ergasto Cordero, Eduardo P.
De Robertis, Carlos Mara Franchi, Walter Acosta Ferreira, Anbal Alvarez, Alberto Vaz Ferreira, Horacio Laborde,
Napolen Pradines Brasil, Blanca Ferreira, HctorSalato, Hugo Zunino, Martha Onetti, Alberto Rivero, Miguel
Fernndez, .

21

Figure 12.- The indoor garden of the Institute with tropical and palm trees soloved by Estable: "where glow
in the sun the light walls, the windows; where the sky interferes with trapezoid shape in the garden
surrounded by semicircular arches; where the air is cut around growing young palm trees as the house,
where noble ambitions inhabiting".

His great friend: Clemente Estable

Saz never hid his admiration for Professor Clemente Estable who began working from the initial moments
of founding the Institute together with Jos Mara Martnez and the great biologist Ergasto Cordero.
As an expression of his sincere affection to Estable, Saz dedicates a valuable article to his personality
(reference 199) expressing his feelings in these select few paragraphs:
This man never drowned the illusions of youth in material needs, which beat shortage making it an
incentive instead of being ruled by it, which beat impatience not worrying about success, not caring that nobody
knew him, as if life had not other horizons and other trails and like nothing it had to look forward to, working and
studying ardently and not satisfied, looking under the microscope, in Nature but not in the books, for what other
men have failed to find.
() Men, who cultivate the scientific work in our Continent have received a legacy of enormous
responsibility with the example of this master, our Estable, who does not know rest, working with passion and
deep faith on his own strengths. The story of his life will print one of the most brilliant pages in the spiritual
history of our countryland.The country should recognize eternally to its illustrious sons, as Estable, who is known

22

and respected throughout the world, thanks to the noble and generous impulse of his great talent, his ardent
enthusiasm and unwavering will

23

Figure13 -Saez and Estable in the laboratory of IIBCE in 1956.

(Photo M.E: Drets).

Figure 14 - Sez staining chromosomes in Coplin jars

in his laboratory of the University of La Plata

24

During 17 years, Sez researched and taught at the Universities of La Plata and Buenos Aires during which
time conducted much of his work.
In March l947, President Pern10, who had intervened Argentine Universities, ordered the dismissal of
hundreds of teachers whose democratic beliefs were contrary to his government. Sez reacted in open rebellion in
graduate centers against the abuses of the President and on March 17 he was fired for "reasons of teaching
convenience" from his positions at the University as Professor of the National Normal School and Chief of the
Laboratory of Cytology of the Faculty of Agriculture, University of La Plata.
Pern constantly repeating his slogan "more espardilles and fewer books", caused many researchers were
fired becomingunemployed at the time including the famous researchers Dr. Eduardo D. P. de Robertis,Bernardo
Houssay11 and Luis Federico Leloir12, both awarded with the Nobel Prize!
Troubled by this critical problem of unemployment, Sez received a telegram signed by Clemente Estable
which deeply surprised him changing the fate of his life. Estable was offering him to return to Uruguay and a
research position at the Institute of Biological Sciences, so Sez decided to move immediately to Montevideo.
Professor Saez remained in the Institute of Biological Sciences of Prof. Estable (later called Institute for
Biological Research) until his death in 1976, a period which involved 29 years of scientific work and Cytogenetics
development in Uruguay.
________________________
10- Juan Domingo Pern (Lobos, 1895 - Olivos, Argentina, 1974) was a political, military and President of Argentina.
He was vice president during the facto government of Edelmiro Farrel (1944-1946) and President during 1946-1952
and 1952-1955 periods, when he was deposed by a military coup in September. In 1973 was again elected president, but
died before finishing his term.
11 - Bernardo Alberto Houssay (Buenos Aires, Argentina, 1887-1971) was a pharmacist, doctor and Argentine
physiologist. He joined the School of Pharmacy at the University of Buenos Aires shortly before the age of 14 and
graduated in 1904. In 1911 he obtained his doctorate in medicine. He received the Nobel Prize in Physiology or
Medicine in 1947 for his discovery of the role played by the hormone of the anterior lobe of the pituitary in sugar
metabolism, the first Latin American Nobel Prize awarded in science
12 -. Luis Federico Leloir (Paris, 1906 - Buenos Aires, Argentina, 1987). Argentine medical doctor and biochemist who
received the Nobel Prize in Chemistry in 1970 for his discovery of sugar nucleotides and their role in the biosynthesis
of carbohydrates.

25

About Vocation
Vocation is the consciousness of a certain skill. Jos Enrique Rod, Motives of
Proteus, 1909

Oh thou!world of the infinitely small, that you meet all the charms and beauties of the infinitely great!
This handwritten phrase signed by Saez in 1919, on the inside cover of one of his study books Trattato
Elementare et Generale di Tecnica Istologica, recorded, in short, his profound aesthetic emotion when he was
only 21! originated when he was observing chromosomes.
Sez never forgot when he observed, for the first time, cell divisions in a bad preparation he could obtain
through the generous help of the Assistant of the Chair of Histology, Faculty of Medicine. Sez was amazed to
watch the images of mitotic figures he had previously observed only in books.
During the evenings in his lab surely there emerged, as a touch of grace, his strong inclination for all that
cell living beings represent. His life beingwas then transfigured in a fiery and urgent need to move forward in this
extraordinary infinitely small arcanum. He had discovered his vocation on the chromosomes of animals and plants!
In 1930, the famous researcher Dr. Clarence Erwin McClung13University of Pennsylvania, USA, founder
of the chromosome theory of sex determination(1902) arrived in Buenos Aires. His stay for a month was a
powerful stimulus to Saez, who was able to meet him personally after several years of exchanging correspondence.
________________
13 - Clarence Erwin McClung (Clayton, 1870 - Phiiladelfia, U.S.A., 1946) was a cytogeneticist who discovered the role
of chromosomes in sex determination studying the grasshopper Xiphidium fasciatum. His studies of the mechanisms of
heredity led to his 1901 hypothesis that an extra, or accessory, chromosome was the determinant of sex.He also made
interesting contributions in microscopical techniques including staining procedures.

26

Figure15 - Sez, (left) first Latin American cytogeneticist and Professor Clarence McClung, (right) first
cytogeneticist of the United States of America, during his visit at the Institute of General Anatomy and
Embryology, Faculty of Medicine, University of Buenos Aires, in 1933.
The same year, Sez published in the Journal of the Museo de La Plata, an extensive work in which, for the
first time in the Americas,the organization of meiotic chromosomes was investigated in four genera of South
American acrididae (grasshoppers) [10].
Sez recalls that early after his arrival in La Plata, he faced a hard time feeling himself alone and unknown
to all, away from family and friends.He suffered great nostalgia remembering the joyful living periods of the life
that he had left forever. Starting a new life, everything depended on himself. However he was able to overcome
with an eye "on a distant star fueled by the fire of an urgent and not renouncing vocation"

Sez and his friends and colleagues of the Americas

Sez was clearly a very caring person, who loved children and liked every expression of art and beauty,
which determined that he had a considerable number of friends. His colleagues appreciated him so clearly that he
was invited to participate as a speaker at numerous international conferences alternating with great geneticists.
Without exception, all geneticists from Latin America and other continents, recognized the high scientific
level of Sez,greatly appreciatinghis kindness and sincere friendship that he always showed to them.

27

Figure 16. Saz with four great geneticists: (Left to Right) Antonio R. Cordeiro, (Brazil),
Michael James Denham White (England), Bruce Wallace (USA), Theodosius Dobzhansky
(Russia and USA) and Sez atCold Spring Harbor, USA, (October 1951).

Figure 17 Sez participating in a Latin American Congress of Genetics.(1985)

28

Figure 18 - Sez with the Brazilian Human GeneticistNewton Freire Maia

(Curitiba, Paran) and ME Drets photographed in 1953 atthe popular
monumentof the La Carretasculpted by Belloni14.
The reader will notice the `unusualbut elegant feet position usually adopted by Prof Sez
trying to dissimulate the length difference of his legs (see p 4-5)
______________________
14. -The monument commemorates the system of transportation of people and goods that existed in Uruguay
during the 17th and 18th centuries. The monument was sculptured by the Uruguayan sculptor Jos Belloni
(1882 -1965).The epic sculpture depicted a covered ox-drawn wagon guided by only one horse-rider but
followed by a wild dog cimarrn extremely loyal to the horse-rider. The bronze and granite base sculpture
was first installed in a Montevideo park in 1934, and was recently restored by the artist's grandson. It
isrecognized as a National Historic Monument by the Government of Uruguay. Besides, Bellonis work La
Carreta was awarded a gold medal in Europe.

29

Figure 19- Sez and the geneticist of Drosophila Prof. Danko Brncic, Chile, visiting to the
Institute in 1959 and surrounded by colleagues from the cytogenetics laboratory.
(from left to right.): M. Drets, A. Laguardia, Sez, A. Diaz, Prof. Brncic, C. Solari and N. Brum.

30

Figure 20 a: Sez with geneticists from Latin America in Mxico City, afterhe was elected
Honorary President of the Latin American Society of Geneticsin 1974.From left to right: Prof. Danko Brncic
(Chile), Sez, Dr. Willy Beak (Brazil),Dr. Cant Garza (Mxico)Photo: M.E: Drets.

Figure 21 -Sez in his laboratory (IIBCE) in 1966 with Barbara McClintock15

(Nobel Prize in Genetics, 1983) accompanied by Fernando Dulout(Argentina),

31

and Orfeo Crosa and Glaucia Prez Mosquera of Uruguay.

It is difficult to improve on what is vividly expressed by Professors. Avelino Barrioand Osvaldo R Vidal
about the personality of Sez: (285)"Anyone who has known him agree that it is hard to imagine him deprived of such unique personality
qualities. Always jovial and cheerful, his constant good humor radiated optimism. Courteous with shameless
ladies had their lips one gallant phrase expressed with hispanic esprit. Simple and open dealing, fleeing of
formalism and protocol. Humanly sympathetic, often hiding the mistakes of others. Faithfully cultivated
companionship, more than once saw him acting as conciliator in bringing together with effort estranged
colleagues. He lavished generously and ungrudgingly transmitting regardless of time constraints knowledge
acquired through long experience. Death overtook him, despite his age, enjoying a full physical and intellectual
health. "

The Eminent Teacher

La Plata, the Universitarian City par excellence, provided an ideal environment for the intellectual work of
Sez. As a student, he founded with his then friend and later remarkable physics researcher, Loedel Palumbo16 a
Center for Teaching that they called "Einstein Institute". Just the 2 April 1925 the great Einstein visited the
Museum of La Plata accompanied by Sez, Prof. Lehman Nitsche and Einstein commented that there was a high
school with my name! Professor Loedel taught physics, chemistry and mathematics and Saez zoology,botany,
biology and English. They were then joined at the Institute by another colleague, Gonzalez, who was
commissioned to teach literature. This Institute was for several years an animated venue attended by teachers and
students.

Figure 22- a:Sez teaching a class for physicians together with ME. Drets during the
First Course of Human Cytogeneticsin the Graduate School
of the Faculty of Medicine of Montevideo in 1970. b: Giving a lecture.
__________________
15- Barbara McClintock (Hartford, 1902 - Huntington, U.S.A, 1992) was a well-known researcher who
worked in maize genetics and on many important cytogenetic problems. She obtained a doctorate in botany at

32
the Cornell University in 1927. She was awarded the Nobel Prize for Medicine and Physiology in 1983 for her
work on the genetic of transposing elements.

With his natural talent to convey not only their knowledge, but also his passion for science, Sez made his
classes and lectures especially attractive, not only for the students and the scientific circle, but for all audiences
(see list in the Publications Section and his Scientific Conference Reviews, and Teaching and Educational
Articles).Maybe that is why he created in 1940, along with two other young talented participants, a cultural
program on Radio Splendid, (Argentina) which lasted from March to September, 1946, and which was transmitted
on Monday, Wednesday and Friday reaching about 71 transmissions. The program was called "There's always
something to learn." His companions, who talked on various issues, especially sciences, were the eminent Ernesto
Sabato17and Jos Luis Lanuza18.
________________________
16 - Enrique Loedel Palumbo (Montevideo, Uruguay, 1901 - La Plata, Argentina, 1962). Uruguayan physicist who
studied at the University of La Plata, Argentina, and obtained his doctorate in 1925. Then spent a few years in Germany
working with Erwin Schrdinger and Max Planck, and returned to Argentina in 1930 where he devoted himself to
teaching. One of his contributions to the teaching of general relativity are the "space-time diagrams of Loedel", a
symmetrical representation of Minkowski diagrams
17- Ernesto Sbato (Rojas, 1911 - Holy Places, Argentina, 2011) was a physicist, activist, painter. andwriter of
Argentine He studied physics and philosophy courses at the University of La Plata, obtaining his doctorate in physics in
1938. Thanks to Bernardo Houssay he won a scholarship to carry out research on radiations in the Curie Laboratory in
Paris. In 1939 his scholarship was transferred to the Massachusetts Institute of Technology (MIT), where he worked on
cosmic rays. Returning to Argentina in 1940 he became a Professor at the University of La Plata. In 1943, due to a
personal crisis that led him to lose his faith in science, he decided to leave and devote himself to writing and painting.
He wrote several essays: One and the Universe (1945), The writer and his ghosts (1963),Apologies and rejections
(1979) among others, and three novels: The tunnel (1948), On heroes and tombs (1961) and Abaddon the exterminator
(1974).

33

Generating Prestige - General Cytology

The internationally renowned name of Sez increased considerably becoming universally known with the
publication of his famous book General Cytology written with his first disciple, E.D. P. De Robertis and with the
biochemist W. Nowinsky, edited in Spanish by "El Ateneo" Pedro Garca SA (Buenos Aires) in 1946. This first
text was revised and re-edited in 1948, 1955, 1960, 1963, 1965, 1968, 1970. In the ninth edition of 1975 the title
changed to Cell Biology (EP De Robertis, FA Sez and E. De Robertis, Jr.).

Figure 23. - Cover of the Japanese version of the General Citology by De Robertis Nowinsky and Saez
published in 1955.
This work was translated into English entitled General Cytology (E.D.P.De Robertis, W. Nowinsky and
F.A. Saez) It was followed by six editions that were published by W. B. Saunders Company (Philadelphia) in
1948, 1954, 1960, 1965 and 1970. In 1975 the title of the book in English was changed to Cell Biology including
as a new author E. M.De Robertis (Jr.) which was edited by W.B. Saunders Company and in Spanish as well for
"El Ateneo" Pedro Garca SA (Buenos Aires)
_________________________________
18 -. Jos Luis Lanuza (Buenos Aires, Argentina, 1903-1976) was a renowned journalist and writer. He presided over
the Argentine Society of Writers and the Argentine PEN Club, and was a professor at the School of Librarians of the National
Library. In 1972 he joined the Argentine Academy of Letters. Among his literary works are: The Dark (1942) Snapshots of
History (1943), Morenada (1946), Painters of the Old Buenos Aires (1961), Genius and Figure of Lucio V. Mansilla (1965) and
The Witches of Cervantes (1973).

34

This popular textbook was also translated into the following languages: French: University of Laval Press
(Quebec, Canada); Portuguese: Editora Guanabara Koogan (Rio); Italian: Editore Nicola Zanichelli (Bologna,
Italy) Japanese: Asakura Editors (Tokyo, Japan); Polish: Panstwowe Wydawinctwo (Warsaw, Poland), and
Hungarian: Akadeniai Kyodo (Budapest, Hungary). An edition in Russian in the Soviet Union and one in Cuba
was also published but we do not have information on these publications.
With the death of the original three main co-authors, the book was not published anymore but there is still a
great appreciation and gratitude for the valuable contribution that meant the use of this first cell biology text in the
biological formation of numerous generations of scholars worldwide.

The cytogeneticist translator

Visualizing the considerable lack of information on cytogenetic problems, particularly at the level of
biologists under training in Latin America, Sez decided to translate the famous books on chromosomes the The
Evolution of Genetic Systems by Cyril Darlington (1948), Chromosomes(1947)and Animal Cytology and
Evolution(1951) both by M.J.D. White. These books on chromosome problems translated by Sez were the first
ones edited in Spanish resulting with very important results both for research and cytogenetics teaching in Latin
America (119, 120, 121).

A compromised scientist
As a great geneticist, Sez was deeply interested in the mutational problems both at the chromosome and
gene level, and a great admirer of Hermann Muller19 Nobel Prize (173), who discovered the mutagenic action of
radiation in Drosophila.Before the tragic use of atomic weapons he gave lectures both in Uruguay and
Argentinaabout the genetic risks of radiations.
Considering the spreading of the terrible use of atomic weapons increased the risks of exposing severely
human populations Sez decided to start informing the governmental authorities and the people of Uruguay about
the genetic risks of radiations.
In the course of time, Sez published a series of letters in which he disclosedthe risks caused by the
indiscriminate use of radiation based on the findings of Mulleron ionizing radiations [180-182] and mutagenic
products [177, 178] capable of causing deleterious mutations or chromosomal changes in humans by incresing the
burden of abnormal genes to be transmitted to future generations. The sustained popular science divulged by Sez
through numerous newspaper articles and radio shows were unconditionally supported by his young collaborators.

35

Sez without doubt, wasa true precursor in the struggle of our genetic stability jeopardized by the irrational
use of radiation and mutagenic agents (pesticides and drugs for medical purposes). Sez wasa pioneer about this
irreducible scientific position, having a huge gravitational reason when countries are solving their energy shortages
through the use of atomic energy minimizing the risks of gene mutationand increased population frequency of
cancer of humans which likely will be exposed. (see Articles and Reports).

Figure 24.Herman Muller Nobel Prize (1946) teaching a class.

__________________________
19 - Hermann Joseph Muller (New York, 1890 - Indianapolis, USA, 1967). American biologist and geneticist
who in 1946 received the Nobel Prize in Physiology or Medicine for the discovery, in 1927, of the production
of mutations in the fruit fly Drosophila by means of X-ray irradiation.

36

ANECDOTES AND FACETS OF SEZ

Recruiting his first Uruguayan disciples
by Mximo E. Drets
One morning during the summer of 1948, two youths, Claudio L. Solari and Hugo E. Silvera, were
enjoying a few hours of rest in Pocitos Beach, Montevideo, when jellyfish werebeing brought in by the tide. They
had an exceptional curiosity and intelligence and began to examine and discuss animatedly the coelenterate
anatomical aspects when a man appeared and himself involved in the conversation. It was Professor Saez who
impressed by the knowledge and intellectual brilliance that these two young swimmers showed, invited them to
attend the course on Cytogenetics he would begin that year in the Faculty of Humanities and Sciences of
Montevideo.
Shortly after, the University Commission organized an Exhibition in honor to the Naturalist Dmaso
Antonio Larraaga, Uruguay, on the occasion of the centenary of his death (July 4, 1948) which was carried out at
the Municipal Subway of Montevideo. All biologists, researchers and enthusiats working in the natural sciences
were invited to participate and Saez presented a stand of genetic illustrations (13 Posters) including a binocular
microscope with a preparation of polytene chromosomes of the giant salivary glands of Drosophila melanogaster.

.
Figure25a, Sez and students making microscope demonstrations during the exhibition in honor to Dmaso
Antonio Larraaga20 in Montevideo in 1948.From left.to right:. H. Silvera, Sez, N. Medina, H. M. Savio, M. Siri,
C. L. Solari. b, Saez in 1948 with his first Bausch & Lomb binocular at the Institute of Milln Avenue (photo:ME
Drets).
Sez had the cooperation of students from his course, Hayde M. Savio, Anglica Soria, Mario Siri,
Claudio L. Solari and Hugo E. Silvera, who explained with great patience, especially the last two, to the common
citizens that every day crowded the exhibition, the chromosomes they were looking with the microscope, installed
by Sez, There were many inquiries about the images and their scientific meaning.

37

Accepting an enthusiastic invitation of my good friend SolariI became convinced me of the interestto visit
such exhibition.Thus, on Sunday July 11th.of1948 not only I went to visit the exhibition but personally met the
distinguished researcher Francisco Alberto Sez. Since that memorable date until his death, I became his student,
collaborator, and friend.
All that day they were showing the material,whichattracted me a lot. This relationship was definitely
strengthened when shortly after Sezgave me a copy of his Spanish translation of the book of Michael White
"Chromosomes" on whose dedication page Sez wrote, in unforgettable words, Mximo Eduardo Drets.The
young researcher under training, a souvenir from his friend(August, 1948).Adedication sentence that fascinated me
both cytogenetically and spiritually forever.

_____________________
20 - Dmaso Antonio Larraaga (Montevideo, Uruguay, 1771-1848) was a priest, politician and naturalist. In 1816 he
founded the National Library - the first public library in Uruguay, and was appointed as its first Director .He had an
ecclesiastical career which saw him become the first apostolic vicar of the province, and an oscillating political life that
culminated as a senator in the first legislature of Uruguay (1830-1834)..He expressed concern for the natural sciences
that unfolded in a myriad of written comments, in neat drawings, and important botanical collections. He made the first
paleontological observations in the country and the first descriptions of fossils. Most of his profuse work published only
starting in 1922.

38

The inspired researcher.

by Mximo E. Drets
In the 60s I was trying to start cytogenetic studies in human beings, but I had not the necessary
infrastructure or any finantialaid. Therefore,I decided to ask locally to help me financing a range of materials for
the new Human Cytogenetics Laboratory I was trying to install. In particular, I requested to the General Electric
Company SA (Uruguay) for a small white refrigerator who they agreed to donateto me. I imagemy surprise to find
that refrigerator installed in my laboratory of Human Cytogenetics which had been received on September 16th,
1966, by Sez in my absence. I noticed that the device had attached a note written by Sez in a kind of memorable
poetic prose:

Here dear I am dressed

all in white
and somewhat distressed
for not having known you;
when arriving I felt your absence
becauseI did not see you on entering your room
The welcome was given in your name,
By one so-called Papico
who, as a real gentleman
I really fell in grace
for its fine aristocracy.
All yours,
Snow-White

The absent-minded professor.

By Mximo E. Drets

In 1973, Sez and I were invited by the Organizing Committee to participate in the 2nd Latin American
Congress of Genetics to be held in Mexico City. We decided to travel together, our route being Montevideo,
Buenos Aires, Rio de Janeiro and finally Mexico. In flight, over the Rio de la Plata, and on route to our goal, Saez
noticed that he had forgotten all the slides selected for his conference. Saez fell into absolute despair. Arriving in

39

Buenos Aires he tried to phone his wife but he was unable to. After that, on our flight to Rio, I tried to calm him
down and to convince him that, with his long teaching experienceand deep knowledge on the subject, using a
simple chalkboard he would be able to present his conferenceovercoming the tremendous problem. During our
brief stay in Rio, awaiting our connection to Mexico, I finally managed to pacify and convince him that my
suggestion was the most plausible solution.
Somewhat comforted and having partly overcame the bad moment, we arrived in Mexico City and,
arrivingin our hotel and to our most extreme and stunning surprise, the receptionist hands to Sez a small package
brought by an unknown traveler containing all the famous slides! What had happened? His wife had noticedhis
oblivion, and immediately decided to go to the Montevideo Airport trying to find a passenger travelling to Mexico
and, by a twist of fortune, found him about to leave. Learning of the problem of Professor Sez,the passenger
politely offered to carry the package not only to the city of Mexico but also to our hotel one day before us! My
reaction was to sayto Sez:you really are a lucky man!

Remembering Sez
By Francisco M. Salzano
Emeritous Professor.
Universidade Federal do Rio Grande do Sul, Porto Alegre, Brasil
My first teaching experience abroad occurred between on 16 to 20 February, 1960 . I had been invited by
Prof. Francisco A. Sez to participate in a round table and to give a course of three lectures in the VI Summer
School organized by theInstitute for Advanced Studies in Montevideo. The event was held at the offices of the
Jockey Club of Montevideo and when I asked to Prof. Saez if I have to wear a formal suit and tie during my
presentation at the Roundtable he replied to me: "With that baby face, if you do not put a tie any of our brainy
Uruguayan counterparts will take you not seriously" When he introduced me at the event, wearing a tie properly,
he showed me with approval, but despite my youth he stressed that I was the President of the Society of Biology of
Rio Grande do Sul (1957-1959) , and currently was the Vice-President of such organization! During the five days
of my stay in Montevideo he was very kind transporting me and my wife all over the town in a fantastic dwarf
Renault stoppingand threatening to stop at every moment, exceptin crosses between the streets!

40

How I learned to respect the wisdom

By Eddy M. De Robertis,
Howard Hughes Medical Institute.
University of Los Angeles, California.
(This anecdote explains how Papico Sez gave me a lesson that was useful for the rest of my scientific life).
During the summers of my 8th to 10th school years I worked in the laboratory of Papico Sez. It may seem
a rather precocious start, but he knew me since I was a small child. In 1961 my project was to go to the back lot of
the Institute of Biological Investigations (which was a veritable zoo) to collect larvae of Chironomus sp. These
insects resemble mosquitoes but have red larvae. They have excellent polytene chromosomes in which the DNA
strands replicate but remain attached to each other. The process of RNA synthesis can be clearly visualized as
thickenings called puffs. My task was to makeI squashed preparations of salivary glands and to then draw the
chromosomes with a camera lucida (a systems of prisms that allows one to see simultaneously the chromosome
and the tip of the pencil one is drawing with on a sheet of paper). I would do rather nice preparations. I had found
a small rusty nail that was perfect to elicit contrast in the ferric Hematoxylin of Heidenhein that Sez used. To
encourage me, Papico would come by every morning and say: Here we have the next Calvin Bridges. After quite
a few days I asked him who was this Bridges guy, and he explained it was the Thomas H. Morgan 21 (Nobel Prize,
1933) graduate student that drew for the first time the giant chromosomes of Drosophila. In this way I, happy as
could be, applied to my designated work with the enthusiasm that Papico instilled in all of us.
One day I obtained a preparation of polytene chromosomes much, much better than we had ever seen.
Since I was drawing in the lab, someone noticed what I was drawing and asked to take a look in the microscope.
Very soon there was great excitement and everyone came to look at my perfect chromosome preparation.
Papico arrived last, took a look and said: Take a photograph right away. I did not know how to use the
photographic camera of the Leitz microscope, so therefore the task had to be done by the researcher (later a great
pathologist) that was my advisor and knew how to do this. It was five oclock. The young researcher sugested
meleave the slide in the refrigerator until the next morning and then take the picture. Papico looked at me intently
and asked what thoughtwe should do. Finally I pronounced myself: Yes, it is best if we take the photos
tomorrow. Papico went into his office silently, but from the look in his eyes I could clearly see I had disappointed
him.

41

The next morning my great slide had dried out in the refrigerator and was ruined. Nevertheless I examined it under
the microscope and, looking at it, realized why it had been such a lovely preparation. The polytene chromosomes
of this particular larva were at least four times wider than in normal Chironomus. When Sez arrived in the lab he
examined my ruined slide and, without saying a word, he limped into his office. A short time later he returned with
an offprint of a publication by the great Brazilian cytogeneticist Crodowaldo Pavn (Campinas 1919. Sao Paulo
2009). He had shown that in another insect infection by protozoans could cause the giant chromosomes of larvae
to continue replicating their DNA, generating truly gigantic chromosomes.
I had obtained the only case of continued DNA replication in Chironomus. As you must realize by now, I
never again found a preparation like this one. It was thus that I missed my first scientific publication to please a
younger man with whom I worked daily, rather than follow the advice of a man of great wisdom.
To this day, I use this personal anecdote to explain to my students why we must document experiments
immediately, never waiting until tomorrow.Papico Sez trained many scientists, creating a great school of
cytogenetics in Uruguay. As so many others, I am forever thankful for his having introduced me to scientific
research.
_____________________
21- Thomas Hunt Morgan (Lexington, 1866 - Pasadena, U.S.A., 1945). Zoologistand geneticistcarried out important
researchon heredity working with the fruitflyDrosophila melanogaster.In 1933 got the Nobel Prize of Phisiology or
Medicinefor his discoveries on the role played by chromosomes in heredity

42

Guidance for young researchers

by Nstor O.Bianchi,
Director, Multidisciplinary Institute of Biology and Experimental Sciences and Multidisciplinary Institute of Cell
Biology (IMBICE), La Plata, Argentina

It was the beginning of 1962, perhaps February or March. A few months before, in December 1961 I had
returned to Argentina after finishing my first external fellowship at the Columbia University of New York where I
was trained by Melvin M. Grumbach and Herbert Taylor in the techniques of human blood cultures, chromosome
labeling with tritiated thymidine and the use of autoradiography to analyze the pattern of DNA synthesis during
chromosome replication. Then, the Authorities of the Scientific Commission of Investigation of the Province of
Buenos Aires (CICPBA) decided to give me space, facilities and a grant to establish a cytogenetic laboratory to
continue in La Plata the research I was carry on in New York. Facing the fears, doubts and tribulations of the task
ahead of me, I suddenly decided to visit Francisco Saez in Uruguay for advice and support.
I had met Sez sometime before during a Symposium held in Argentina in which I had delivered a
scientific presentation and Sez a conference. At that event I introduced myself to Sez, I congratulated him for his
inspiring conference and we shook hands and exchanged greetings; in summary just a brief encounter between one
of the most, or perhaps the most outstanding Latin-American cytogeneticist of that time and a newcomer to the
chromosome discipline. Having taken the decision to visit Sez my anxiety overcame my common sense: I
gathered my few savings and travelled to Montevideo without a previous announcement to professor Sez. A soon
as I entered in the Institute Clemente Estable I realized that I had no appointment and not enough savings to cover
the hotel and meal expenses for more than a couple of days. Moreover, I was sure that the giant of the cytogenetics
very likely would not remember our fleeting exchange of good wishes in Argentina. What a surprise! The
receptionist, after consulting Sez, gave me the instructions on how to reach his office telling me that the professor
would receive me. In a few minutes, I was knocking his door and when, once again. I met the
Professor.Surprisingly, I discovered that he not only remembered me, but he also remembered my name and my
presentation to the symposium. We talked for several hours. Saez was always focused, he addressed me as a
colleague and not as a professor to a beginner.He analyzed and discussed my ideas and he appeased my fears and
doubts. When I entered in his office Sez was an admired cytogeneticist, when I went out of his door Sez was an
admired cytogeneticist and an admired friend. His suggestions and advices were useful to carry on my project in

43

Argentina, yet, still more important, the treatment he gave me during my visit taught me how my behavior should
be with students and colleagues in the future.
In Latin-America Papico was the friendly nickname gave to Sez when he was a child. Moreover, since
Saez was very fond of visiting the laboratories of other Latin-American cytogeneticists I used to call him Papicus
migratorius it is a pity that the passing away of Papicus implied as well the pass away of one of the most
representative specimens of that rare and endangered species of great and lovable men.

Cyrcadian Rhythm and Sez Forgetfulness .

by Nstor O.Bianchi,
Director, Multidisciplinary Institute of Biology and Experimental Sciences and Multidisciplinary Institute of Cell
Biology(IMBICE), La Plata, Argentina.
Francisco Saez, Crodowaldo Pavan, Juan Hunziker and I were part of the small delegation of SouthAmericans that in 1968 were invited to Tokyo as speakers or to chair sessions, round tables or symposiums in the
XII International Congress of Genetics. All South-Americans, including ourselves, suffered the sleep-day
vigil-night resulting from the +10-12 hour difference between Tokyo and our fatherlands. We drowsed during
the conferences and other diurnal Congress activities and at night we chatted, drank infusions and reveled
enhancing the discomfort of not adapting to the circadian rhythm changes in a futile attempt to get the circadian
rhythm adapted to us instead otherwise. Thus, in daytime we experienced changes in the appetite and other habits,
some disorientation in time and space and we looked and were more absentminded than usual. In one of those
days in which the circadianism severely mistreated us, Don Francisco Sez felt the urgent need to go out
shopping to buy some presents for his family and he asked Juan and me to join him. We picked up a taxi up to the
Ginza22area where we wandered hours visiting shops with plenty of potential gifts too expensive for the Papicos
wallet,or that pleased his budget but not his taste. Finally, more from exhaustion than conviction Papico Sez
bought a pair of local objects and that then packed for gifts.
Once again we took a taxi shared the fare and entered the hotel and then, when searching in our pockets for
the room key, Don Francisco discovered that his wallet where he kept the yens to pay the hotel expenses and meals
was lacking. Despair sheer despair! After exchanging memories we retrospectively assumed that the missed
money purse had been forgotten on the shop counter where Sez had bought the presents Once again the
circadian runaway had played a trick on us, especially since no one remembered the name and address of the
business and Don Francisco had also kept the purchase receipts in the forgotten wallet. From 9-to 11 that night we

44

held an emergency meeting in a Crodowaldo Pavansroom. After using our best hours of nocturnal lucidity we
concluded that the only way out was to share a room and the money withDon Francisco, DArtagnan and his three
musketeers: Juan, Crodowaldo and Nstor. Immediately we went down to the hotel reception desk to make the
arrangements and then surprise!!
The shop owner where Papico had bought the gifts found a hotel card in the wallet and sent it back with the
money intact with a messenger in bicycle who spent two hours for the trip... The same two hours the emergency
meeting had lasted. Don Francisco displayed the distractions of people who only think about transcendent things
and also of those who suffer from circadian rhythm disorders.
________________________
22 - Ginza is a Chuo district of Tokio where are an important number of stores, restaurants and
cafeterias. It is known as one of the most luxuries commercial areas in the world-

45

Figure 26The four authors of the anecdotes on Prof. Sez. a: E. de Robertis in his laboratory
at the Howard Hughes Medical Institute, University of Los Angeles,California; b: M.E. Drets
and N.O. Bianchi in Houston, USA, returning from their participation in the Somatic Cell
Genetic Conference developed in Galveston, Texas, in October 28-30, 1970; c: F.M. Salzano
in his Department of Genetics, Porto Alegre, Brazil, 1975.

The actor on stage

(By Prof Juan Hunziker23, Argentina)

Professor Hunziker (1976) (288) described Sez as "not the kind of reconcentrated, shy and reclusive
scientist and suffering abounds whenever trauma must climb to the stage" Although it was never said and they
were good friends - Prof. Hunziker considered that Saez "loved the limelight" consequently, he could participate at
scientific meetings often transcending the purely scientific and academic personality. In 1968, at the end of the
International Congress of Genetics held in Tokyo, Japan, several geneticists from different countries sang songs
from their countries. Suddenly, without any request or any begging from anyone, Sez, determined and confident,
went to the stage and taking the microphone interpreted in a clear, pleasant voice, showing his intense love for the
land and the River Plate idiosyncrasies, sang the tango "A media luz" which was warmly celebrated (288).

46

Figure 27 -Professor J.H. Hunziker

Beloved friend of Professor Sez.
23. Juan Hctor Hunziker. (Buenos Aires 26/08/1925
17/03/2003) Licentiate in AgronomicalEngineer, University of
Buenos Aires (1949); Master in Genetics and Ph.D. from the
University of California; Researcher in Genetics and Evolution;
Professor, Faculty of Exact and Natural Sciences, Univ. of
Buenos Aires; California, and Washington (USA). Investigator,
Inst. of Botanics, Minister of Agriculture and Cattle (19471958).Member of the National Council of Scientific and
Technical Research.Crops, Ecology and Genetic Resources
UnitConsultant.(FAO), Rome (1975); Member of Science
Society of Argentina, Botanical Society of America (USA),
International Association for Plant Taxonomy (Holland) and
Botanical Society of America. Honor Diploma, University of
Buenos Aires (1948); "Eduardo Holmberg Award" (1966);"
Konex Platinus Award in Genetics and Citology" (1983) and
"Len Cherny Award" (1983).He published more than 100
research papers and three books.

47

SCIENTIFIC CONTRIBUTIONS
(Critical Racconto)
The reader is encouraged to look, fora morecomplete information, the Section Additional References
(pages 129-130), particularly forthose references briefly mentionedin this text.

Cytological methods for chromosomes

Throughout his life Sez performed numerous investigations in various biological materials, both animal
and vegetalstarting in 1925he introduced multiple technological improvements applied to the study of nuclei and
chromosomes by microscopy [1,2].
In 1929 he developed the method of ferric acetic carmine [8] for chromosome staining, a technique
subsequently used for decades by numerous cytogeneticists. In 1939, Sez published a study on the Feulgen
reaction which was the first in South America paying attention on the technical aspects of chromosome staining
[40].

Figure 28.-Sez drawings of meiotic chromosomes of Schistocerca paranensis for his work of 1930 on
Orthopteran chromosomesSchistocerca paranensis,(1 and 2), Elaeochlora viridicata(3 and 4),Diedronotus
discoideus (5 and6) [10]these made using the Camera Lucidaand Indian ink.Note that the different tonalities
appearing in the drawing indicates that black color the chromosomes were closer to the observer eye and the
chromosomesin gray color were at a lowerimage level.(see also the metaphases appearing in the cover of this
book)-

48

His slides were always the product of a true artist-artisan obtained mainly from the microtome cuts made at
the time, which were firmly adhered to the slide using the "Mayersalbumin " (a personally prepared mixture of
fresh egg white and glycerin).
Sez also used hematoxylin for chromosomes staining and counterstaining the cytoplasm with Safranin or,
alternatively, with Crystal Violet or Gentian Violet methods that required clove oil to remove patiently the excess
of dye to achieve a precise differentiation obtaining the most splendid staining hues forprecise microscope
observations. No doubt, these technological procedures were part of the secret that allowed Sez to arrive to such
highly precisekaryological diagnoses.
In 1950 Sez devised a technique for studying the long somatic chromosomes usually observed in plants
[184]He used the procedure particularly in the karyological analysis of Brodiaeauniflora [50]. In this case, Saez
used the method of "crushing" or squashing to produce the preparations with extended wellall the chromosomes
in the cell.
During his research work, Sez constantly perfected a number of cytological staining procedures for
accurate chromosomes observations of various materials as is mentioned in the particular areas he studied always
obtaining chromosomal preparations with beautiful images [19, 75].

Orthoptera
In 1924 Sez began his research on chromosomal behavior in relation to inheritance, sex determination,
phylogeny and evolution to establish the evolutionary cytotaxonomic structural interrelationships in several
species of Orthoptera insects of South America.
It is really admirable that despite his deep interest in studying chromosomes, Sez was devoted primarily
with high scientific rigor and critical essays to all cytological procedures available at the time (1912-1926),
discussing all the technical steps (fixation, paraffin embedding, microtome cuts, coloring with safranin, gentian
violet, iron hematoxylin, picro-indigo-carmine Cajal, among others [4] in order to achieve high-quality
chromosomes, following the advice of McClung (Fig.10) with whom he had frequent correspondence and also
with his mentor Prof. Miguel Fernndez.
Whereas scientific loneliness and the absence of colleagues at that time that were interested in
chromosomes, Sez began his cytogenetic studies at the Museum of La Plata in an ingenious way. First, he had the
clever insight to choose for his chromosome studies, a uniquely suited biological material as were theOrthoptera,
due to the large chromosomes size and the remarkable beauty of thecytological images that could be obtained with
this material [5,6]. With particular intellectual clarity he started, then, in the detailed analysis of the morphology of
the chromosomes [21],orientinghis studies to the chromosomes of the Orthoptera of South America
[22].Simultaneously, Sez studied in a detailed way the literature available at the time covering the main findings
in the subject published until 1927 [3].
Although Sez certainly lacked laboratory material, in his own laboratory he was able to publish in 1930,
his historical work on Orthoptera [10] obtaining a considerable impact of which was, at the time, of such
international magnitude, that the famous English cytogeneticist Cyril D. Darlington24 used the Spanish title to cite

49

the first great work of Saez in his monumental book "Recent Advances in Cytology" published by JA Churchill
Ltd. (First ed.) London, 1932 and in the 2nd.Ed. in 1937 (pp. 534).

50

Darlington made several references to Sezs work which appeared in his book. In particular in Table 2 on
"Organisms with chromosomes of extreme size" section Animals mentioning Schistocerca paranensis (and other
South American Acrididae) (p. 84) as well as in the referencetoChromacris miles et al.in Table 14:Classification
ofOrganisms according to the distribution of chiasmata in metaphase,column 1:Chiasmata locatedor semi
located/Minimal terminalization " and in the reference that appears included in full in Appendix IV: Bibliography,
(p. 534).

Figure 29 The famous cytogeneticist Professor Cyril Dean Darlington (F.R.S) (1948)
Director of the John Innes Horticultural Institute, England.
Photo: National Portrait Gallery, London. (Background re-processed).
In 1931 [20] and 1935 [27] Sez published two papers on chromosomes of Orthoptera species of the Genus
Aleuas (Aleuas vitticolis and Aleuas lineatus) describing the formation of multiple chromosomes by translocation
of autosomes which generated great international interest and that can be considered as true classical papers.
Saez continued expanding his studies on the chromosomal constitution of several Orthoptera from South
America, Schistocerca, Elaechlora Diedronotus and Chromacris and of the subfamilies Acridinae and Truxalinae
as well as the chromosomal complex of orthoptera of the Ro de la Plata [11, 12].
_________________
24- Cyril Dean Darlington (F.R.S.) (Chorley, 1903 - Oxford, England, 1981) was a biologist and geneticist
whose research on chromosomes influenced the basics of hereditary mechanisms. He discovered the
mechanism of genetic crossingover, its role in heritage and its importance for evolution.

51

In 1938-39 Sez performs other karyological studies on Schistocerca paranensis[36]. With Rosa Gershanik
Sez published in 1939 a note on the cytology of Trygonophymus arrogans[42] and in 1950 with his first two
Uruguayan disciples Silvera and Solari, describe the cytogenetic structure of Schistocerca cancellata[52]
analyzing some cytogenetic features such as location, frequency and terminalization of chiasmata, as well as the
recombination rate which were considered key factors in the genetic system for understanding the inheritance, the
rithm and the direction of the species evolution or within its intercrossing group, aspects in which he emphasized
in a further work [56]In 1955 Saez examined the chromosome interspecific variation in the genus Scotussa[60] In 1956 he
published in Nature an extensive list summarizing his research on the chromosome structure and the sexual
constitution of a considerable number of species of Orthoptera of the southern hemisphere [61, 62]. Employing the
same analytical criteria mentioned in the paper on S. cancellata[52], he studied,with Amalia Laguardia (1959), the
cytogenetic structure of Laplatacris dispar [82]. Together with Claudio Solari he extended his studies on the genus
Scotussa (1959)describing the chromosomes of three new species [74] and in 1960 the chromosomes of genus
Scyllinops [79]. With Alvaro Diaz (1960) he described the sex system neo-Xneo-Y in Xyleus laevipes [78]. With
Glaucia Perez-Mosquera he studied in 1964 the evolution of the sex chromosome of Schistocerca infumata
showing in this paper that the X chromosome follows a differential alocyclia during spermatogenesis, either by
condensation or through spiralization degree [87].

Figure30- .Diagram of the gradual process of chromosome

heterochromatinization as Sez hypothesized (83).
One of the most interesting contributions from Saez was his extensive analysis of the evolution of the
sexual system Neo -X Neo -Y in relation to theheterochromatinization gradient [83] an hypothesis published in
1963 which afterwards he developed and deepened further in 1968 [94] (Fig. 30). The importance of the
observation was that when the neo-X-neoY system (X-autosomal-translocation) occurs differentiationin the
chromosome incorporated in the neo system, in which the chromatinof one of the autosomes is gradually
transformed into heterochromatin, appearing as positive heteropycnotic segments in the neo-Y chromosome during
the meiotic prophase.

52

The sex-determining mechanism is provided by the system X0 in some species, a marked change from the
X0 to XY type is produced [105]. This mechanism is originated by a translocation between anodd sex
chromosome X coming from an autosomepair, that sometimesform a new kind of sexual mechanism in the genetic
system of the species [65]. Saez issued his heterochromatinisation gradient theory as an explanation of successive
differentiation steps leading to the eventual genetic isolation of the Y chromosome [106].
Manuel Diaz studied in 1968 with Sez, the interesting behavior of the DNA synthesis of Dichroplus bergi
by autoradiographic methods using tritiated thymidine [93]. They observed that a special pattern of late X
chromosome replication that relates the chromosome X with the neo-Y was genetic restricted to the male line
becoming heterocromatinized. The autoradiograph showed that the process of DNA replication was parallel to the
heterochromatisation and that, in both processes, the original X chromosome and neo-Y were late replicating.
Moreover, the homologous autosomal regions of the neo-X and neo-Y were not isolated and paired during
meiosis, showing a pattern of reverse replication. Besides the homologous autosomal part of the neo-X and neo-Y
that was not isolated and paired during meiosis showed a reverse replication pattern. The facts observed by Sez at
the microscope level led him to propose that the heteropycnosis of the chromosome neo-Y could consist of a
regional differentiation of the chromosome, with a characteristic chronological behaviour with its own cycle of
coiling.They thought that it was not exaggerated to admit it should have a defined and constant function, a
hypothetical view of Sez that turned out very interesting resulting from his observations and deep knowledge on
the karyotypes of these orthoptera which was then confirmed by the autoradiographic observations mentioned.
As a culmination of his research on the Orthoptera cytogenetics, Sez discovered in 1956 [63] the
extraordinary case of the grashopper Dichroplus silveiraguidoi, whose study wasextended in 1957 [69],carrying an
extreme and unique karyotype resultingfromthe decrease of the chromosome number from 2n ()=18/22/23 a
kariotype supposedly an ancestral characteristic of most acrididae, to an species of 2n = 8 with achange in the sex
system determining from type X0 to XY [70] (Fig. 31).
This rare insect was found near the Batov Hill, Department of Rivera, Uruguay. An outstanding feature of
these grasshopers with eight chromosomes is the fact that they do not fly, and only can jump, because they are
brachypterous mutants. So, they are confined to a limited living area, (see Fig. 31). One wonders: what could have
happen in the area of Cerro Batov to cause such drastic evolutionary change?
This remarkable finding increased the interest of Sez on the problem which led to a series of studies on
the cytogeneticsof the genus Dichroplus [95] carried out with his collaborators, trying to clarify the origin of the
chromosomal changes andkaryotype evolution of the different species. In this regard, Sez and Prez- Mosquera
published (1969) [97] a detailed cytogenetic research on the evolutionary mechanisms of the insect, proposing that
the reduction of the chromosome number could have been produced by two mechanisms: (1) centric fusion or
tandem fusion between non homologous chromosomes or, (2) structural rearrangements by pericentric inversions.

53

Figure 31-The eight chromosomes pair of

Dichroplus silveiraguidoi.The fourth pair
corresponds to the neo-X neo-Y elements.
Above: The Batov hill.
The localization, structure and behaviour ofheterochromatin C (constitutive) during the meiosis of
Dichroplus silveiraguidoi were investigated in detail by Cardoso, Saez and Brum in the years 19731975[109,113,115], who detected an unusual behaviour of the heterochromatic blocks located in the synaptic
region of the sex bivalent(Neo-X Neo-Y) as well as a heterogeneous configuration of the C blocks characterized
by the existence of small positive granules interconnected by euchromatic filaments. In addition, they observed a
large amount of heterochromatin distributed in the form of granular material and centromeric and telomeric
centromeric blocks along the autosomes,all of which the authors assumed must have had an evolutionary
significance in this species.
With Lucy Peixoto, Sez return to analyze a population of Laplatacrisdispar [99]an easily collectible
grashopper in Uruguay, which he had studied from its beginnings as cytogeneticist. At that time (1970) Sez
associated with Knapper studied the cytogenetics of the most frequent annelids species of the genus Pheretima of
the family Megascolecidaeto contribute to the taxonomic clarification, confirming his interest as cytogeneticist in
other biological materials [98].
Sez emphasizes again thisattractive subject and, in 1974, Cosen and Saez observed in Dichroplus vittatus
the sexual system of the type neo-X neo-Y also found in Xyleus laevipes, Trygonophymus arrogans, Dichroplus
bergi, D. obscurus and D. silveiraguidoi [114].

54

Sez never ceased to study the structure, behavior and chromosome evolution of D. silveiraguidoi and thus
produced a new manuscript with Prez- Mosquera on these issues that was accepted by Geneticaon March 1st.,
1976, just few days before his death, having been published in 1977, so this research became his posthumous work
[118].
In this paper, the authors studied again the chromosomal changes occurred in the insect according to Saez
hypothesis published in 1968-71 but presented in a diagrammatic form. They described the stages of the meiosis
performing a detailed description of the X chromosome. The interesting thing about this structure is that presents
heteropicnosis of the sex chromosome located at the centromere distal segment while the proximal shorter segment
is isopicnoticso that the authors suggested the existence of a translocation between the original X chromosome and
an autosome. Another aspect of interest of this work is that they presented, for the first time, a comparative
cytophotometric analysis in spermatids of four species (D. elongatus, D. silveiraguidoi, D. bergi and D. punctatus)
a matter of great evolutionary importance but, unfortunately, this research could not be continued.

Figure 32. - Prof. Saez and Drets watching the first cultures of
Drosophila willistoniobtained at the IIBCE in 1956.

Amphibians
The anuranamphibians have always constituted a group of great cytogenetic interest for the possibility to
investigate their phylogenetic and evolutionary relationships as well as to studytheir sex determining mechanism.
In 1934 Sez [23] and Sez et al [24] published several studies on the chromosomes of the toad Bufo
arenarum (Hensel) analyzed by various methods of fixation (Minouchi variant,strong Fleming, diluted Fleming,
and Allen and La Cour procedure) presenting the first illustrations of spermatogonial metaphases and images of
tetrads observed during meiosis andhand drawn with the Abbe Camera lucida. In both studies, they observed 22

55

and 11 chromosomes respectively, concluding that the chromosomes of the toads were all metacentric and that the
formula of the material was 22 diploid chromosomes and 11 haploid chromosomes.
Sez was already, at that time, an expert cytologist in allprocedures for staining chromosomes (e.g.,
Heidenhaim hematoxylin, crystal violet, Feulgen, supravital methods with methylene blue). With this extensive
technological knowledge, he decided to study in detail together with Rojas and De Robertis the meiosis of South
American toad Bufo arenarum (Hensel) [29, 30].
Detailed research was published in 1936 [31, 32]showing that amphibians did not presentmorphologically
differentiated sex chromosomes contrary to the previously held view by the Japanese school and several other
authors. It was only possible to carry out such an important discovery, which at that time was opposed to the
commonly held view on the sex chromosome constitution because of the detailed cytological observations made
by Sez and his collaboratorsand the excellent and numerous cytogenetic images that documented their work.

Figure 33 - Early stages of meiosis of Bufo arenarum (Hensel) as appear illustrated in the work of Saez et al.
published in 1936 [32].Note the high quality and detailed images drawn.

It remains admirable in a superlative degree, even today, the high quality achieved through hand drawings
patiently made with Camera Lucida using progressive dilutions of Chinese ink in his drawings to differentiate tiny
and subtle structures stained with different cytological methods, a really laborious almost artistic work. All meiotic
stages were carefullyexamined and illustrated highlighting faint images of the earliest images including the precise
synaptic leptotene meiotic"bouquet" stages (Fig.33).It is no exaggeration to say that even the most accurate
photomicrography system perhaps could not beat the quality of these remarkable hand drawn illustrations.

56

We transcribe some conclusions of the authors that are really historical about this discovery about the
problem of the existence of the sex chromosomes in Anurans.
"In Bufo arenarum 22 metacentric chromosomes(twelve large and ten small chromosomes) were
observed during metaphase, being the chromosomes arranged radially in the equatorial plane. In
metaphaseI11 tetrads were observed. We found one differential tetrad similar to the sex chromosome
described by the Japanese authors. This element, however, was not assigned the designation of sex
chromosome, because it is not certain that is the only chromosome complex that behaves this way. The sex
chromosome of anuran amphibians has not been identified sincethey represent as just tetrads whose
peculiar behaviour can affect different chromosomes of the complex. In fish and amphibians the sex
chromosomes arein a morphologically undifferentiated state. It cannot inferred thus, that the male sex is
homozygous in anurans simply because theydo not show cytological differences. The sex chromosomes
have not been identified with certainty with respect to the other autosomes"
In adition, Sez analyzedin 1937 [33]the cytology of Atelopus stezlneri (Weyenbergh.) (Amphibian:
Anura) showing that they had 22 chromosomes, a fact well demonstrated studying the animal meiosis.
Saez takes up the theme and published a following paper in 1939 [37] on the problem of sex determination
in lower vertebrates. Sez did a fine critical study on the problem identifying the cytogenetic mistake about the
existence of sex chromosomes mainly sustained at the time mainly by the cytogenetic Japanese School. [Iriki S,
1930; Makino1932; Minouchi & Iriki; 131; Sato 1933)
This superb contribution on this problem by Sez was an exemplaryand detailed analysis aboutthe
determination of 22 diploid chromosomes, which deserves tobe mentioned in more detail because of the historical
nature of the contents of this work.
At that time this was a matter of great interest to cytologists, when it came to determine the sex in fish and
amphibians, particularly after the work of McClung (1902)
It was observed in Lebistes reticulatus (Vaupel J, 1929) that a primitive condition of the mechanism of the
sex chromosomes in which a high degree of mutability was observed suggesting that the XY condition was in a
"nascent" stage (Winge 1932). Winge noted inLebistes that not only that the sex chromosomes should be
considered as part of the autosomes but that it was possible to detect the transformation of the X chromosomes into
autosomes and that autosomes into X or Y chromosomes and that, under certain conditions, the difference between
the X chromosome or the Y was of a quantitative nature, being unable to confirm the presence of sex
chromosomes morphologically distinguishable from the autosomes in fish.
Aquarium enthusiasts have always found that, in some species, the number of females is higher than males
as it is frequently observed in Lebistes reticulatus, Xiphophorus helleri (swordfish), Xiphophorus maculatus
(platis),Mollienisia sphenops (mollies).Sexual reversions from males to fertile females, or the contrary direction
occur, showing that there is no a fixed sexual determinism as in higher vertebrates (Innes, 1954). It is a classic fact
that in frogs sexual inversions are also observed. [Witschi E (1931 a, b, c)

57

The Japanese authors had observed a V-shaped chromosome during the first meiotic division which was
considered the sex chromosome.This fact was not confirmed,since Galgano (1933 a) and Witschi (1929) had found
chromosomes in variable numbers, from 2 to 5, and of different size thatwere named pseudosexual chromosomes
as well the existence of variability in the shape and size of tetrads in cells of the same animal. These events were
consistent with what Saez et al. found in Bufo arenarum[31, 32]in Atelopus and Leptodactyllum where they
established the tetrad with a shape of a V, sometimes is observed and sometimes not, and that were of different
size in different cells the same individual, so Sez called it the "differential tetrad " [38].
In 1939 Sez conducted a study [39] on another amphibian with similar karyotype to the previously studied
Bufonidae, the Atelopus stelznery (Weyemberh) Investigating the origin and evolution of the annular tetrads in
amphibians with a penetratingjudgment, hecomparedhis observations with the literature of the time, and concluded
that sex chromosomes did not exist in anura.

The phenomenon of polyploidy in amphibia

Two decades later, Saez and Nadir Brum (1959) described the chromosomes of Odontophrynus
americanus, Ceratophrys ornata and Odontophrynus cultripes, [73], in relation to their geographical distribution
in South America, finding karyotypes of 22, 42 and 44 chromosomes [81]. Hunziker (1976) [116] argues that these
studies were the first ones published in the international literature describing the phenomenon of polyploidy in
amphibians (Fig. 34).

Figure 34- Polyploid metaphases of anuran amphibians

first described by Saez et al. in 1959.
In 1960, Saez and Brum published in Nature [77] a broad overview of South American amphibians
pointing out that the ancestral diploid number was 22 metacentrics [80, 81, 84], confirming the observations of
Galgano (1933b).

58

Afterwards, Saez and Brum (1965) described the karyotypes of other Odontophrinus (O. occidentalis and
O. cultripes of Argentina and O. americanus, O. americanus of Uruguay [88]. Subsequently, Saez and Brum
extend their observations in a note on the karyological variation observed in anurans of the genus Odontophrynus
[91] finding in Ceratophys ornata (Uruguay) 92, 96 chromosomes and 108 chromosomes in Odontophrynus
americanus concluding that polyploid number of chromosomes found in O. americanus might indicate the first
stage of the evolutionary process of separation between forms found in Uruguay and Argentina.
Polyploidy, as an evolutionary factor, was also described in vertebrate species by researchers from Brazil
who published a series of papers on the subject confirming the existence of the phenomenon of polyploidy in
amphibians [Beak Beak ML &Beak W 1998) Beak, ML, Beak W, & Rabello -Gay (1965); Beak ML, Beak
W (1998).
As a result of his research on frogs, Brum and Saez published in Experientia(1968) [92] a list on the
chromosome composition of 23 species of Leptodactylidae (amphibiananurans) not finding sex chromosomes
cytologically diffentiated in these frogs. This list was completed in 1971 [101] describing the chromosome
complement of South American Bufonidae,confirming the absence of sex chromosomes.
Brum and Saez also studied several species of Hylidae [96] and Barrio, Sez and Barrio Sez and RinaldiChieri [100] studied the somatic chromosomes of a South American apodalamphibian, Chthonerpeton
indistinctumin peripheral blood establishing a karyotype of 2n = 20, notfinding sex chromosomes. In this species
was difficult to establish the meiotic sequential stages, although images wereobserved with different degrees of
coiling and two to four chiasmata in late leptotene. In particular, the presence of sex chromosomes was not
ascertained.The resulting karyotype number was lower than in Ichtyophis glutinosusand in Uraeotyphlus
narayani,species that carry 42 and 36 chromosomes respectively, suggesting that these three legless amphibian
species belong to three different families. On the other hand, the authors considered that these different
chromosome constitutions would reveal an expression of the evolutionary process independent of the Apoda
karyotype in relation to the Urodeles and Anura, since in the lattermicrochromosomes have been observed, an
aspect of taxonomic and phylogenetic importance.
With WaldoVenegas, Sez studied a small toad Rhinoderma darwini (2n = 26) by determining the chiasma
frequency, the coefficient of terminalization and the recombination index.These dataindicating that the
independent segregation of some chromosomal segments is restricted. The authors discuss, on the basis of these
observations, whetherRhinoderma darwini belongs to the subfamily of the Leptodactylidae,eventhough variable
karyotypes from 18 to 30 chromosomeshad beenobserved,which indicates that is a very heterogeneous and
complicated family [107]
In 1973, Brum and Saez [108] examined the chromosomes of eight species of South American toads (Bufo
marinus, B. paracnemis, B. ictericus, B. arenarum, B. spinulous, B. fernandezae, B. d'orbignyi and B. crucifer)
establishing a karyotype for all of them 2n = 22 without detecting bivalents with the characteristic morphology and
behavior of sex chromosomes confirming, once again, the discovery mentioned at the beginning of this section.

59

Sumarizing, during the period 1959-1973 Saez and Brum investigated about 80 species of South American
anura amphibians, which clearly indicates their great interest in this topic,that was driven by the problems of the
lack of sex chromosomes and the presence of polyploidy in these animals [76].

Plants
In 1943, Sez and Nez [47] studiedthe somaticchromosomes ofSorghum almum (2n = 40)suggesting that
ithad beencaused by fertilization of a normalgamete haploid of S. halepense (2n = 40) with a diploid gamete of a
species of the Arundinacea subsection[51].
The interesting history of this elegant work was that Sezs attention had been caught by the external
appearance of black sorghum (Sorghum almum) [57]that had some similarities to other sorghums, for
instance,Sorghum halepense, known as sorgum of Aleppo. On the other hand,Sez supposed not improbable the
existence of some genetic relationship, unlikely with other sorghums,such as Sorghum sudanense or Sudan grass.
The sorghum of Aleppo (Sorghum halepense) had 40 chromosomes and S. sudanense 20. As an hypothesis,
he thought it could have happened a cross between sorghum Aleppo and Sorghum sudanense whose gametes
carry20 and 10 chromosomes respectively, and therefore the black sorghum should have a karyotype of 30
chromosomes.
However, contrary to expectations, Sez by cytogenetic analysis revealed the existence of 40 chromosomes
and not 30 as it was expected. Such finding complicated the problem of the origin of the black Sorghum. After a
careful study of several hundred plants of this sorghum Sez concluded that the black Sorghum was a new
perennial species with 40 chromosomes supposedly caused by fertilization of a normal gamete with 20
chromosomes of Sorghum Aleppo and with an exceptional gamete originating at ramdom from a sorghum of the
type of kind of sorghum sudanense,whichinstead of having 10 chromosomes.carrieda set 20chromosomes(Fig.
35).
Simultaneously, with the first publication of the work of Sez and Nez a paper by Parodi (1943)
appeared based upon of taxonomic studies, who named this new species with the name of Sorghum almum based
on the results of Saez.
Some years after, Sez received the information that Dr. Randolph,from Cornell University ( USA), had
conducted an experiment obtaining, by synthesis, a sorghum virtually identical to the black sorghum (S. almum).
based on the results of Sez. Randolph made a cross between S. halepense and the autotetraploid
S.sudanensewhich produced gametes with 20 chromosomes.
This synthetic species became the definitive experimental confirmation of the Sez hypothesis on the origin
of the new sorghum. It is highly recommended toread a newspaper article on this extraordinary work that Saez
published in 1974 on the origin of a new species (215).

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In 1949 he published the first studies of plant karyotypes made in South America with modern squash
techniques on Hypochoeris [46, 49, 53] and Brodiaea uniflora [50]. Sez studied the species of the Hypochoeris
group (Compositae) because they are advanced forms of group judging fromthe structural changes with decrease in
the number of chromosomes that occurred in the course of their phylogeny. Sez investigated the Liliacea plant
Brodiaea uniflora from the point of view of the differential morphology of chromosomes to establish its
karyotype,which consisted of 6 chromosomes. He discussed theoriginof this karyotype in view of the discrepancies
existing between various authors.
An experienced geneticist as Sez,both in animals and plants cytogenetics, knew that chromosomes exhibit
constancyof number, morphology, size, behavior, structure and other characteristics,and that these aspects of the
karyotype that must be studied carefully.
It is worth mentioning a historical comment made by Saez (168), concerning a number of alterations
observed by Garber [276] in the chromosomes of a plant, characterized by having different diploid numbers and
different types of chromosomes (long, metacentrics and telocentrics) suggestinga significant numerical and
structural variation.In this regard, Saez pointed out that Garber had not studied the morphology and somatic
chromosome number or the chromosome configurations that can be observed during meiosis. Surprisingly, this
author did not mention whether these chromosome abnormalities could have some action on the gamete viability
and fertility of the plant, and that, according to the cytogenetic knowledge of Sez, such omissions invalidate his
conclusions.
His commentary shaped a masterful example of how a cytogeneticist with vast experience in these matter
was able to evaluate the work of other colleagues and detect errors preventing them from making a correct
interpretation of chromosomal changes in genotypes and their impact on the viability or fertility of living beings.

61

Figure 35.The Sorghum almum of 40 chromosomes originated by fusing a normal gamete from Sorghum
halepense (20 chromosomes) with an abnormal one of Sorghum sudanense (20 chromosomes) according to Saez
and Nez [47]. Note the large size of the new species.

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Mammals
As early as 1928, Sez was interested in the problem of the chromosomes of mammals [7] mentioning,
first, the hypothesis of McClung (1902) on the chromosome dimorphism found in the male gametes of some
insects, in which half of the individuals had an additional chromosome compared to others,suggesting that this
chromosome, which he named accessory, was related with the sex determination in males (sex chromosome Y).
This discovery was largely confirmed and Saez mentioned the discovery of the Y chromosome in human cells
discussing the problem of heterochromosomes in mammals in a publication in 1939 [41] (Painter 1924).
With the discovery of Y chromosome it was thought, of course, thatthe chromosome formula of man was
completed (46A + XX, 46A + XY totalizing 48 chromosomes) assuming it was correct at the time(De Winiwarter
&Oguma, 1926). Only modern cell culture techniques employed decades laterclarified this classic
error,establishing the correct number of 46chromosomes of humans(Tjio & Levan, 1956).
It isinteresting to mention that Varela, Rojas and Saez published in 1934 [25, 26] a method for studying
human chromosomes from bone marrow cells using a technique that was ahead than that employed by Tjio and
Levan in 1956 so, we justly consider this contribution of Varela and Saez as a historical precursor effort to
establish the normal number of human chromosomes.
In 1930 Sez began studying the karyotypes of some mammals in South America achieving important
results in Marsupials, setting the number of chromosomes (2n = 22) of the opossum Didelphis paraguayensisOken
(Marspialia) [14, 18], indicating that the male had a chromosome pair of XY chromosomes and thatits karyotype
was comparable to the North American species Didelphys virginiana.Years later Saez found the same number and
sexpair in the red weasel (Lutreolina crassicaudata)(Desmarest) [34, 35].
Saez and Castellengo [28] made an attempt to establish the chromosome number of the rabbit and the
metaphase analyzed suggested that this animal had more than 40 chromosomes with XY sex system . In this
regard, it should be noted the extraordinary difficulties faced at that time to make a correct microscopic
observation. In their paper the authors admit that "the spermatocyte 1 is, in rabbits, a small cell which accentuates
the difficulty of studying the chromosome complement that, in most cases, is present in the preparations closely
clustered, makingimpossible the clear differentiation of the metaphase plates from the side view (equatorial
view)or from the poles (polar view)" The difficulty was originated by the biological materialthat had to be included
in paraffin and cut with a microtome to a thickness close to the average diameter of the cell nucleus of the species
under study.In addition,the chromosomes were stained with iron hematoxylin. These difficulties, added to the lack
of precisionof the photomicrograph systems availableat that time,forced the researcher to make
painstakingobservations with the microscope who had to raise and lower the micrometer to achieve an idea on how
the individual chromosomes were positioned in a metaphase plate, without the possibility of separating them or
holding them on the same plane as is possible using the present techniques.

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This therefore meant, as a test of patience, perseverance and heroism not only for they had to make direct
observations but also documenting images in the study, which were achieved using the camera lucida and drawing
the chromosomes with Indian ink, a task in which the researcher went back and forth for hours
todocumentaccurately the karyotypes. When you compare those historical and primitive technologies with the
comfortable, accurate and rapid current observations made using modern photomicroscopes no onestops admiring
those Herculean efforts of classic cytogeneticistswho obtaining accurate results in most cases.

NativeAnimals
Sez was always interested in native animals of Uruguay. In 1964 he investigated with Drets and Brum the
somatic and meiotic chromosomes of the armadillo Dasypus hibridus(2n = 64) [86]. They also described the
karyotype of the world'slargest rodent Hydrochaeris uruguayensis, the(carpincho or capibara) (2n = 66), in
1971[102] and in 1973 [111],two typical animals of our country using modern techniques of chromosomal
spreadsusing cell suspension as starting materials.

Figure 36 - a: Dasypus hibridus 2n=64 (Mulita). b: Hydrochaeris uruguayensis 2n=66(Carpincho)

Because of his permanent interest in the chromosomes of mammals Sez (1972) conducted a joint research
with Pablo Kiblisky, Nadir Brum and Glaucia Prez-Mosquera about the chromosomal variability of different
rodents populations of the genus Ctenomys of Uruguay, and discussed the origin of their multiformity[104, 117].

64

Arachnids
Among the topics that Sez investigated with Manuel O. Daz, werethe karyotype of eleven species of
Araneida [89 90] belonging to eight families:Dysderidae , (2n = 9, X-0); Segestridae , (2n = 9, X-0, and 2n = 14,
XX-0); Sicaridae , (2n = 14, XX-0, and 2n = 20, XX-0); Amaurobiidae, (2n = 40, XX- 0);Sparassidae , (2n = 42 ,
XX- 0); Lycosidae , (2n = 22, XX- 0 and 2n = 19, X -0) ; Theriididae ,(2n = 22, XX- 0) , and Argiopidae , (2n =
24, XX-0).
These karyological analyses allowed establish: a) the absence of centromeric constriction in three
species,which they interpreted as cases of diffuse centromeres; b) theyobserved a system of X sex determination in
eight speciesand of the X0 system in only three of them. In the illustrations appearing in the publications
mentioned it is observed many species possessing acrocentric chromosomes, except Dysdera magnaand
AriadnamollisX that wereprobably metacentrics,and that the family Dysderidaeshowed the largest chromosomes.
These were the first cytogenetic investigations on Arachnids carried out in Uruguay.

Experimental Studies.
Sez was always deeply concerned about how the various physical agents could affect the chromosome
structure. Thus, in 1941 Sez described, for the first time, chromosomal alterations induced by gravity obtained by
centrifugation of somatic cells of the root meristem of Lathyrus odoratus (Leguminosae) [43]. In this paper he
pointed out that gravity was able to displace nuclei, chromatin and the nucleolus. Metaphasic chromosomes were
also displaced but instead in anaphase chromosomes were resistant and that the achromatic spindle acted as a rigid
system. Sez also concluded that translocations were produced during prophase since he observed dicentric bridges
during anaphase. In addition, chromosomal abnormalities were obtained by the action of low temperatures [167].
Sez described similar alterations induced by gravity inSchistocerca paranensis since the nuclei showed
unequal number of chromosomes, breaks, dicentric bridges and other disturbances determining that the most
sensitive meiotic stagesare leptotene and gonial metaphases. These studies meant that was possible to induce,for
the first time, chromosomal aberrations by centrifugal force, which would cause gene mutations [44, 45].

65

Cytotaxonomy
The relation of cytology to taxonomy had highlighted by McClung (1908) based upon the existence of the
relationship between the chromosomes of acrididae and their somatic characteristics.
When Sez was investigating the grasshoppers heexplained in 1929 and 1930 that cytogenetic
investigations were clearly related to taxonomy since they were able to influence the direction of the taxonomic
problems exceeding the criteria of those times on the differentiation and classification of species merely based on
morphological aspects. In this respect, Saez analyzed in 1931 the reality of heterohomeotypic scheme of Gregoire
[16] drawing attention to the value of cytology and genetics for the future of systematics and noting that the latter
should be based upon the behavior of chromosomes from a phylogenetic point of view [17].
Then, he made an extensive review of the correlation between chromosome composition and the
mechanisms of numerical changes observed in the individual, the species, the genus and higher taxonomic groups,
realizing that they werecritical to establish correlations between cytological and somatic phenomena.
It is extremely enlightening to visualize, after so many years, the precision of the reasoning displayed by Sez in
this field in 1929!:
If the chromosomes are the key elements of the characters development in the individual and, therefore, in
the species and across taxonomic groups,their importance is evident in the establishment of the phylogenetic
relationships of a given biological group.
The criteria of evidence based on the composition of the external specific characters are often insufficient,
and the work of taxonomists should be made with caution, particularly when working with material showing
marked somatic variability.
The systematic interpretation of future natural forms must be based primarily on hybridization experiments
and on the investigations carried out about the behaviour of chromosomes from a phylogenetic viewpoint.
Nor should we think that the groups created by taxonomists are necessarily definitive and it seems to us
reasonable to take into account chromosomes material, since chromosomes are factors of the characters
development of an individual, forming each biological group with its species a chain of genetically related
organisms
This basis led him to say in 1935, in relation to the contribution of cytogenetics in taxonomic problems the
following:
"A zoologist or botanist, and I refer to them especially, although this concerns of course to anyone
who cultivates any biological discipline, that they at present cannot ignore or remain with
indifference to the progressesoccurred in this fundamental branch of knowledge " [9, 13,15].

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Sez thus became the first Latin American cytogeneticist that issued such important concepts between the
biological link between cytogenetics and taxonomy.

Sex chromatin
In 1959, Brum, Laguardia and Saez conducted a study on the sex chromatin [72] originated by the XX
female constitution, to establish whether itsdetection depended on the zoological group, the staining technique or
the observation method. For this analysis, they used cells of Laplatacris dispar, Bufo arenarum, cat cells and
Didelphis. Based on the statistical data analysis, the authors concluded that they confirmed the observation of the
sex chromatin in cells of the cat but, in other materials, such as grasshoppers or Bufo,they could not identify the
sex chromatin,arguing that the said corpuscle was not readily identifiable depending on the biological materials
used and the personal ability of the observers.

Experimental mutagenesis
In the 50s Clemente Estable was studying the nervous system of a rare arachnid (Acanthopachylus) when
he observed that, without apparent cause, Paramecia in culture died mysteriously. For his research, Estable
maintained a large number of Acanthopachylus in a box that was located close to a vessel containing paramecia.
His scientific deductive power suggested to him that somerelationship between arachnids and myterious death of
paramecia might exist.
Estable intelligently concluded that the aracnids kept in a nearby box that the arachnids secreted
spontaneously a high volume of a volatile substance and that this substance was capable of destroying paramecia at
a distance. Surely only the genius and insight from Estable was able to link the origin and action of this substance.
The substance was secreted by two glands skin located cephalotorxically in the arachnid,emiting a yellowish
liquid of very pungent odor that evaporated very quickly. As at that time the arachnid was called Gonyleptes as it
was considered taxonomically amember of the family Gonyleptidae,the secretion was thus namedby Estable
"Gonyleptidine " (Fig. 58 a).
At the Institute, they were able to withdraw a considerable volume of the substance observing that it had a
great bactericide power when testing in bacterial cultures,so it was considered as an antibiotic of animal
origin(Estable et al, 1955).
Saez and Drets [58, 66, 71] studied the effects of goniyleptidine into meristematic cells in Allium cepa and
in meiotic cells of the grasshopper Laplatacris dispar observing that the secretion was able to induce varying
degrees of interchromosome coalescence, C-metaphases, diplochromosomes, polyploidy and interference with cell
division .

67

An interesting aspect of these experiments was that the Orthoptera were placed in sealed containers so they
were forced to breathe thevapor of the volatile Gonyleptidine.This methodology devised by us,thusappeared
described in the literature for the first timeit turned into a technique widely used today to testnewly synthetized
volatile drugs for their capacity to induce chromosomal aberrations.

Cytospectrophotometry
With the advancements of knowledge about the chemical nature ofthe chromosomes and the nucleolus in
the 1950s [48], it was realized that there was a need to study the phenomena related to the location, identification
and behavior of certain intracellular components particularly the nucleic acids, a fact that initiated the
cytophometrical development of methods for analyzing chromosomes and nuclear components using
metachromatic stainings [55, 59].
In the Department of Cytology, University of Columbia, where Sez worked, A. W. Pollister25 and
colleagues were engaged in the study of the cytochemical localization of cell compounds, including nucleic acids.
They used the Feulgen reaction for detecting deoxyribonucleic acid (DNA), which stains in red, while ribonucleic
acid is colorless. The staining intensity of the microscope slide, asmeasured directly by microphotometry, was
interpreted as an index of its concentration, allowed investigating the variations of this important constituent of
cell nuclei, during different physiological stages in different types of tissues.
During the period of 1950-1951, Sez was in the city of New York thanks to a grant from the Rockefeller
Foundation for research at Columbia University together with Prof. AW Pollister on the microphotometrical
analysis of nuclei and chromosomes. As an extension of his training in this area, during the following year Sez
made a short stay at the Karolinska Institute in Stockholm under the direction of Torbjrn Caspersson26. The
purpose of this stay was to become familiar with microspectrophotometric analysis in order to install such
instrumentation in his laboratory of cytogenetics.
In the Pollister laboratory Saez made,for the first time, direct measurements of the absorption of Feulgenstained chromosomes in the visible spectrum. The maximum absorption measured using a Perkin Elmer precise
monochromator corresponded to 550 millimicrons wavelength approximately, and registered variations in the
heterochromatin and euchromatin, using in the microspectrophotometer a photomultiplier cell detector according
to the Pollister and Moses technique. The problem consisted of discriminating possible changes during the stages
of meiosis of sex chromosome of Orthoptera of the genus Melanoplus and other species. The limitations of the
method generated a considerable margin of error when it was used to explorate minute areas, as was the case
when, instead of measuring the absorption of whole nuclei, the technique was applied to analyzing chromosomal
segments. This issue was a real challenge to Sez who was always very interested in this analytical technology to
differentiate between the heterochromatic and euchromatic regions.

68

To develop this kind of research was necessary to employ a more refined procedure. Therefore, Sez tested
in the sex chromosome of Orthoptera, the metachromatic dye Azure B used in the techniques of Flax and Himes,
which allowed differentiating euchromatin and heterochromatin during meiosis.The nucleolus showed a maximum
absorption at 545, 590 and 650 millimicrons wavelengths of the visible spectrum, characteristic of the
metachromatic substances. Instead, the heterochromatic sex chromosome originated a curve with an absorption
maximum peak at 650 m that corresponded to a typical orthochromatic curve, but the euchromatic chromosomes
generated a different absorption curve. Thus, using the Azur B stain he found that there are differences between
euchromatin and heterochromatin so he concluded that both chromatins differ in their molecular organization.

Figure 37 - Professor Torbjrn OskarCaspersson

by the time when, with Lore Zech,he discovered
the florescence banding patterns (1969).
_______________________
25 - Arthur Wagg Pollister (Turner, 1903 - Harleysville, USA, 1994) was a zoologist and cell biologist who first
proposed that DNA is the key substance that carries genetic information. In 1949 he built a device developed over
seven years which he called "microspectrophotometer" and allowed him to measure the amount of visible light passing
through the nucleus of a cell, showing the main chemical components and the amount by which each was present. This
allowed him to define the amount of DNA present in the nucleus of a cell, which was an unprecedented breakthrough.
Through his measurements, Pollister could discern how much DNA was present in a chromosome.

Saez worked with Torbjrn Caspersson and colleagues during three additional months, aided by a research
fund at the Institute of Cell and Genetics Research of the Karolinska Institutet, Stockholm. There, Caspersson had
been developing intensively for twenty years quantitative cytochemistry, designing and building original precise
and complex instruments in the Institute workshop as its ultramicroespectrophotometer.

69

With this impressive universal instrument(see Fig 38) Casperssoncould automatically perform
spectrographic measurements using wavelengths between 200 to 2000 millimicrons. In the ultraviolet range using
wavelengths of 250 to 300 millimicrons, the area explored was reduced to a tenth of a micron square, with an error
that did not exceed one percent. The intense chemical action of the radiations used required him to take all sorts of
precautions, using a very low intensity of illumination to preserve intact the biological material, in order to obtain
accurate quantitative information. In this way,could study from the absorption curves obtained,fractions
corresponding to the various groups of nucleic acid and amino acids.

Figure 38 - Fourhistorical cytospectrophotometerss..a: First cytospectrophotometerfor ultraviolet

lightdevelopedin the 40ths decadeby Caspersson in the Karolinska Institutet laboratories ofSweden (1950); b:
Second improved version of the ultramicrospectrophotometermade by Caspersson(1953);c: The instrument of
Casperssonwas later made by Carl Zeiss (Oberkochen, Wrttemberg, Germany) and called
Microspectrophotometer UMSP1.d:Analytical spectrophotometer for visible light developed by Pollister and
Moses in 1949.
In addition from using this equipment, Sez worked with photomicrographs, exposing the slides to
Cadmium radiations of 214 to 257 millimicrons and Magnesium, ftom 280 to 309 millimicrons. Thus, by
densitometry, he obtained the extinction coefficients corresponding to various areas of mouse and cockroach sex
cells employing fresh and fixed material, prepared by the crushing method, in order to determine the absorption of
desoxyribonucleic acid, in sex chromosomes andin autosomes as well.

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During this stay in Europe Sez visited other research centers. Thus, King's College London, met a selected
group of biophysicists: Walter Davies, Yates, Chayen, Selman among others, informed Sez about their research
methods of nuclear components from tissue cultures using visible and ultraviolet light in cell culture preparations.
They localized points of interest by phase contrast microscopy, and then scanned them in the ultraviolet spectrum
immediately to avoid alterations in the reflection objectives, avoiding the use of expensive quartz optics.
At the Institute of Genetics,University of Milan, directed by Professor Camilo Barigozzi, Sez was
informed about the cytochemical researchinDrosophila tumors carried out by fluorescent chromatography, being
of particular interest the studies of giant chromosomes fromAphiochaeta whose chromatin filaments appear
together or separately depending on the diet regime used.
In addition, Sez contacted and discussedhisresearch problems with the famous cytogeneticists Cyril
Darlington and La Cour of England.In this trip he also met other distinguished researchers at the Institutes of
Pavia, Rome, Paris, Toulouse, Switzerland, Belgium, Spain and Sweden. In the United States he also discussed
with colleagues his acetic hematoxylin staining method giving conferences in the Universities of Pennsylvania,
Fordham, Chicago, Cornell, Princeton, Harvard and the Atomic Energy Laboratory of Brookhaven where he
showed slides stained with acetic hematoxylin.
_______________________
26 - Torbjrn Oskar Caspersson (Motala, Sweden, 1910-1997) was cytologist and geneticist.Together with the
biochemist Einar Hammersten conducted research on the molecular structure of DNA, which led him to discover that it
was a polymer-or-macromolecule composed of small repeating units. After completing his doctorate at the University
of Stockholm in 1936, he developed a first photomicroscope for ultraviolet light to determine the nucleic acid content of
cellular structures. In the late 1930s, with the geneticist and biochemist Jack Schultz, performed studies in which
reunited the principles of cell biology and biochemistry, bringing together techniques such as spectroscopy and
ultraviolet microscopy. After many years of research, he concluded that RNA play a role in protein synthesis. The
scientific community was not very interested in nucleic acids in 1944. Only a few scientists were active in this field.
One of them was Einar Hammarsten, Professor of Chemistry at the Karolinska Institute in Stockholm. In 1924 he had
published a thesis about the preparation and properties of DNA or thymonucleic acid as it was called at that time. In the
ensuing 20 years he worked with both DNA and RNA. Only a few students joined his endeavor. Most notable among
them was Torbjrn Caspersson who by 1944 already was Professor and head of a Nobel Institute at the Karolinska
Institute.. Caspersson headed one of the most active research centers at the Karolinska Institute. He had developed a
new ultraviolet microscope to study nucleic acid and protein metabolism (see Fig. 37a and b), and he and his
collaborators applied the new methodology to many different biological systems. In 1969 when working with Lore
Zech at the Karolinska Institute, they found a stain (quinacrine mustard) that caused chromosomes to show light and
dark lateral bands along their length. This banding method permitted the accurate identification of all 22 autosomes and
the X and Y chromosomes. This technique highlighted slight structural abnormalities and helped the identification of
extra chromosomes involved in conditions such as Down's syndrome.

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On his return to his country, the Rockefeller Foundation financed the acquisition of the components for the
first citoespectrofotometer to be introduced in Uruguay. All the instrumental parts were assembled with the
technical assistance of the Head of the Department of Biophysics of the Institute, Professor Carlos Maria Franchi,
a researcher of extreme skill and creativity both in the areas of electronics and in optics and mechanics. Franchi
not only installed the instrument of Sez in 1953 but also designed additional optical and mechanical components.
Sez had the first cytospectrophotometer existing in both countries of the Rio de la Plata.
Although Sez was familiar with the methodology and had already made their first contributions in
microspectrophotometry he tried to create not only a new instrumental environment but also to give an opportunity
to train new researchers in this field of investigations. In fact, in 1954 Sez decided to send one of his brightest
research collaborators, Claudio Leopoldo Solari to work with Prof. Lucien Lison (Fig. 39) in Curitiba, Brazil.
Lison, with J. Pasteels, had developed in 1950 a new histophotometer (different from the one installed in
the laboratory of Sez) that Lison used to quantify DNA in different cell types. This approach became a popular
laboratory tool. In 1951, using this technique with the Feulgen reaction, both authors studied the amount of DNA
in the nuclei of urchin sea eggs demonstrating that during their morphogenetic changes intense mitoses occur
increasing the amount ofDNA. Lison also conducted studies on metachromasia, and histochemistry of
phosphatases and lipids. In 1953, Lison published an important text on animal histochemistry (Histochimie et
cytochimie animales: principes and methodes, Gauthier- Villars, Paris Ed.), which became a classic book. Because
of this, Lison was called in Brazil, "the father of histochemistry". Undoubtedly, this research activity of Lison
provided Solari with an excellent opportunity to expand his training on the microspectrophotometrical
possibilities.

Figure 39.a: Sez with the new microspectrophotometer installed in his Laboratory of Cytogenetics in
1953, b: Two cytophotometrists from Brazil, Prof. Lucien Lison and Prof. Giorgio Schreiber discussing research

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on cytophotometry during the First Scientific Meeting of the Latin American Association of Physiological
Sciences, held in Punta del Este, Uruguay, in 1957.
Besides Lison, Schreiber of the Biology Institute of Minas Gerais, Belo Horizonte, Brazil, was also
interested in this technology making, in 1949, statistical and physiological studies on the growth of interphase
nuclei using the cytometricalmethod to analyze the problem during the increase of the nuclear volume of 1.5 times
from their initial size and its intermediate stages. (Schreiber, 1949).
Furthermore, the geneticist Willy Beak (born in Mulhouse, France, 1932) also introduced cytometry
methodology at the Butantan Institute in So Paulo, Brazil.

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HONORS AND AWARDS

Due to hiscontinued and valuable contributions to cytogenetics Sez earned significant scientific and
academic recognition throughout his life as researcher, recevingdistinctions detailed below:

In 1934 Sez received the National Prize for Natural and Biological Sciences of Argentina, which marked the
beginning of a successful career and recognized for their contribution to science not only in the River Plate area
but also worldwide.
Elected President of the 7th. International Congress ofGenetics, held in Edinburgh, Scotland, August 23 to
30, 1939.
The National Commission of Culture of Argentina invited Sez as a Fellow to give a lecture in the frame of the
1942 cycle. (165)
In 1965 was appointed Vice-Chairman of the Organizing Committee of the International Symposium of
the Nucleolus, Montevideo, Uruguay (212, 213)
The Society of Biology of Montevideo pays tribute to Dr. Francisco A. Saez on the occasion of its 70th
anniversary (1968)
In 1968 the Faculty of Humanities and Sciences of the University of the Republic, Montevideo, Uruguay, gives
him the title of Professor Honoris Causa.
Sez is awarded in 1968 with the title of Professor Ad-Honorem by the Faculty of Medicine Montevideo,
Uruguay,
On December 11, 1971 Sez is appointed Honorary Member of the Genetics Society of Argentina by the
Assembly of the Institution held in the city of Santa Catalina, Argentina.
Sez receives (1972) the Lucio Cherny Award (given on behalf of the Lucio Cherny Foundation by Dr. Luis
Federico Leloir Nobel Prize and Honorary Member of the Society of Genetics,Argentina (The Award was
presented by Mr. Ewald A. Favret of the Genetics Society Argentina) at the Medical Association of Argentina on
June 28, 1973 at 18 am.
In 1974 Sez is appointed Honorary President of the American Genetic Association (ALAG) during the 2nd
Latin American Congress of Genetics held in Mexico City

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On behalf of his disciples, M.E. Drets delivered the following speech in 1968 on the occasion of the
ceremony to award Sez the title of Professor Ad Honorem held in the Auditorium of the Faculty of Medicine of
Montevideo, Uruguay.
Mr. Dean ,Members of the Council, Ladies and Gentlemen :
The great researcher Clemente Estable once said that:"In general, human progress is mainly due to early
maturing youth and men who do not grow old Let me start with these lucid words because I think they are
extremely defining the spiritual state that this tribute brings us here jubilant. And I think they are a miraculous
synthesis of the significance of the personalityand work of Prof Sez, particularly applicable to our environment
and its scientific development, we celebrate here hisanniversary,that is also a facet of the culmination of the
revolution started in the last century of which he has been a distinguished protagonist
This remarkable change that occurred in just over half a century has taught us to understand not only the
gene properties and functions but also the cytogenetic mechanisms involved. This understanding was based, of
course, upon the detailed knowledge of chromosome structure and physiology at different stages of cell division.
But this knowledge, that every day is deeper and more varied,which continues reveal mew perspectives in
the hope of obtaining a full view on the hereditary systems could not be integrated without the solid foundation
built by Jansens on chiasmata, by Carothers on the independent segregation of structurally heterozygous bivalent,
by Balbiani on polytene chromosomes, by the brilliant Bridges on the mechanism of non-disjunction, gaps and
duplications in Drosophila and his gene balance theory of sex determination, extraordinary concepts prevailing
today; by Belling father of now common chromosome crushing in his old acetic carmine; the inspired Muller,
inducing structural changes and gene mutations with X rays; Heitz, creator of the concept of heterochromatin;
McClung who discovered the sex chromosome Y; Goldschmidt whose observations the Lymantria butterfly led him
to the idea of the intersex and the theory of the quantitative relationship of genes; in short, from Darlington solid
scaffold builder of the cytological chiasma hypothesis and gene exchange that still survives intact, among many
others that one should also mention at this time.
These were three successful decades of progressive illuminations. And there, by the thirties approximately
appears in these latitudes and by lucky gene recombination, as in historical times, a new and curious conqueror.
His vocational mision was not to conquer lands and unknown beings, but to start studying our wild flora
and fauna were waiting to reveal their secrets. That unexplored biological world was andalso still is our America

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and the master Sez of white uniform, microscope in hand and chromosomein his heart, began to studyour living
beings without pause 40 years ago!
Perhaps this is not the most adequate time for a detailed analysis of the vast work of Prof Sez, it is in the
domain of this select audience. Besides, a thorough job of this nature has the flavor of the old and the past when it
is intended to bring all us here propitiatory auguries and an expression of our faith in the continuation, in the
multiplication of his future work.
However, it is now appropriate to try to make a retrospective overview and synthesis reviewing very
briefly the contributions that, in our opinion, are relevant examples of his scientific interests.But, It is absolutely
impossible to separate the personality from hisproduction:.they are children of the intellect that always go along
with his father.
His research has been, within a common thematic denominator, extremely varied. He conducted research
on cytological techniques, chromosomal cytotaxonomy and evolution, studies of natural populations, studies in
Orthoptera, amphibians, mammals, sex chromosomes and sex determining mechanisms; hybrids and polyploids;
action of chemical substances and physical agents, cytotophotometrical and cytochemical studies. Fortunately, all
this was produced showing a wide experience and brightly didactic sense, as it appears in his widespread text
"General Cytology" in colaboration with De Robertis and Nowinski, now called Cell Biology that was edited not
only into Spanish but also in English, French, Japanese, Russian and Italian. No doubt this work contributed to
the development of important cytological investigations and to the increase of his deserved international prestige.
I think it is highly instructive to recall a few typical examples of his research indicating the directions that
followed his work and which has remained firm over the course of the years.
First, I wish to mention his old paper published in the thirties entitled "lnvestigations on the chromosomes
of some Orthoptera of South America". He examined the chromosomes of the sex cells in four genera of South
American grasshoppers. Object of particular study was the called at the time accessory chromosome" or the sex
chromosome now . In this investigation Sez clearly related purely morphological features of chromosomes with
the systematics of the genus and its phylogenetic implications. This contribution, in the style McClung school, of
which he was a fervent disciple, became a classic not only for the early date of its publication but because it
attracted great international interest. Sez, loyal to this type of cytotaxonomic studies in Orthoptera, reported in
more recent paper (1956) the extraordinary case of a grasshopper carrying eight diploid chromosomes and XY sex
mechanism.That was followedby a series of studies on this insect. In 1936 he showed, investigating the sex cells of
anura, that in amphibians there is no sex chromosome, definitively clarifying doubts at that time.
Another prototypical example of research line that has been pioneered by Sez, was carried on "The
cytologyof Sorghum findinga new species with 40 chromosomes. He demostrated that this new plant species was
actually an interspecific hybrid caused by species with a lower number of chromosomes. Finally,and according to

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my personal preferences, perhaps the work that raises most suggestions from the theoretical and conceptual point
of viewis the gradient of heterochromatinisation in the evolution of neo-X neo-Y sexual system. This is a splendid
work of problem synthesis which, based on morphological features of chromosomes, made excellent physiological,
biochemical and genetic correlations.
Nevertheless, in my personal opinion, the most significant aspect of his work for our countries has been the
task of training young people who have continued developing our discipline. The broad and spontaneous Sez
concept of Disciple made his laboratory a permanent crossroads with aconsiderable number of young people
learning cytogeneticsand receiving his example of a great scientist and thethoroughness of expert cytologist.
That constant flux of young people as I witnessed firsthand for almost 27 years now, entitles me to say,
with the safety that comes from such a long period, that Professor Sez always received all of them with the same
permanent warmth and genuine affection in his Laboratory of Cytogenetics. I do not know too much about his
relationship with the people who wanted to know more on chromosomes before 1948, but by simple extrapolation,
genotype and phenotype to the grave.
Considering the legion of disciples trained by Sez he can take legitimate pride that many of them reached
sometimes the distant sky of Science. So,Iremember now Eduardo D.P. De Robertis , Ovidio Nez, Isaac Tarasiuc
, Alfonso Castronovo , our unforgettable Claudio Solari , Alejo Mesa, Alvaro Diaz , Nadir Brum , Amalia
Laguardia, Glaucia Prez Mosquera, Manuel Daz, Horacio Cardoso , among others, who have been his followers
, some of them continuing their personal research lines, performing other new approaches. But it is justice to say,
he encouraged them to develop within and outside your lab, becoming receptive to all new problems, all ideas,
good or inadequate , continually joining usby the abstract but fortunate matter called Cytogenetics and that has
joined us intellectually.
The words of the wise men are like goads," says Ecclesiastes. The immense responsibility to continue the
work in progress with personal style, the ineffable concerns that bring us new facts, the endless renewal of
cytological problems are constantly darts piercing the spirit of his disciples. The growing incessant complexity
and of the Discipline, withits new lines and sub-lines seem likea cosmic explosion and the separations beween
themmake more and more difficult the task delegated to his School.
However, the hard challenge of Gorgias is still alive. And like him in this hour of joy and optimism
celebrating the Master who does not abandon us but, on the contrary, he will continue with us as the direct
successors of his work. I express, in the dimension of Rod words, my most intimate and profound hope that
someone among them "put the foot forward of his last foot stepand his forehead even more in the clear and
spacious" This act will also be a vivid and permanent of their accomplishments. And there, in the ardent hope this
happens is our sincere emotion and our warm tribute and recognition. It is the most authentic feeling of his
Disciples.

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Mximo E. Drets, March 15, 1968.

Figure 40 Sez receiving the Lucio Cherny Prize.

At right, Luis F. Leloir Nobel Prize of Chemistry (1970).
Speech given by the Academic Prof. Dr. Eduardo D.P. de Robertis during the ceremony of Prof. Francisco A. Sez
receiving the Corresponding Membership of the National Academy of Exact, Physical and Natural Sciences
of Buenos Aires, Argentina, Saturday, May 4, 1968 (222).

Members of the Academy, Ladies and Gentlemen:

As a corresponding member in Montevideo we incorporate today to our Academy a researcher of
international stature, who founded Cytogenetic studies in Latin America, and who made important original
contributions and formed disciples who follow his teachings with devotion and affection.
Descended from a distinguished family of the Republic of Uruguay, Francisco Alberto Sez represents an
admirable example of the commonwealth of the people of Rio de la Plata. From a young age he joined our milieu,
studied at the Faculty of Humanities at the National University of La Plata, graduating with the title of Professor,
with high scores in l927, being formed by the wisdom of Argentinian zoologist Miguel Fernndez and received the
decisive influence of Professor Clarence McClung of the United States, who decidedly pushed him to study of
chromosomes and their role in inheritance, sex determination, variation and evolution of living beings. Sezs
work "Research on theChromosomes of some Orthoptera" published in 1930, was considered a classic in its genre
by the international interest generated immediately. It was a few years later, while I was still a novice student I
met him and received the impact of his boundless enthusiasm and passion for awakening vocations. From our
association resulted that was rewarded with work on the chromosomes of amphibians, we published in the

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Zeitschrift fr Zellforschung in 1936 that was rewardedwith the National Science Award and to me a German
nickname by my hospital colleagues.
Although our research steps diverged, the friendship between us contributed to the publication in 1946 and
with the help of Nowinski we published in 1946 the first edition of the book "General Cytology", which has since
been our common spiritual son. After serving as Professor of Natural Sciences at the University of La Plata, where
he worked for about 27 years, at the Faculty of Agronomy, Museum and in the Normal School in the year of 1946,
so terrible for our country and that did not succeed in separating him from returning to his country of origin, Sez
became Head of Department of Cytogenetics at the Research Institute of Biological Sciences, Montevideo, and
formed an important and numerous group of researchers who have had great influence in the Latin American
cytogenetics.His research interests ranged from general cytogenetics related to taxonomy and evolution, to the
cytospectrophotometry and mutagenic action of physical and chemical agents. Classes were his contribution to
classical cytogenetics of the genus Sorghum, so studied from the botanical point of view, by our always
remembered colleague Lorenzo Parodi,as well ashis studies on the neo-X, neo-Y sexual system. In his lecture
today he will give us an outstanding example of the influence of chromosomal variations in the evolution of living
organisms.
His curriculum and achievements are so numerous, that I could not even mention them all without
exceeding the limit of my time. More than 200 published works; fellow at the University of Pennsylvania, the
American Phylosophical Society, the National Commission of Culture of Argentina, and the Rockefeller
Foundation. He perfected his studies at Columbia University and the Karolinska Institute in Stockholm. He has
given lectures and seminars at numerous universities in the United States, Canada, Belgium, Switzerland,
England, Italy, Portugal and Sweden. He is a member of numerous institutions of his specialty and was nominated
Vice-President of the International Congress of Genetics, Edinburgh(1939) and Vice-Chairman of the Organizing
Committee of the International Symposium on the Nucleolus (1965) [213].Recently, reaching his seventh decade,
the forever young Sez receives the distinctionHonoris Causa from the University of Montevideo. The
diplomaaccredits him as as corresponding member of our corporation is an additional laurel on that crown that
he has garnered through so many years of efforts, the service yo the ideal of science, training of disciples and the
brotherhood of the peoples of America
Eduardo D.P. de Robertis (May 4, 1968)
Transcription of some of the issues raised by Prof. Saez on occasion of receiving the Prize "Animal and human
genetics " (1972 ), awarded by the Foundation L. Cherny , (fragment) .
" ... The participation of Latin America in genetic research began with us, as we said before, half a century
ago, the corresponding to Argentina the roleof precursor of the discipline in the continent. Its advancement

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through these years has been remarkable; then Brazil, Chile and Uruguay provided valuable progress on various
issues. The Argentina counts with researchers of international level working in different fields of research, that
have given new and brilliant impetus to basic and applied genetics. UNESCO some time ago, made a division of
the scientific world into two zones, one clear and bright, the other dark. In the latter was Latin America. While it is
true that, it has been often said that in our America, there is no comprehensive scientific development, but instead
few points of light,few institutes and laboratories where first-class scientific research is carried out.Some men
insist to do their best of themselves to carry out inquisitive research despite all the difficulties Therefore, we can
add that we have already transposed the wall of shadow, and penetrated the mid-light in an upward march
towards clearness.Science is advancing rapidly in these latitudes and becomes increasingly stronger by the
contribution of its researchers of high intellectual level"

Figure 41- a: Saez and Drets visiting the Anthropological Museum of Mexico (DF) in 1974
few days after they were elected Honorary President of the American Genetic Association (ALAG)
and President of ALAG (period 1974-1977) respectively, b: Sez talking with the eminent
human geneticist Curt Stern27during their visit to Ouro Preto, Brazil, in 1968 accompanied
by theIIBCE researcher Jos Roberto Sotelo.
_______________________
27 - . Curt Jacob Stern (Hamburg , Germany, 1902 Sacramento, California 1981 ) was an eminent
human geneticist . In 1949 he published the first book on human genetics, Principles of Human
Genetics, edited by WH Freeman and Co., which was translated into Spanish by ME Drets and edited
by El Ateneo in Argentina in 1963.

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Disciplesand Coworkers
(Uruguay) -Claudio L. Solari ; Hugo E. Silvera ; Mximo E. Drets; Horacio Cardoso; Nadir Brum; Mario Stoll;

Hctor N. Seunez; lvaro Daz; Amalia Laguardia ; Glaucia Prez-Mosquera; Manuel O. Daz, Hugo Montero,
Gustavo A. Folle, Alejo Mesa ; and Orfeo Crosa.
(Argentina)Ovidio Nez; Eduardo P. De Robertis ; Fernando Dulout ; Alfonso Castronovo; Isaac Tarasiuk ;
Avelino Barrio; Primarosa R. Rinaldi; Pablo Kiblisky and Alberto J.Solari.
(Brasil)Lucy I. S. Peixoto; and C.F.U. Knapper
(Chile)Waldo Venegas
(United States of America)Edward M. De Robertis

Figure 42 Professor Sez with his first Uruguayan disciples. The picture was taken after participating in the First
South American Congress of Agronomical Research developed in the Phytotecnical Institute and National Seed
Repository La Estanzuela Colonia, Uruguay, in November,13-19, 1949. Left to right: ME. Drets, Hayde
Castelo, F.A. Sez, Claudio Solari and Hugo E. Silvera

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Scientific Publications
1.
Sez FA (1925): Sobre algunos mtodos generales empleados en tcnica microscpica. Rev Centro Est
Med Veter Univ Nal La Plata, 2: 1-42.
2.
Sez FA (1927): Un caso de pseudoparasitismo en un carpincho por larvas de dptero. Resumen de la
comunicacin a la Sociedad Uruguaya de Ciencias Naturales. Physis, 8: 659.
3.

Sez FA (1927): Notas de bibliografa citolgica. Physis, 8: 666-684.

4.

Sez FA (1927): Algunas consideraciones tcnicas sobre el estudio de los cromosomas. Physis, 8: 481-502.

5.
Sez FA (1927): Los cromosomas en la espermatognesis de algunos ortpteros sudamericanos.
Comunicado a la Soc Uruguaya Cien Nat (sesin del 29 de enero).
6.
Sez FA (1928): Exhibicin de microfotografas de cromosomas de ortpteros sudamericanos y
consideraciones relativas al tema. Resumen. Physis, 9: 101-106.
7.
Sez FA (1928): Estado actual del problema sobre los cromosomas de los mamferos. Real Soc Espaola
Hist Nat, 2: 109-119.
8.
Sez FA (1929): Le carmin actique ferrique dans ltude des chromosomes des animaux. Arch Soc Biol
Montevideo, 1: 258-261.
9.
Sez FA (1929): Puede la citologa influir en la orientacin de los problemas taxonmicos? Rev Soc
Entomol Arg, 10: 251-262.
10.
Sez FA (1930): Investigaciones sobre los cromosomas de algunos ortpteros de la Amrica del Sur. I.
Nmero y organizacin de los complejos en cuatro gneros de acridios. Rev Mus La Plata, 32: 317-361.
11.
Sez FA (1930): Organizacin y nmero de cromosomas en los ortpteros de Amrica del Sur. Actas Cong
Intl Biol, Montevideo (oct 7-12), pp. 1-8.
12.
Sez FA (1930): Nota sobre los complejos de cromosomas en cuatro especies de ortpteros (Acrididae) del
Ro de la Plata. Arch Soc Biol Montevideo, 2: 35-43.
13.
51.

Sez FA (1930): Sobre las relaciones de la citologa con la taxonoma. Arch Soc Biol Montevideo, 1: 25-

14.
Sez FA (1930): Los cromosomas de algunos mamferos de Amrica del Sur. Nota sobre el complejo
cromosmico de Didelphis paraguayensis Oken (Marsupialia). Actas Cong Intl Biol, Montevideo (oct 7-12), pp.
1-7.
15.
Sez FA (1931): Las conquistas de la biologa celular y su influencia en los grandes problemas de la
biologa contempornea. Bol Cien Nat Inst Estud Sup, 1: 3-12.

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16.
Sez FA (1931): Las mitosis de maduracin y la realidad del esquema heterohomeotpico de Grgoire.
Physis, 10: 378-391.
17.
Sez FA (1931): Sobre las relaciones de la citologa con la taxonoma. Cong Intl Biol, Montevideo, 1930,
Arch Soc Biol Montevideo, S1: 1-27.
18.
Sez FA (1931): The chromosomes of the South American opossum, Didelphis paraguayensis. Am Nat,
65(698): 287-288.
19.
Sez FA (1931): Un sencillo dispositivo para la inclusin en parafina que elimina ventajosamente el
empleo de la estufa termorreguladora. Physis, 11: 82-85.
20.
Sez FA (1931): Variacin numrica en correlacin con la existencia de cromosomas mltiples en Aleuas
vitticolis sp. (Orthoptera: Acrididae). Rev Mus La Plata, 33: 189-193.
21.
Sez FA (1932): Individualidad de los cromosomas. La constancia de la forma y el criterio de
identificacin. Arch Nac Biol Med, 3: 46-51; Soc Nac Biol Med, Buenos Aires, 10(2): 46-51.
22.
Sez FA (1932): Organizacin y nmero de cromosomas en los ortpteros de Amrica del Sur. Arch Soc
Biol Montevideo, 7: 1912-1919.
23.
Sez FA (1934): Formule chromosomique du crapaud Bufo arenarum (Hensel). Comp R Soc Biol Paris,
117: 1242-1243.
24.
Sez FA, Rojas P & De Robertis E (1934): La frmula cromosmica del sapo Bufo arenarum (Hensel).
Rev Soc Arg Biol, 10: 340-344.
25.
Varela E & Sez FA (1934): Un nuevo mtodo para el estudio de los cromosomas somticos del hombre.
Rev Soc Arg Biol, 10: 421-422.
26.
Rojas P & Sez FA (1935): Estado actual del problema relativo a los cromosomas de la especie humana.
Cong Nal Med, 3: 69-74.
27.
Sez FA (1935): Estructura de los cromosomas en dos especies del gnero Aleuas. Actas Cong Nal Med,
Rosario, pp. 93-96.
28.
Sez FA & Castellengo L (1935): Nota sobre los cromosomas del conejo. Actas Cong Nal Med, Rosario, 3:
81-83.
29.
Sez FA, Rojas P & De Robertis E (1935): El problema de los cromosomas sexuales en los anfibios. Rev
Soc Arg Biol, 11: 173-176.
30.
Sez FA, Rojas P & De Robertis E (1935): Le problme des chromosomes sexuels des amphibiens. Comp
R Soc Biol Paris, 120: 368-369.
31.
Sez FA, Rojas P & De Robertis E (1936): Investigaciones sobre las clulas sexuales de los anfibios
anuros. El proceso meitico en Bufo arenarum (Hensel). Inst Mus Univ Nal La Plata, 2: 95-143.

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32.
Sez FA, Rojas P & De Robertis E (1936): Untersuchungen ber die Geschlechtszellen der Amphibien
(Anuren). Der Meiotische Proze bei Bufo arenarum. Zeit Zellforch Mikro Anat, 24: 727-777.
33.
Sez FA (1937): Sobre la citologa de Atelopus stelzneri (Weyenbergh) (Anfibios: Anuros). Arch Fitot
Uruguay, 2: 452-454.
34.
Sez FA (1938): Formule chromosomique et complexe sexuel chez Lutreolina crassicaudata (Desmarest).
Comp R Soc Biol Paris, 129: 865-866.
35.
Sez FA (1938): Investigaciones citolgicas sobre los marsupiales sudamericanos: frmula cromosmica y
complejo sexual en la comadreja colorada Lutreolina crassicaudata (Desmarest). Rev Soc Arg Biol, 14: 156-161;
Physis, 18: 153-159 (1939).
36.
Sez FA (1938): Estado de las investigaciones sobre citologa de las variedades de Schistocerca
paranensis. Rev Fac Agron La Plata, 22: 129-137; Physis, 17: 253-259 (1939).
37.
122.

Sez FA (1939): El problema de la determinacin del sexo en los vertebrados inferiores. Physis, 18: 111-

38.

Sez FA (1939): Sobre el origen y evolucin de las tetradas anulares en los anfibios. Physis, 18: 165-174.

39.

Sez FA (1939): Nota sobre la citologa de Atelopus stelzneri (Weyenbergh). Physis, 18: 161-164.

40.

Sez FA (1939): Observaciones sobre la reaccin nuclear de Feulgen y Rosenbeck. Physis, 18: 123-126.

41.

Sez FA (1939): El problema de los heterocromosomas en los mamferos. Physis, 18: 161.

42.
Sez FA & Gershanik R (1939): Nota sobre la citologa de Trygonophymus arrogans (Orthoptera:
Acrididae). Cong Nal Med, Crdoba, 1: 890-893.
43.
Sez FA (1941): Alteraciones experimentales inducidas por la accin de la gravedad en las clulas
somticas de Lathyrus odoratus (Leguminosae). An Soc Cient Arg, 132: 139-150.
44.
Sez FA (1941): Produccin experimental de alteraciones en los cromosomas de Schistocerca paranensis.
Rev Arg Agron, 8: 61-62.
45.
Sez FA (1941): Efectos de la centrifugacin sobre las clulas sexuales de Schistocerca paranensis. 7th Intl
Cong Genet Edimburgo, J Genet, Suppl, Cambridge, p. 253; Centro Est Agron, pp. 9-10.
46.
Sez FA (1941): Nota sobre la citologa del gnero Hypochoeris (Compositae). 1 Reunin Arg Agron, Rev
Arg Agron, 8: 62-63.
47.
Sez FA & Nez O (1943): La citologa de Sorghum almum (Parodi): nueva especie con 40 cromosomas.
I. Los cromosomas somticos. Notas Mus La Plata, 8: 333-348; Rev Arg Agron, 11: 152-157.
48.

Sez FA (1945): La naturaleza qumica de los cromosomas y del nuclolo. Bol Soc Arg Bot, 1: 4-35.

49.
Sez FA (1949): Estudio citolgico comparativo de algunas especies del gnero Hypochoeris (Compositae)
de la Amrica del Sur. Actas 2 Cong SA Bot, Tucumn, Lilloa, 19: 97-105.

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50.
Sez FA (1949): Los cromosomas de Brodiaea uniflora. Actas 2 Cong SA Bot, Tucumn, Lilloa, 19: 105110; Inst Inv Biol Pub, 1: 207-212 (1951).
51.
Sez FA (1949): En torno a la meiosis de Sorghum almum (Parodi). Actas 2 Cong SA Bot, Tucumn,
Lilloa, 19: 111-118; Inst Inv Biol Pub, 1: 185-197 (1951).
52.
Sez FA, Silvera H & Solari CL (1950): La estructura gentica de Schistocerca cancellata. Rev Fac Hum
Cien, Montevideo, 5: 23-26.
53.
Sez FA (1951): Estudio citolgico de algunas especies del gnero Hypochoeris (Compositae). Inst Inv
Biol Pub, 1: 199-205.
54.
Sez FA (1952): Differentiation of meiotic heterochromatin and euchromatin by microspectrophotometric
techniques. Anat Rec, 113: 65.
55.

Sez FA (1953): Espectrofotometra del cromosoma. Hoja Gentica, Soc Riopl Gent, 1: 10-12.

56.
Sez FA (1953): Anlisis citogentico de los cromosomas de Schistocerca cancellata. Arch Fitotec
Uruguay, 5: 395-396.
57.
Sez FA (1953): La citogentica del sorgo negro (Sorghum almum) y el problema de su origen. Arch
Fitotec Uruguay, 5: 404-405.
58.
Sez FA & Drets ME (1954): Cytological alterations induced by gonyleptidine during cell division. Rec
Genet Soc Am, 23: 76.
59.
Sez FA (1955): Estudio microespectrofotomtrico de la heterocromatina y eucromatina. Riv Istoch Norm
e Patol, Miln, 1: 485-502.
60.
Sez FA (1955): Interspecific cytological
Cyrtacanthacridinae). Rec Genet Soc Am, 24: 593-594.
61.

variation

in

the

genus

Scotussa

(Orthoptera:

Sez FA (1956): Cytogenetics of South American Orthoptera. Nature, 177: 490-491.

62.
Sez FA (1956): Estudios citogenticos en ortpteros sudamericanos: el cariotipo de treinta y dos especies.
Biolgica, Chile, 22: 21-26.
63.
Sez FA (1956): Caso extraordinario de un ortptero acrdido con ocho cromosomas diploides y
mecanismo sexual XY. Biolgica, Chile, 22: 27-30.
64.

Sez FA (1956): Citofotometra de la heterocromatina y eucromatina. Biolgica, Chile, 22: 31-36.

65.
Sez FA & Daz (1956): Sistema sexual neo-X neo-Y en Xyleus laevipes (Orthoptera: Romaleinae).
Arch Soc Biol Montevideo, 23: 13-27.
66.

Sez FA & Drets ME (1956): Chromosome alterations induced by gonyleptidine. Biolgica, Chile, 22: 37.

67.
Sez FA (1957): Una mutacin cromosmica excepcional que no afecta la viabilidad del individuo. 1
Reunin Cient, Asoc LA Cien Fisiol, Montevideo, pp. 154-155.

85

68.
Sez FA (1957): Caracterizacin morfofisiolgica de la heterocromatina. 1 Reunin Cient, Asoc LA Cien
Fisiol, Montevideo, pp. 200-201.
69.

Sez FA (1957): An extreme karyotype in an orthopteran insect. Am Nat, 91: 259-264.

70.

Sez FA (1958): Chromosome evolution in Dichroplus. Intl Cong Genet, Montreal, 2: 247-248.

71.
Sez FA & Drets ME (1958): The action of gonyleptidine on the mitotic and meiotic chromosomes and on
the interphase nucleus. Portugaliae Acta Biol, 5: 287-302.
72.

Brum N, Laguardia A& Sez FA (1959): A study of sex chromatin. Tex Rep Biol Med, 17: 73-81.

73.
Sez FA & Brum N (1959): Citogentica de anfibios anuros de Amrica del Sur. Los cromosomas de
Odontophrynus americanus, Odontophrynus cultripes y Ceratophrys ornata. An Fac Med, Montevideo, 44: 414423.
74.
Sez FA & Solari CL (1959): Chromosome studies in three species of Scotussa (Orthoptera:
Cyrtacanthracridinae). Caryologia, 11: 358-368.
75.
Sez FA (1960): El empleo de la hematoxilina actica o propinica para el estudio de los cromosomas con
la tcnica de aplastamiento. Com Soc Biol Montevideo (mimeografiado) (11 de mayo).
76.
Sez FA & Brum N (1960): Consideraciones citogenticas de nuevas especies de anfibios anuros
sudamericanos. Resumen. Sesiones Cien Biol, Mendoza, pp. 88-89.
77.

Sez FA & Brum N (1960): Chromosomes of South American amphibians. Nature, 185: 945.

78.
Sez FA & Daz (1960): Neo-X neo-Y system of sex determination in Xyleus laevipes (Orthoptera:
Romaleinae). Tex Rep Biol Med, 18: 116-128.
79.
Sez FA & Solari CL (1960): Estudio citogentico del gnero Scyllinops (Orthoptera: Acrididae). Arch
Soc Biol Montevideo, 24: 36-42.
80.
Sez FA (1961): Investigaciones citogenticas de anfibios sudamericanos. Simp Sul-Am Genet Fac Filos
Cien Letras, So Paulo, pp. 286-288.
81.
Sez FA & Brum N (1961): Investigaciones citogenticas de anfibios anuros de Sudamrica. 1er Cong
Sudam Zool, La Plata, 5: 303-305.
82.
Sez FA & Laguardia AM (1961): La estructura citogentica de Laplatacris dispar (Orthoptera:
Acrididae). 1er Cong Sudam Zool, La Plata, 5: 307-315.
83.
Sez FA (1963): Gradient of the heterochromatinization in the evolution of the sexual system Neo-X NeoY. Port Acta Biol, Lisboa, 7: 111-138.
84.

Sez FA & Brum N (1963): Cytogenetics of South American amphibia. Genet Today, The Hague, 1: 141.

85.
Sez FA & Prez-Mosquera G (1963): Anlisis, citogentica y evolucin del cromosoma sexual en
Schistocerca infumata. Ann Soc Biol Rio Grande do Sul.

86

86.
Sez FA, Drets ME & Brum N (1964): The chromosomes of the mulita (Dasypus hybridus Desmarest): a
mammalian edentate of South America. Mammalian Cytogenetics and Related Problems in Radiobiology
Symposium, So Paulo, 1962.Pergamon Press, Oxford, pp. 163-170.
87.
Sez FA & Prez-Mosquera G (1964): Evolucin del cromosoma sexual en Schistocerca infumata. Arch
Soc Biol Montevideo, 26: 50-62.
88.
Sez FA & Brum N (1965): Los cariotipos de las especies del gnero Odontophrynus. Actas 2 Cong LA
Zool, So Paulo, 2: 287-290.
89.
Sez FA & Daz MO (1965): Investigaciones citogenticas sobre algunas especies de araneidos uruguayos.
Actas 2 Cong LA Zool, So Paulo, 2: 3-9.
90.
Daz MO & Sez FA (1966): Karyotypes of South-American Araneida. Symp Intl So Paulo, Mem Inst
Butantan, 33: 153-154.
91.
Sez FA & Brum N (1966): Karyotype variation in some species of the genus Odontophrynus (Amphibia,
Anura). Caryologia, 19: 55-63.
92.

Brum N & Sez FA (1968): Chromosomes of Leptodactylidae (Amphibia: Anura). Experientia, 24: 969.

93.
Daz MO & Sez FA (1968): DNA synthesis in the neo-X neo-Y sex determination system in Dichroplus
bergi (Orthoptera; Acrididae). Chromosoma, 24: 10-16.
94.
Sez FA (1968): On the heterochromatinization process in the X-autosome fusion of some orthopteran
insects (Session V: Induced changes in chromosomes). Intl Sem Calcutta, India, p. 16.
95.
Sez FA (1968): The extraordinary karyotype of Dichroplus silveiraguidoi: a possible explanation of its
origin. Intl Cong Genet, Tokyo, 1: 198.
96.
Brum N & Sez FA (1969): Los cromosomas de varias especies de Hylidae (anfibios anuros). Cong LA
Zool, Caracas, p. 62.
97.
Sez FA & Prez-Mosquera G (1969): Posibles mecanismos evolucionarios en la formacin de algunas
especies crticas de Dichroplus (Orthoptera: Acrididae). 21 Reunio An Soc Bras Progr Cin, So Paulo, p. 247.
98.
Knapper CFU & Sez FA (1970): Estudo citolgico das espcies mais freqentes do gnero Pheretima no
Ro Grande do Sul. 22 Reunio An Soc Bras Progr Cin, Salvador de Bahia, p. 295.
99.
Peixoto LI & Sez FA (1970): Estudo da estructura citogentica de uma populao de Laplatacris dispar.
22 Reunio An Soc Bras Progr Cin, Salvador de Bahia, p. 95.
100. Barrio A, Sez FA & Rinaldi-Chieri PR (1971): The cytogenetics of Chthonerpeton indistinctum
(Amphibia: Gymnophioma). Caryologia, 24: 435-445.
101. Brum N & Sez FA (1971): Chromosomes of South American Bufonidae (Amphibia: Anura). Experientia,
27: 470-471.

87

102. Sez FA, Drets ME & Brum N (1971): Karyotype of the Carpincho Hydrochaeris hydrochaeris
uruguayensis (Rodentia: Hydrochaeridae). 5 Cong LA Zool, Montevideo, 1: 187-191; Experientia, 27(5): 584585.
103. Sez FA & Prez-Mosquera G (1971): Citogentica del gnero Dichroplus (Orthoptera: Acrididae). En:
Meja RM & Moguilevsky JA (eds.): Recientes adelantos en biologa. Bona, Buenos Aires, pp. 111-120.
104. Kiblisky P, Brum N, Prez-Mosquera G & Sez FA (1972): Variabilidad cromosmica en diversas
poblaciones de roedores del gnero Ctenomys (Rodentia: Octodontidinae) en el Uruguay. Cin e Cult, So Paulo,
24: 208.
105. Sez FA (1972): El proceso meitico en Dichroplus silveiraguidoi (Orthoptera: Acrididae). Soc Arg Gent,
p. 23.
106. Sez FA (1972): Correlaciones entre conversin y heterocromatinizacin en el sistema neo-X neo-Y. 3
Reunin Anual Soc Arg Gent, p. 9.
107. Sez FA & Venegas W (1972): A cytogenetic study of Rhinoderma darwini (Amphibia: Anura). Chrom Inf
Serv, Tokyo, 13: 11-15.
108. Brum N & Sez FA (1973): Chromosomes of South American Bufonidae (Amphibia: Anura). Caldasia,
Colombia, 11: 51-61.
109. Cardoso H, Sez FA & Brum N (1973): Localizacin y comportamiento de la heterocromatina constitutiva
en Dichroplus silveiraguidoi (Acrididae: Orthoptera). Soc Arg Gent, p. 22.
110.
36.

Sez FA (1973): El sistema sexual neo-X neo-Y de Spathalium audouini. Reunin Anual Soc Arg Gent, p.

111. Sez FA, Drets ME & Brum N (1973): Cromosomas somticos y meiticos del carpincho Hydrochaeris
hydrochaeris uruguayensis (Rodentia Hydrochaeridae). Cong LA Zool, Montevideo, 1: 187-191.
112. Sez FA & Venegas W (1973): Estudio citogentico de Rhinoderma darwini (Amphibia: Anura). 5 Cong
LA Zool, Montevideo, 1: 181-186.
113. Cardoso H, Sez FA & Brum N (1974): Location, structure and behaviour of C-heterochromatin during
meiosis in Dichroplus silveiraguidoi (Acrididae: Orthoptera). Chromosoma, 48: 51-64.
114. Cosen RH & Sez FA (1974): Cariotipo y sistema neo-X neo-Y en Dichroplus vittatus (Acrididae:
Orthoptera). Physis, 33: 237-242.
115. Cardoso H, Sez FA & Brum N (1975): La heterocromatina constitutiva en Dichroplus silveiraguidoi
(Acrididae: Orthoptera). Cin e Cult, 27: 420-426.
116. Hunziker JH & Sez FA (1976): Citogentica y gentica evolutiva vegetal y animal en la Argentina.
Historia y Bibliografa (1923-1972). Bol Acad Nal Cien, Crdoba, 51: 283-323.

88

117. Kiblisky P, Brum N, Prez G & Sez FA (1977): Variabilidad cromosmica entre diversas poblaciones
uruguayas del roedor cavador del gnero Ctenomys (Rodentia: Octodontidae). Mendeliana, 2: 85-93.
118. Sez FA & Prez-Mosquera G (1977): Structure, behaviour and evolution of the chromosomes of
Dichroplus silveiraguidoi (Orthoptera: Acrididae). Genetica, 47: 105-113 (trabajo pstumo).

Figure 43 -El Prof. Sez making chromosome observations

with a large optical microscope available in the Laboratory of Microscopy
of the Andes University, Mrida, Venezuela, (circa September, 1968).

Book Translations
119. White MJD (1947): Los cromosomas.Sez A (trad.), Col. Nueva ciencia -Nueva tcnica, Espasa Calpe,
Buenos Aires, 146 pp. [Trad. de: Chromosomes. 2 ed., Saunders, Philadelphia, 1942].
120. Darlington CD (1948): La evolucin de los sistemas genticos. Sez A (trad.), Espasa Calpe, Buenos Aires,
197 pp. [Trad. de: The evolution of genetic systems.Cambridge Univ. Press, 1939].
121. White MJD (1951): Citologa animal y evolucin. Sez A (trad.), Espasa Calpe, Buenos Aires, 511 pp.
[Trad. de: Animal cytology and evolution. Cambridge Univ. Press, 1945].

89

Reviews of scientific publications, conferences, and educational articles

Professor Sez not only conducted research, but also dealt with often draw a large number of popular articles on
various issues related to cytogenetics, as well as scientific review articles and commentaries on the life and work
of their colleagues.
122. Sez A (1925): Variacin y herencia en los animales domsticos y las plantas cultivadas, por Jos
Fernndez Nondez, Madrid, 1922, 200 pp. Rev Centro Est Med Veter Univ Nal La Plata, 4(2): 107-108.
123.

Sez A (1927): Museo de La Plata. Physis, 8(31).

124.

Sez A (1927): Doctorado en Ciencias Naturales en el Instituto del Museo de La Plata. Physis, 8(31).

125. Sez FA (1927): La Herencia Mendeliana y su fundamento citolgico. Conferencia pronunciada en el

Saln de Actos del Museo Pedaggico de Montevideo (2 de febrero).
126. Sez FA (1927): El mecanismo de la Herencia Mendeliana. Conferencia dictada en el Laboratorio de
Ciencias Biolgicas del Consejo Nacional de Enseanza del Uruguay.
127.

Sez A (1927): Sociedad Uruguaya de Ciencias Naturales. Physis, 8.

128.

Sez A (1927): Congreso Internacional de Gentica y de Zoologa. Physis, 8.

129.

Sez A (1927): Nueva revista de bibliografa Biological Abstracts. Physis, 8(31).

130. Sez FA (1928): La Biologa Celular a la luz de las ms modernas investigaciones. Conferencia dictada en
el Laboratorio de Ciencias Biolgicas del Consejo Nacional de Enseanza del Uruguay.
131. Sez FA (1928): La determinacin del sexo, la intersexualidad y la herencia ligada al sexo. Conferencia
dictada en el Laboratorio de Ciencias Biolgicas del Consejo Nacional de Enseanza del Uruguay.
132. Sez FA (1929): La herencia y sus manifestaciones en el individuo. Conferencia por invitacin de la
Comisin Pro-fomento de la Escuela N 105, Montevideo.
133. Sez FA (1929): Herencia y evolucin en la Biologa moderna. Conferencia dictada en el Saln de Actos
de la Universidad de Mujeres, Montevideo.
134. Sez FA (1929): Algunos grandes problemas de la Biologa. Conferencia dictada en la Seccin
Universitaria de la Asociacin Cristiana de J-venes, Montevideo.
135. Sez FA (1930): Explicacin tcnica de la Conferencia del Director del Laboratorio de Zoologa de la
Universidad de Pennsylvania, Prof. G.E. McClung. Conferencia dictada en la Facultad de Ciencias Mdicas,
Buenos Aires.
136.

Sez A (1931): Albina E. Bonjour.Crnica. Physis, 10: 435-436.

90

137. Sez FA (1932): Los artesanos del sexo y de la herencia. Conferencia dictada en el Colegio Bartolom
Mitre de La Plata, resumida en El Da, La Plata.
138. Sez FA (1932): Complicacin del problema sobre determinismo del sexo y la moderna teora de la
herencia. Conferencia dictada en el Colegio Bartolom Mitre de La Plata, resumida en El Da, La Plata.
139. Sez FA (1933): Sexo, intersexualidad y herencia en la Biologa contempornea. Conferencia dictada en la
Asociacin Sarmiento, La Plata.
140. Sez FA (1934): Vida y obra de Santiago Ramn y Cajal. Conferencia dictada en el Ateneo de la
Asociacin Juan Agustn Garca (7 de noviembre).
141. Sez A (1934): Thomas H. Morgan, Premio Nobel 1933. El Da, La Plata, ao 10, n 303. Artculo
transcripto en La Maana, Montevideo, n 5908 (13 de enero).
142. Sez A (1934): La Gentica: la ms novel de las Ciencias Biolgicas. Estados de los estudios en Argentina
y Uruguay. El Pueblo (reportaje), Montevideo (26 de febrero).
143.

Sez A (1934): Dr. ngel Gallardo: su personalidad de hombre y de sabio. Rev Arg Agron, 1: 138-145.

144. Sez FA (1935): Panorama de la Biologa a medio siglo de la muerte de Mendel. Conferencia dictada en la
Estacin Radiotelefnica, Universidad Nacional de La Plata.
145. Sez FA (1935): Problemas actuales de la Biologa. Conferencia dictada por invitacin del Centro de
Estudiantes del Doctorado en Ciencias Naturales, Universidad de Buenos Aires.
146.

Sez A (1935): Homenaje al Profesor Houssay. Physis, 11: 528-530.

147.

Sez A (1935): Santiago Ramn y Cajal (1852-1934). Physis, 11: 550-552.

148. Sez FA (1936): Una era nueva en el estudio de las Ciencias Naturales. Conferencia publicada en An Soc
Cient Arg, 121: 291.
149.

Sez FA (1936): El ejemplo de Ameghino. Conferencia dictada en el Instituto Juan A. Garca, La Plata.

150. Sez A (1937): El porvenir de las Ciencias Naturales en el Uruguay. El Debate, Montevideo (19 de
febrero).
151. Sez FA (1937): Herencia y evolucin en la Biologa contempornea. Conferencia dictada en el Museo de
La Plata.
152.

Sez A (1937): Las leyes de la herencia y el sexo. La Razn, Buenos Aires.

153. Sez A (1937): Transcripto en Curso de Zoologa Argentina por J.L. Liebermann. Buenos Aires (19 de
agosto).
154. Sez FA (1938): Florentino Ameghino y la vocacin cientfica. Conferencia dictada en la Escuela Normal
Nacional de La Plata, con motivo de la celebracin del cincuentenario de dicha institucin.

91

155. Sez FA (1938): Origen, determinacin y herencia del sexo. Conferencia dictada en el Centro y Biblioteca
Alborada, La Plata.
156. Sez FA (1938): ltimos descubrimientos relacionados con la biologa del sexo. Conferencia dictada por
invitacin del Centro de Estudiantes de Ciencias Naturales, en el Museo de La Plata.
157. Sez A (1939): Lisandro de la Torre, su vocacin cientfica. Cursos y Conferencias, Homenaje a Lisandro
de la Torre, Rev Colegio Libre de Estudios Superiores, ao VIII, n 9, v. 15.
158. Sez A (1941): La meiosis y la determinacin del sexo. En: Calatroni JC & Ruiz V (eds.): Teraputica
ginecolgica. 2 ed., El Ateneo, Buenos Aires.
159.

Sez A (1943): La vocacin de Ameghino. El Da, La Plata, p. 7 (25 de mayo).

160. Sez FA (1943): Florentino Ameghino y su vocacin por la ciencia. Alas, Rev Ilustrada Chivilcoy,
Argentina, 3: 1-4.
161.

Sez A (1944): El gene: maravilla invisible. El Mundo, Buenos Aires (2 de agosto).

162.

Sez A (1944): Molcula u organismo? El Mundo, Buenos Aires (12 de diciembre).

163. Sez FA (1944): E. Garber. Spontaneous alterations of chromosome morphology in Nothoscordum fragans
(Am J Bot, 31: 161-165).Rev Arg Agron, 4: 336-340.
164.

Sez FA (1944): Thomas Hunt Morgan (1866-1945). Obituario. Cien e Inv, 2: 83-85.

165. Sez FA (1945): Algunas conquistas recientes de la biologa. Conferencias del ciclo 1942 dictadas por los
becarios. Com Nal Cultura, Buenos Aires, 5: 71-159.
166.

Sez FA (1945): El mecanismo de la herencia humana(por Combes TC). Cien e Inv, 1: 19-20.

167.

Sez FA (1945): Accin de las bajas temperaturas sobre la qumica del cromosoma. Cien e Inv, 1: 24.

168.

Sez FA (1945): Una alteracin intraespecfica espontnea de los cromosomas. Cien e Inv, 1: 39-41.

169.

Sez FA (1945): Un nuevo alcaloide capaz de inducir la poliploida. Cien e Inv, 1: 74-75.

170.

Sez FA (1945): La actividad de los cromosomas inertes y el cncer. Cien e Inv, 1: 121.

171.

Sez FA (1945): Partenognesis y determinacin del sexo. Cien e Inv, 1: 271-272.

172.

Sez FA (1946): Hay posibilidad de producir hijos de un determinado sexo? Cien e Inv, 2: 434-435.

173.

Sez FA (1946): Hermann J. Muller: Premio Nobel de Medicina y Fisiologa. Cien e Inv, 3: 538-540.

174. Sez FA (1947): Los fenmenos qumicos de la divisin celular: el cido nucleico y la iniciacin de la
meiosis. Cien e Inv, 3: 510-511.
175.

Sez FA (1947): Consejos a los agrnomos (comentario). Cien e Inv, 1: 25.

92

176. Sez FA (1947): Un catlogo sobre el nmero de cromosomas en los vegetales (por C.D. Darlington). Cien
e Inv, 3: 243.
177. Sez FA (1948): Influencias de los insecticidas gamexane y DDT sobre el mecanismo de la divisin
celular. Cien e Inv, 4: 330-331.
178.
208.

Sez FA (1949): Es posible inducir mutaciones hereditarias con sustancias carcingenas? Cien e Inv, 5:

179.

Sez FA (1949): Un diccionario de gentica(por R.L. Knight). Cien e Inv, 5: 245.

180.

Sez FA (1949): Investigaciones genticas sobre la accin de la bomba atmica. Cien e Inv, 5: 289-290.

181.

Sez FA (1949): Microscopa ultravioleta de la clula tumoral viviente. Cien e Inv, 5: 331-332.

182.

Sez FA (1949): Cul es el momento ms sensible de los cromosomas a los rayos X? Cien e Inv, 5: 333.

183. Sez FA (1949): Habla para Accin el Prof. Francisco A. Sez, autor de un libro de xito en Estados
Unidos y Europa. Ciencias Biolgicas. Accin 1, 201: 3.
184. Sez FA (1950): Una tcnica para el estudio de los cromosomas somticos largos en los vegetales. Cien e
Inv, 6: 281-282.
185.

Sez FA (1950): El contenido nucleoproteico del ncleo celular. Cien e Inv, 6: 319-320.

186.

Sez FA (1950): Gentica y evolucin. Cien e Inv, 6: 531-539.

187.

Sez FA (1951): La estructura gentica de Schistocerca cancellata. Cien e Inv, 7: 171-172.

188.

Sez FA (1951): Anlisis cromatogrfico de los aminocidos del cromosoma. Cien e Inv, 7: 172.

189.

Sez FA (1951): Cmo se comporta el cido nucleico en la divisin celular? Cien e Inv, 7: 411-413.

190. Sez FA (1953): La fragmentacin nuclear como fenmeno normal del ciclo mittico de la clula
cancerosa. Cien e Inv, 9: 224.
191.

Sez FA (1953): Metabolismo de los cidos nucleicos en las clulas tumorales. Cien e Inv, 9: 225-226.

192.

Sez FA (1954): Charles Leonard Huskins (necrologa). Cien e Inv, 10: 336.

193.

Sez FA (1954): Citogentica: una nueva ciencia. Cien e Inv, 10: 552-554.

194.

Sez FA (1956): El gene, maravilla invisible de la Biologa.Rev Inst Est Sup, Montevideo, 1: 72-80.

195. Sez FA, Drets ME & Novizki I (1958): Mestres de renome internacional proferiro hoje conferncias no
Conclave de Gentica. I Reunio de Gentica. Diario do Paran, Curitiba, p. 8 (14 de noviembre).
196. Sez FA, Novizki I & Drets ME (1958): Geneticistas, cientistas da vida, so a favor do aplo de Pauling.
O Estado do Paran, Brasil (16 de noviembre).

93

197. Sez FA (1959): Homenaje al Dr. Alberto Boerger. Homenaje al Ing. J. Fischer. Arch Soc Biol
Montevideo, 24: 13.
198. Sez FA (1959): Reflexiones sobre mutacin: es posible modificar la naturaleza hereditaria de un
individuo? Rotauruguay, 26: 13-19.
199. Sez FA (1959): Algunos rasgos de la personalidad cientfica de Clemente Estable. Rev Inst Est Sup,
Montevideo, 4: 534-543.
200.

Sez FA (1959): Biologa de la mutacin. Marcha, Montevideo, XXI, 3 Secc., 992: 7 (31 de diciembre).

201. Sez FA (1959): La radiacin atmica puede provocar muy graves efectos de tipo gentico. El Pas,
Montevideo, p. 5 (12 de marzo).
202. Sez FA, Drets ME, Solari CL, Brum N, Daz & Laguardia AM (1959): La radioactividad: suicidio del
mundo. Un problema sin fronteras, minuto a minuto. Marcha, Montevideo, XXI, 969: 32 (24 de julio).
203. Sez FA, Drets ME, Solari CL, Brum N, Daz & Laguardia AM (1959): Nuestros hombres de ciencia
opinan. Marcha (carta al director), XXI, 969: 32 (24 de julio).
204.

Sez FA (1960): El drama gentico de la era atmica.Com Nal Prot Radiac, Uruguay JUS, pp. 1-6.

205. Sez FA (1960): Amenazan las radiaciones dar formas monstruosas al hombre. La Maana, Montevideo,
44: 4 (1 de agosto).
206. Sez FA (1961): Son criminales las pruebas atmicas? Habla el Prof. Francisco Sez. La Maana,
Montevideo, pp. 3 (12 de setiembre).
207. Sez FA (1962): Las radiaciones y el destino del hombre.Conferencias radiales (SODRE). Bol Com Nal
Prot Radiac, pp. 3-4.
208. Sez FA (1962): Esta pesadilla. Sez: un testimonio contra las explosiones. Marcha (reportaje de Eduardo
H. Galeano), Montevideo, XXIII, 1110: 32 (8 de junio).
209. Sez FA (1962): El Departamento de Citogentica del Instituto de Investigacin de Ciencias Biolgicas.
Bol Soc Bras Genet, Curitiba, 4: 144-150.
210. Sez FA (1963): Alteraciones hereditarias inducidas por las radiaciones. Conferencias radiales (SODRE).
Bol Com Nal Prot Radiac, pp. 5-6.
211. Sez FA (1964): Instituto de Ciencias Biolgicas. Departamento de Citogentica. El Plata, Ediciones del
Cincuentenario, Montevideo, 9 pp.
212. Sez FA (1965): Un Simposio sobre la funcin del nuclolo (en el Instituto de Ciencias Biolgicas):
International Symposium The Nucleolus, its structure and function, Montevideo, Uruguay. Hechos, p. 5 (1 de
diciembre).

94

213. Sez FA (1966) (ed. asoc.): International Symposium The nucleolus, its structure and function (held at
Montevideo, Uruguay, December 5-10, 1965). National Cancer Institute Monograph 23, U.S. Dept. of Health,
Education, and Welfare, Public Health Service, Bethesda, 610 pp.
214. Sez FA (1973): Historia de la biologa en Amrica Latina con especial referencia al desarrollo de la
gentica. Conf Interam Enseanza Biol, Asuncin, pp. 82-88.
215. Sez FA (1974): Cmo se forma una nueva especie en la naturaleza. (origen del sorgo negro descubierto
por un investigador uruguayo). El Da, Supl, v. 41, 2135: 8-9 (30 de junio).

Articles on Professor Sez

216. Comisin Nacional de Cultura (1937): Becados en el pas. Francisco Alberto Sez. Comisin Nacional de
Cultura, su labor en 1936. Buenos Aires, p. 64.
217. Comisin Nacional de Cultura (1940): Francisco Alberto Sez. Comisin Nacional de Cultura, su labor en
1939. Buenos Aires, p. 31.
218. Asociacin Uruguaya para el Progreso de la Ciencia (1949): Instituciones Cientficas y Cientficos del
Uruguay. Francisco Alberto Sez. Asociacin Uruguaya para el Progreso de la Ciencia, Montevideo, 1: 62-64;
Instituciones Cientficas y Cientficos Latinoamericanos. Unesco, Mon-tevideo, 2: 205-206 (1953); idem, 31: 1312 (1965).
219. La Estanzuela (1949): Sez FA, Espectrofotometra del cromosoma. Hoja Gentica, Colonia, 1: 9; idem, 1:
10-12 (1953).
220. Argone C (1968): Francisco Alberto Sez. En: Garibaldi CA (ed.): Vidas admirables: biografas y
semblanzas. Consejo Nacional de Enseanza Primaria y Normal, Montevideo, pp. 95-97.
221.

Conteris H (1968): Sez a los 70. Marcha, XXIX, 1397: 11 (5 de abril).

222. De Robertis EDP (1969): Discurso pronunciado por el Acadmico Prof. Dr. Eduardo D.P. de Robertis en
presentacin del Acadmico Correspondiente Prof. Francisco A. Sez, en el acto de entrega del Diploma de
Acadmico Correspondiente en la Academia Nacional de Ciencias Exactas, Fsicas y Naturales de la ciudad de
Buenos Aires, Repblica Argentina, el sbado 4 de mayo de 1963. En:UNESCO (ed.): Homenaje al Dr. Francisco
A. Sez en su 70 aniversario. Arch Soc Biol Montevideo, Nmero extraordinario, 27: 4-6.
223. Drets ME (1969): Discurso pronunciado por el Dr. Mximo Eduardo Drets en ocasin del acto de entrega
del Ttulo de Profesor Ad-Honorem al Prof. Francisco A .Sez, en representacin de sus discpulos, en el Saln de
Actos de la Facultad de Medicina de Montevideo, Repblica Oriental del Uruguay, el viernes 15 de marzo de
1968. En: UNESCO (ed.): Homenaje al Dr. Francisco A. Sez en su 70 aniversario. Arch Soc Biol Montevideo,
Nmero extraordinario, 27: 7-12.
224. UNESCO (1969) (ed.): Homenaje al Dr. Francisco A. Sez en su 70 aniversario. Arch Soc Biol
Montevideo, Nmero extraordinario, v. 27, 152 pp.

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225. Fundacin Lucio Cherny, Argentina, honra al Dr. Sez (1974): Ciencia Interamericana, Washington D.C.,
15: 13.
226. Santos Prez M (1998): Hace 100 aos naca un sabio: Dr. Francisco Alberto Sez. Entrega 2000, Temas,
p. 16.

Papers from other authors reviewed by Sez

227. Fernndez Nondez J (1923): Variacin y herencia en los animales domsticos y las plantas cultivadas.
Bibl. Agrcola Espaola, Espasa Calpe, Madrid. [Reseado en: Rev Centro Est Med Veter Univ Nal La Plata, 4:
42-81, 1925].
228. Metalnikov S: Sobre herencia de inmunidad adquirida. [Reseado en: Rev Centro Est Med Veter Univ Nal
La Plata, 4: 106-107, 1925].
229. King RL (1923): Heteromorphic homologous chromosomes in three species ofPseudotrimerotropis
(Orthoptera: Acrididae). J Morph, 38(1): 19-63. [Reseado en: Physis, 8: 666-667, 1927].
230. Oguma K & Kihara H (1923): tude des chromosomes chez lhomme. Arch Biol, 33: 493-515. [Reseado
en: Physis, 8: 667-668, 1927].
231. Painter ThS (1924): The sex chromosomes of man. Am Nat, 58(659): 506-524. [Reseado en: Physis, 8:
668-669, 1927].
232. Maln DJ & Maln DE: The spermatogenesis of Locustana pardalina (Walker),the Brown treck locust.
[Reseado en: Physis, 8: 669-670, 1927].
233. Causey D (1925): Mitochondria and Golgi bodies in Entamoeba gingivalis (Gros) Brumpt and
mitochondria in Leishmania brasiliensis. University of California Press, 28 pp. [Reseado en: Physis, 8: 670-671,
1927].
234. De Winiwarter H & Oguma K (1926): Nouvelles recherches sur la spermatogense humaine. Arch Biol,
36: 99-166. [Reseado en: Physis, 8: 671-673, 1927].
235. Walker CE (1926): The meiotic phase in certain mammals. Proc R Soc Lond B, 99(698): 366-374.
[Reseado en: Physis, 8: 673-674, 1927].
236. Harman MT & Root FP (1926): Number and behaviour of the chromosomes in Cavia cobaya (the common
guinea pig). Biol Bull, 51(2): 73-84. [Reseado en: Physis, 8: 674-675, 1927].
237. Altmann SCA & Ellery MEW (1925): The chromosomes of four species of marsupials. Quart J Micr Sci,
69: 463-470. [Reseado en: Physis, 8: 675, 1927].
238. Hargitt GT (1926): The formation of the sex glands and germ cells of mammals. II. The history of the male
germ cells in the albino rat. J Morph, 42(1): 253-305. [Reseado en: Physis, 8: 675-676, 1927].
239. Palmer R (1926): The chromosome complex of Gammarus chevreuxi, Sexton. I. Spermatogenesis. Quart J
Micr Sci, 70: 541-551. [Reseado en: Physis, 8: 676-677, 1927].

96

240. Gallstegui C (1926): Nmero de cromosomas en algunas especies del gnero Brassica. Bol R Soc
Espaola Hist Nat, 26: 185-191. [Reseado en: Physis, 8: 677, 1927].
241. Nadson GA & Rochline-Gleichgewicht EJ (1926): Le chondriome est la partie de la cellule la plus sensible
aux rayons X. C R Soc Biol, 95: 378-380. [Reseado en: Physis, 8: 677-678, 1927].
242. Bowen RH (1926): Notes on the form and function of the Golgi apparatus in striated muscle. Biol Bull,
50(2): 108-116. [Reseado en: Physis, 8: 678, 1927].
243. Bowen RH (1926): The Golgi apparatus: its structure and functional significance. Anat Rec, 32(2): 151193. [Reseado en: Physis, 8: 678-679, 1927].
244. Guilliermond A (1926): Sur laction des mthodes imprgnation osmique sur les cellules vgtales:
nouvelle contribution ltude de lappareil de Golgi. C R Soc Biol, 95: 442-445. [Reseado en: Physis, 8: 679680, 1927].
245. Heilbrunn LV (1926): The centrifuge method of determining protoplasmic viscosity. J Exp Zool, 43(2):
313-320. [Reseado en: Physis, 8: 680, 1927].
246. Heilbrunn LV (1926): The absolute viscosity of protoplasm.J Exp Zool, 44(1): 255-278. [Reseado en:
Physis, 8: 680-681, 1927].
247. Fetter D (1926): Determination of the protoplasmic viscosity of Paramecium by the centrifuge method. J
Exp Zool, 44(1): 279-283. [Reseado en: Physis, 8: 681, 1927].
248. Popovici H (1926): Nouvelle mthode de coloration du noyau par le vert Janus. C R Soc Biol, 94: 991.
[Reseado en: Physis, 8: 681-682, 1927].
249. Horovitz S (1926): Estudio de cromosomas durante la formacin del polen.Rev Centro Est Agron Vet
UBA. [Reseado en: Physis, 8: 682-683, 1927].
250. Lahille F (1927): La clula: breve resea histrica y aparatos generales(apuntes). Rev Centro Est Agron
Vet UBA, 131: 7-19. [Reseado en: Physis, 8: 683, 1927].
251. Hance RT: El sexo y los cromosomas en la gallina domstica (Gallus domesticus). [Reseado en: Physis,
8: 683-684, 1927].
252. Guynot (1930): La variation et lvolution. Vol. 1. La variation. Col. Encyclopdie scientifique, Doin
diteurs, Paris, 457 pp. [Reseado en: Physis, 10: 343, 1931].
253. Patterson JT (1929): The production of mutation in somatic cells of Drosophila melanogaster by means of
X-rays!J Exp Zool, 53(3): 327-372. [Reseado en: Physis, 10: 344, 1931].
254. Wilhelm O (1927): La Rhinoderma Darwini D. y B. Bol Soc Biol Concepcin, 1(1-2): 11-39. [Reseado
en: Physis, 10: 347, 1931].
255. Lipschtz A (1927): El experimento de la hiperfeminizacin de Steinach. Bol Soc Biol Concepcin, 1(1-2):
41. [Reseado en: Physis, 10: 348, 1931].

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256. Baur E (1923?): Las bases cientficas de la fitotecnia: un texto para agrnomos, horticultores y
silvicultores. Fischer GJ (trad.), Palacio del Libro, Montevideo, 140 pp. [Reseado en: Physis, 10: 349, 1931].
257. McClintock B (1929): A method for making aceto-carmin smears permanent.Stain Tech, 4(2): 53-56.
[Reseado en: Physis, 10: 349, 1931].
258. Rosas Costa JA (1930): Literatura nacional y extranjera producida sobre ortpteros argentinos.Mero Jard
Zool La Plata, 2: 72-91. [Reseado en: Physis, 10: 350-351, 1931].
259.

Gates RR (1932): Nuclear structure. J R Micr Soc, 52(1): 1-19. [Reseado en: Physis, 11: 200, 1932].

260. Makino S (1931): The chromosomes of Diestrammena japonnica (Karny) (an orthopteran). Doubutsugaku
zasshi, 43(517): 635-646. [Reseado en: Physis, 11: 200-201, 1932].
261. Hareyama S (1932): On the spermatogenesis of an orthopteran, Gampsocleis brgeri (D. H.). J Sci
Hiroshima Univ Ser B, 1: 91-143. [Reseado en: Physis, 11: 201, 1932].
262. Carothers EE (1931): The maturation divisions and segregations of heteromorphic homologous
chromosomes in Acrididae (Orthoptera). Biol Bull, 61(3): 324-349. [Reseado en: Physis, 11: 201-202, 1932].
263. Matthey R (1931): Chromosomes des reptiles sauriens, ophidiens, chloniens: lvolution de la formule
chromosomiale chez les Sauriens. Rev Suisse Zool, 38(9): 117-186. [Reseado en: Physis, 11: 202-203, 1932].
264. Robertson WRB (1931): Chromosome studies. II. Synapsis in the Tettigidae, with special reference to the
presynapsis split. J Morph, 51(1): 119-145. [Reseado en: Physis, 11: 203-204, 1932].
265. Babcock EB (1931): Cyto-genetics and the species concept. Am Nat, 65(696): 5-18. [Reseado en: Physis,
11: 204-205, 1932].
266. Gates RR & Catcheside DG (1931): Origin of chromosome linkage in Oenothera. Nature, 128: 637.
[Reseado en: Physis, 11: 205-206, 1932].
267. Darlington CD (1932): Chromosomes and plant-breeding. Macmillian & Co. Ltd., London, 112 pp.
[Reseado en: Physis, 11: 206, 1932].
268. Gates RR (1931): The cytological basis of mutations. Am Nat, 65(697): 97-120. [Reseado en: Physis, 11:
206-207, 1932].
269. Winge (1932): On the origin of constant species-hybrids. Svensk Bot Tidskr, 26: 107-122. [Reseado en:
Physis, 11: 207, 1932].
270. Frandsen HN & Winge (1932): Brassica napocampestris, a new constant amphidiploid species hybrid.
Hereditas, 16(1-2): 212-218. [Reseado en: Physis, 11: 207, 1932].
271. Randolph LF (1932): Some effects of high temperature on polyploidy and others variation in maize. PNAS,
18(3): 222-229. [Reseado en: Physis, 11: 207-208, 1932].

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272. Beadle GW (1932): A possible influence of the spindle fibre on crossing-over in Drosophila. PNAS, 18(2):
160-165. [Reseado en: Physis, 11: 208, 1932].
273.

Gallardo A (1931): Oruga mimtica del rosal. Piselli, Buenos Aires. [Reseado en: Physis, 11: 211, 1932].

274. Gallardo A (1931): Deux nouvelles espces de Pogonomyrmex de la Rpublique Argentine

(Hymenoptres formicides). Rev Mus La Plata, 33: 185-188. [Reseado en: Physis, 11: 211, 1932].
275. Gallardo A (1932): Las hormigas de la Repblica Argentina. Subfamilia Mirmicinas. Seccin
Promyrmicinae. An Mus Nal Hist Nat Bernardino Rivadavia, 37: 37-87. [Reseado en: Physis, 11: 211-212,
1932].
276. Garber E (1944): Spontaneous alterations of chromosome morphology in Nothoscordum fragans. Am J
Bot, 31: 161-165. [Reseado en: Rev Arg Agron, 11: 336-340, 1944].
277. Combes TJC (1945): El mecanismo de la herencia humana. Ruiz, Rosario, 319 pp. [Reseado en: Cien e
Inv, 2: 19-20, 1946].
278. Boerger A (1946): Agronoma: consejos metodolgicos. El Ateneo, Buenos Aires, 538 pp. [Reseado en:
Cien e Inv, 3: 25-26, 1947].
279. Darlington CD & Janaki Ammal EK (1945): Chromosome atlas of cultivated plants. Allen & Unwin Ltd.,
London, 397 pp. [Reseado en: Cien e Inv, 3: 243-244, 1947].
280. Knight RL (1948): Dictionary of genetics. Chronica Botanica Co. Waltham, Massachusetts, 200 pp.
[Reseado en: Cien e Inv, 5: 245, 1949].
281. Matthey R (1949): Les chromosomes des vertbrs. Librairie de lUniversit F. Rouge, Lausanne, 356 pp.
[Reseado en: Cien e Inv, 6: 71-72, 1950].
282.

Brncic D: Un estudio moderno sobre sistemtica de Drosophila. [Reseado en: Cien e Inv, 14: 26, 1958].

283. Brachet J (1957): Biochemical cytology. Academic Press Inc., New York, 516 pp. [Reseado en: Cien e
Inv, 14: 124-125, 1958].
284. Turpin R & Lejeune J (1965): Les chromosomes humains. Gauthier-Villars, Pars, 535 pp. [Reseado en:
Cien e Inv, 22: 25-26, 1966].

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Figure 44.Sez celebrating his 78th birthday on March 10th 1976,with his disciples and colleagues. Top Row, left
to right:Horacio Cardoso, Mario Stoll, Berta Oliveras, NadirBrum,Hctor Navarrete,Gustavo Folle, Alicia
Arias, Mximo Drets;Bottom Row: lvaro Novello, Mnica Brauer,Prof. Sez, Gloria Martnez-Drets.

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Figure 45 Last picture of Professor Sez accompanied by his disciple Drets

taken on March 10th, 1976, during his 78 anniversary celebrated
at the Institute of Biological Investigations Clemente Estable.

OBITUARY
Francisco Sez Alberto passed away unexpectedly on April 27, 1976. His disappearance caused great
consternation in the scientific community, as well as its reputation as a professional, was a beloved figure. His
contributions to Science will remain forever attached for those who knew him to the memory of a great personality
and for,many,of noble and unforgettable friend.
285

Barrio OA, Vidal R (1976) Profesor Dr. Francisco Alberto Sez. Necrolgica. Physis, 90:197-203

286

Beiguelman B (1976) Francisco Alberto Sez. Cincia e Cultura. So Paulo, 2:1372-1374

287

EL DIA (1976) Obituario. Prof. Dr. Francisco Alberto Sez, Abril 28, pp.7

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288
Hunziker JF (1976) Francisco Alberto Sez y su contribucin al desarrollo de la citogentica rioplatense.
Mendeliana, 1:69-74

ADDENDA

THE MASTER AND HIS SCIENTIFIC LEGACY

Examples of three areas of investigations initiated by Sez
and continued by his disciples.
(1960 - 2007)

Introduction
This section is only intended to illustrate tothe reader how some cytogeneticists disciples of Professor Saez
developed at least three areas of research initiated by him, making original contributions that indicated not only
the thematic continuity of these studies in our laboratory, but they were only possiblebecause of the strong
scientificbackground obtained with him which allowed the continuation of theinvestigation of various aspects of
contemporary cytogenetics and the published contributions which are cited and discussed below, are associated to
this volume and attempt to render hommage to the Masterwho guidedall of us.

(1) Microphotometry
A )Instrument developments
Despite the scientific interest ofSez in developing this new analytical methodology in his laboratory,the
instrument installed was based on imported equipment (Pollister & Moses, 1949) designed for the range of the
visible spectraand not manufactured by a single company but composed of a considerable number of parts
produced by different companies, so it was really difficult to assemble, the instrument artisanal, in a single unit
and also to get it functioning properly. For this reason, wishing to collaborate with Sez we tried to facilitate the
prolonged task of making photometric measurements of cell nuclei performed manually.Therefore, we developed a
more comfortable scanning system,but keeping high measurement precision for analyzing biological materials.

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The tedious practical problem with the operation of the original instrument required that the photometric
measurements of cell nuclei be obtained by moving the objectfordiaphragming the area of interest. By contrast, our
instrument which we called image discriminator apparatus, was based on the use of a movable mirror(Fig. 46) that
allowed exploring the image of the microscopic object without moving it, which greatly facilitated the selection of
the areas to be measured (less than 0,5m, depending on the final optical magnification) and analyzed with a fixed
diaphragm [289].

Figure 46.The image discriminator apparatus for cytophotometry.H indicates the moving mirror for exploring the
cell image generated by the microscope. Note that the object is stationary and the reference beams (sample and
background beams)of the selected object are obtained through a system of mirrors.C and Iare diaphragms for
limiting the area to be measured. Both light beams are collected by the higher mirrors system which arealternately
focused on the photomultiplier.
With the previous Pollister system it was only possible to obtain a few photometric measurements of the
different parts of the same object, for example, a nucleus with an average size of 10 m. This situation is
illustrated in Fig 47a. Fig.47 b shows that the optical scanning using the image discriminator increased the number
of measurements withoutoverlaps which would originate measurement errors. Years later, the replacement of these
systems by photometrical scanning under computer command significantly increased the number of measurements
(Fig. 47, c) providing a higher analytical precision.

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Figure 47Comparative diagram illustrating three systems for measuring the optical density in a cell nucleus.a:
manual method with the Pollister apparatus; b: the nucleus scanned with the image discriminator which yielded a
greater number of step measurements without risk of overlappings, c:computerized scanning using discrete steps
noticing the considerablemeasurements increase obtained and, therefore, ahigher precision of the analysis.
This system was later associated with a high resolution monochromator (Fig. 49, 4) and a proportional
recorder allowing to make analog cytochemical analysis automatically (for example, to obtain DNA curves of
Feulgen stained nuclei). This instrumentation was the first one installed and operated in Latin America [290].
289

Drets M E (1960) Image discriminator apparatus for cytophotometry. Mikroskopie, Wien. 15: 14-20.

290
Drets M E (1961) Ratio recorder for cytophotometry based on an image discriminator apparatus.
Mikroskopie. Wien. 16:341-348.

B )Software for microphotometric quantitative analysis

ofnuclei and chromosomes
The initial old cytospectrophotometer was later replaced in our Laboratory of Quantitative Microscopy by
the (MP01 and MSP65 photometric systems developed by Carl Zeiss, Oberkochen, (Germany). (Fig. 49).
The replacement of the old analog instruments by the Zeiss system which operated under computer
command allowed us to develop thefirst dedicated computer program (Bandscan)(Fig. 48) [291]aimed at exploring
"online" banded chromosomes and obtaining high resolution quantitative chromosome maps on the band locations
because the stage is moved under computer command generating a pixel, or imaging unit, capable of analyzing
areas>0.1 m, with extreme accuracy. Also, we could appliedthis methodto other problems such as replacing
manual count of silver grains in autoradiographs with this rapid automated quantitative analysis with highly
reliable results [292].

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Figure 48. -First quantitative detection of G-bands of the long arm of human chromosome number 1 obtained
using the Bandscan program developed in our laboratory. The use of different levels of sensitivity used,
facilitated the location of the main bands in their relative positions. The resulting chromosome arm diagram on the
right is a drawing. The result is highly accurate, providing a quantitative picture of the chromosome arm structure
first reported in the literature [292].

291
Drets ME Folle GA & Comings DE (1978) Mechanisms of chromosome banding X. Chromosome and
nuclear changes induced by photo-oxidation and their relation to R-banding with anti-C antibodies. Chromosoma,
69, 101-112.
292
Drets ME (1978) Bandscan - A computer program for on-line linear scanning of human banded
chromosomes. Computer Programs in Biomedicine. 8:283-294.

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Figure.49. Incorporation of the Zeiss MP01 scanning cytospectrophotometer associated online to a Wang
programmable calculator and to the microscope system (1975). With this device we could develop the first
computer program reported internationally for the quantitative chromosome analysis [291)]. 1. Electronic
Modules; 2 photomultiplier detector; 3 microscope system with motorized stage for object scanning; 4
monochromator for illumination with wavelengths of the visible spectra; 5, programmable calculator connected
online with the system, 6 , printer of quantitative data obtained during analysis; 7 controls of the photomicroscope

C )Chromosome banding
Since the discovery in 1882 of chromosomes by Flemming28 the descriptions of chromosomes were mainly
confined to recognizing the centromeric constriction and chromosome arms. Caspersson et al discovered in 1968
that DNA-binding fluorescent agent (quinaquine mustard) allowed the identification of all human chromosomes
showing

ineach one a specific fluorescent banding pattern. Using simple methods of incubation in saline

solutions, differential chromosome structures stained with Giemsa stain were also discovered. Later, it was found
that these structures reflected different molecular structures along the chromosome arm. The year 1971 was critical
when it was shown that the centromeric regions were specifically stained showing the C bands, called centromeric
banding. In the same year, performing incubations at different temperature, G bands (Giemsa) and the opposite R
bands (reverse) were also discovered at that time. Figure 50 illustrates the nvestigators that played key roles in
these discoveries in 1971-

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Figure 50 Pioneers of chromosome banding. a-b: (Banding G), M.W. Shaw (1970 photo) and M.E. Drets (1990
photo); c-d: (Banding C) T.C. Hsu (1971 photo) and F.E. Arrighi (1977 photo); and e: (Banding R) B. Dutrillaux
(1966 photo).
293
294

Arrighi FE & Hsu TC (1971): Localization of heterochromatin in human chromosomes. Cytogenetics, 10:
81-86.
Drets ME & Shaw MW (1971): Specific banding patterns of human chromosomes PNAS, 68: 2973-2977.

295

Dutrillaux B & Lejeune J (1971): Sur une nouvelle danalyse du caryotype humain. C R Acad Sci Paris,
272:238-2640.
___________________________
28 - Walther Flemming (Sachsenberg, Mecklenburg At present part of Germany, 1843 - Kiel, Germany,
1905) was a german anatomist and physiologist considered the founder of cytogenetics. He was a pioneer
using staning trchniques to study the cell stuctures. He studied the process of cell division (that he called
mitosis). He used hexomatine for staining the cell nucleus discovering structures intensely stained that during
cell division separated as filaments. Flemming named these stuctures chromatins which later were called
chromosomes. These studies were published in 1882 in his paper Zellsubstanz, kern und zelltheilung (Cellular
substance, nucleus and cell division) (F.C.W. Vogel, Leipzig, 424 pp.

The story of the G banding discovery

In 1969 I was trying to obtain a fellowship from the U.S. National Health Organization that, at that time,
was administered by the Faculty of Medicine of Montevideo under the responsibility of Prof. Roberto Caldeyro
Barcia. During my encounter with Prof. Curt Stern during our visit to Ouro Preto (see Fig 41) invited by the
organizers of the International Symposium of Nuclear Physiology and Differentiation (organized by Alexander
Hollaender) and developed in Belo Horizonte, Brazil, I mentioned to him my interest in perfecting my knowledge
on the human chromosome structure. I mentioned I was looking for a good US Laboratory working in this area. He
immediately said: the best place is the laboratory of Margery Shaw in Houston, Texas. As soon as I returned to
Uruguay I sent a letter to Dr. Shaw explaining hermy scientific interest in becoming familiar with the chromosome
autoradiography method and other techniques for identification of human chromosomes. Instead of replying by
mail she telephoned me telling me: forget the fellowship: I offer you a position of Research Associate for two
years.I confess that this call confused me a little because apparently we were talking about two different things.
But she insisted and I finally accepted her offer. When I arrived in Houston, on the following day I went to the

107

M.D. Anderson Hospital, where Shaws Laboratory was located, to meet her. She was extremely kind to me telling
me: you do not have any compromise with me to develop any kind of research. Even more, you are not obliged to
any daily schedule to work with us The only thing that I ask you is to enjoy your stay in my Laboratoy These
words deeply astonished me increasing my compromise with her as a responsible researcher.
So, I immediately started repeating the techniques used in her laboratory to obtain slides of normal
chromosomes. After a period of about 20 days, I was not really satisfied at all with the quality of chromosomes
that I was obtaining with the methods followed in the laboratory. They appeared as very small structures that were
practically impossible to study. So after a careful analysis of the situation I decided to change the method and I
first started by looking in the lab closets for different chemicals especially crystalline Colchicine trying to avoid
the use of high doses of Colcemid used there, which produced an excessive contraction of the chromosomes. The
second change was a matter of luck. In another place I found a forgotten bottle containing a solution of authentic
German Giemsa MERCK. So the main changes that I introduced were to reduce the time of exposure to the cell
cultures to Colchicine and to use Giemsa from Merck for chromosome staining. Then, I started obtaining beautiful
metaphases with long informative chromosomes very similar to the ones published by Jerme Lejeune who had
taught me in a detailed way how to obtain high chromosome quality during my stay in Laboratory in Paris in 1966.
Using these much improved chromosome preparations I started using the method that had been recently
published by Arrighi and Hsu (1971) [293] for the C banding in that time, using my beautiful chromosomes with
excellent results.
Hsu had participated in a scientific meeting where he learned the discovery of Gall and Pardue [296]. Then,
Hsu asked Gall authorization to send his coworker Francis Arrighi to his laboratory in order to learn their
technique. In the original paper of Gall and Parduethey mentionedthat the centromeric regions were stained
intensensely with their method of formation and detection of RNA-DNA hybrid molecules in cytological
preparations. Returning to her Laboratory, Arrighi mentioned this fact to Hsu.They immediately decided to use he
procedure in human chromosomesfinding the same reaction at the centromeric regions in form of intensely stained
centromeric blocks later named C bands.
The technique included the treatment of chromosomes with NaOH so everybody (including me) used a
glass bottle containing NaOH. After a period of time I noticed the sodium hydroxide was slowly losing his activity
and that its action on the chromosomes progressively was lower. It was under these conditions that I noticed that
chromosomes were losing their capacity to show the centromeric blocks of C-bands but Giemsa staining in the rest
of the chromosome arm reappeared progressively. The C-band procedure was so drastic that in practice produced
empty unstained chromosome arms.

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In these new conditions, one of my slides treated for C banding showed after the saline incubation step a
surprising array of alternate chromosome segments strongly stained by Giemsa. In order to verify if I was looking
real structures, I selected at random the long arms of many normal chromosomes number 1 and started drawing
them. When I was convinced that I was seeing and drawing the same pattern of bands on these chromosome
segments I called Dr. Shaw, who was very impressed by my findings. While still exploring other areas of the slide,
and becoming convinced, she called her husband Charles [Hez] Shaw (the Director of the Section), showing him
that these unknown structures were the result of a specific reaction of the chromosomes. This happened at the
begining of January, 1971. We called the chromosome structures G-bands (or G-banding From Giemsa stain)
[294].
The next problem was to try to find the best treatment to obtain consistent results in all the chromosomes of
the human karyotype from different normal individuals and in both sexes. This task took us a lot of time to get,
and also to have our paper submitted to the PNAS accepted. The first version written by Margery was rejected, so
she asked me to re-write again our contribution and fortunately the second version was finally accepted. The last
anecdote of this story is when I received a reply of the letter that I had sent to Prof. Lejeune describing him our
findings and the main comment was that they had another proceduresimilar to ours. But, ourG bands were the
opposite of theirs. He was referring to the discovery he made with Bernard Dutrilllaux of the Reverse bands (Rbanding) in 1971.!! [295].
296
Gall, J G & Pardue, M L (1969). "Formation and detection of RNA-DNA hybrid molecules in cytological
preparations.". PNAS USA 63 (2): 37883.

D )Microphotometry and chromosome banding

With the discovery of methods of chromosome G-banding we adapted our equipment in order to explore
banded chromosomes for the quantitative location of the densitometric curves peaks allowing to report at an
international level the first quantitative maps of the C- and G banding patterns of human chromosomes [293-303].

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Figure52.-a:Historical human first metaphase found (Houston, 1970) showing G bands.-b:First map of human
female and maleG-bands published by us [294];c: computer graphics image showingthe maximum values of
density peaks and the band-interband regions detected in the long arm of a normal human chromosome number 1
increasing remarkably the quantitative information for each band and, therefore, of the analyzed chromosome arm.
The subsequent addition of a graphic terminal and a modern computer significantly expanded the analytical
capabilities of our microphotometer system allowing the development of other computer programs for a detailed
analysis of chromosome differential structures in humans (Fig. 52, 53) [297-304].
The technology developed by us thus originated very precise chromosome and nuclear images allowing
tovisualize unsuspected structures particularly at the subtelomeric region.

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Figure 53 -Examples of quantitative microfotometric exploration under command of computer programs developed at the
Division of Quantitative Microscopy that achieved new insights into the structural densities of human nuclei and
chromosomes of other species [303].a: graphic image of a normal human chromosome number 13 showing chromatin of
high density in a segment near the centromere; b: the same chromosome seen in pseudo-third-dimension;c: chromocenters in
a nucleus of an insect; d and e: rotation of a human chromosome number 9;f: chromatid exchange. g-iThe system allows
amplification of the terminal bands showing a onelocated in a chromatid higher than the other. Boxes illustrate
chromosomes as are seen with the usual microscope.

297.

Drets ME & Seuanez H (1974): Quantitation of heterogeneous human heterochromatin:

microdensitometric analysis of C- and G-bands. In: Coutinho EM & Fuchs F (eds.): Physiology and
genetics of reproduction. Part A. Plenum Press, New York, pp. 29-52.

298

Drets ME (1975): Human chromosome band mapping. The Lancet, 306:1035.

299

Drets ME & Seuanez H (1975):Cuantificacin de las bandas C del segmento heterocromtico terminal del
cromosoma Y humano normal. Rev Med Chile, 103: 307-311-

300.

Drets ME & Folle GA (1978):On line computerized band localizationin human chromosomes. Abstract.
VIth Enginering Foundation Conference on Automated Cytology. Schloss Elmau, p.121.

111

301.

Drets ME & Folle GA (1979):Computational aspects of banded chromosome scanning. Microsc Electron
Biol Cel, 6: 45-51.

302.

Drets ME & Monteverde FJ (1987): Automatedcytogenetics with modern computerized scanning

microscope photometer systems. In: Obe G & Basler A (eds.): Cytogenetics: Basic and applied aspects.
Springer Verlag, Berlin, chap 3, pp. 48-64.

303.

Drets ME, Folle GA& Monteverde FJ (1989): Quantitative detection of chromosome structures by
computerized microphotometrical scanning. In: Obe G (ed.): Chromosome aberrations: basic and applied
aspects. Springer-Verlag, Berlin, Heidel berg, New York, chap 1, pp-1-12.

304.

Drets ME,Drets GA, Queirolo PJ & Monteverde FJ (1995): Computer graphics as a tool in cytogenetic
research and education.Comput App Biosci (CABIOS First Byte), 11: 463-468.

F) Special features of the subtelomeric region

The powerful Instrument available in our laboratory allowed the generation of graphic images of the
chromosomal structure of human and Chinese hamster (Fig. 53), and the quantitative analysis of the fine structure
of the subtelomeric region (terminal segment of the chromosome) detecting the existence of differential density
patterns in this segment as well very small sister chromatid exchanges of chromosome material (t-SCEs). Also,
allowed us to analyze graphically chromosome digestedby restriction enzymes in this chromosome region which
induced holes as well as to explore quantitatively chromatid "gaps" [305-308].
We devoted considerable efforts to the study of telomere region,which were reported in a series of papers
(296-302) because sister chromatids patterns of differential density and the detection of small sister chromatid
exchanges (t-SCEs) were of great biomedical interest.
Traditionally in medicine, mental retardation and congenital malformations identified by chromosomal
aberrations were first described in Down's syndrome (Lejeune, et al1959) and in the case of fragile-chromosome X
(Sutherland 1977). However, it had observed that a large number of individuals with unexplained mental
retardation and birth defects existed so they were called "cryptic syndromes". Modern cytogenetic techniques
allowed to detect a significant number of cryptic chromosomal aberrations in the subtelomeric region in
individuals with mental retardation and birth defects, a fact totally ignored until recent years [305-315].(Fig. 54 af)

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Figure54 - a-c: Graphic images of high density patterns (black areas) of Chinese hamster chromosomes. df,:confirmation of density patternsin endoreduplicated CHO chromosomes.

Figure 55 -Different examples of the existence of minute sister chromatid exchanges (t-SCEs) detected in the
subtelomeric region of chromosome CHO (a-c) and in human chromosomes (d-f).

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Apparently, the small sister chromatid exchanges (t-SCEs) detected previously by us by means of our
photometric system, could have been an expression of the cryptic aberrations mentioned, so that the
microfotometric and graphic image analysis, provided a new microscopic view of the telomeric/sub-telomeric
region,has increased the interest and projections of these biomedical studies.
305

Drets ME, Obe G, Monteverde FJ, Folle G A, Medina II, De Galvez MG, Duarte JE, & Mechoso BH
(1992):Computerized graphic and light microscopic analyses of T-banded chromosome segments of
Chinese Hamster ovary cells and human lymphocytes. Biologisches Zentralblatt, 111:204-214

307

Drets ME, Mendizbal M, Boccardo EM & Bonomi R (1995);Further analyses of subtelomeric and
paracentric holes induced in human and Chinese hamster ovary cell chromosomes. Biologisches
Zentralblatt, 114:329-338.

308

Martnez-Lpez W, Bonomi R, Folle G A& Drets ME (1996):Microphotometric scanning of chromatid

gaps by AluI and BamHI in Chineses hamster ovary cells. Brazilian Jour Genet, 19:577-582.

309

Drets ME & Mendizbal M. (1998): Microphotometrical image analysis of the subtelomeric region of the
T-banded endoreduplicated chromosomes of. Chinese hamster ovary (CHO) cells.Genet Mol Biol 21: 219225.

310

Drets ME & Mendizbal M. (1998):The underlying structure of the subtelomeric region detected by
microphotometrical scanning and chromosome graphic imnage analysis. Mutation Res, 404:13-16.

311

Drets ME (2000) Insights into the structure of the subtelomeric chromosome segments. Genet Mol Biol, 23,
4:1087-1093.

312

Obe G, Pfeiffer P, Savage JRK, Johannes C, Goedecke W, Jeppesen P, Natarajan AT, Martnez-Lpez W,
Folle GA&Drets ME (2002):Chromosomal aberrations: formation, identification and distribution.
Mutation Res, 504:17-36.

313

Drets M E (2004): Cytological indications on the complex structure of the subtelomeric region. Cytogenet
Genome Res, Chromosomal Aberrations, 104:137-141.

314

Drets ME& Santiaque FF, (2007): A biomedical perspective of the telomeric structure and function.
In:Chromosomal alterations: methods, results and importance in human health. Obe G, Vijayalaxmi, eds.
Springer-Verlag, Chapter 14, pp. 225-240.

315

Santiaque FF & Drets, ME (2007): Non-random distribution of high density chromatin detected at
opposite ends of T-banded human metaphase chromosomes. Short Communication. Genet and Mol Biol,
30:1054-1057.

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(2). Humanand Insect Cytogenetics

A )Selective Endoreduplication
In the course of the cytogenetic studies on human chromosomes that we started in in Uruguay in 1961 we
observed that a malformed patient carried a cell mosaic with an abnormal long chromosome number 16 due to an
extra segment which was detected in three generations. This abnormal chromosome showed, in a number of cases,
the phenomenon of selective endoreduplication (also named partial trysomy), that is, a chromosome with a
duplicated segment [316]. This rare chromosomal configuration was first observed by Lejeune et al(1968) in
human chromosome No. 2. Therefore, our finding represented the second reported case in the literature of selective
endoreduplication found in a human chromosome.

Figure 56.a: First observation of selective endoreduplication of chromosome number 2 discovered by Lejeune et
al (1968);b, diagrammatic Interpretation of the abnormal chromosome 16. c, endoreduplicated chromosome 16
found in our laboratory.
316. Drets ME, Cardoso JH Delfino AH & Carrau J (1970) Familial normal partial trisomy 16 with selective
endoreduplication in malformed proband. Cytogenetics. 9: 333-350.

B )Meiotic terminal associationof the autosomes

During a cytogenetic study of chromosomes of the common cricket (Gryllus argentinus) we found that all
chromosomes carried intensely stained telomeric segments (C bands). Moreover, we found that all the

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chromosomes of this insect (except the X chromosome) were associated end to end (i.e. through both telomeres)
during meiosis.

Figure 57 a: Total telomeric association during meiotic pachytene final stages of the cricket through
heterochromatic bivalent telomere segments. b: progressive separation of the associated bivalent chromosomes
suggesting football (soccer)players. The sex X chromosome is compact and not associated during meiosis.
This finding was the first example of the cytogenetics literature of an unexpected universal chromosome
association. The uniqueness of this discovery, led us to dedicate this work to Professor Saez in honor of his 75th
anniversary [317].

317
Drets ME & Stoll M (1974) C-banding and non-homologous associations in Gryllus argentinus.
Chromosoma, 48: 367-390.

C )The fine structure of the parachute chromosome

The cytogenetic study of a small coccinellid (Epilachna paenulata) usually contaminating plantations of
common pumpkin (Cucurbita pepo) showed that had 16 autosomes and the sexual system xyp presented large
heterochromatic segments located in the centromeric region. The particularity of this insect is the behavior of the
sex bivalent Xyp (called parachute chromosome). In this case, both the heterochromatic regions of the X
chromosome and the Y enter in contact. Subsequently, the long arms of both chromosomes become folded
contacting each otherside to side, and the two fused heterochromatic segments generate an image that resembles
the parachute and long arms folded the parachutist.
Until this paper was published, the complex sexual Xyp was interpreted differently, under the supposition
that the compact X was the cap (the parachute) and that the Y the segment not compacted. Therefore, our detailed

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description which included all stages of this process, illustrated a mechanism of formation of the sexual bivalent
Xypdifferent from that previously reported in the cytogenetic literature [318] (Fig. 58 d).

Figure 58 -Stages of formation of the parachuteXypchromosome. In a-dare observed the progressive

approximation of the heterochromatic regions of the X chromosome (large) and the Y (small) forming the cap of
the sexual bivalent of E. paenulata with the long chromosome arms folded resembling the "parachutist". (d) The
meiotic separation of the two sex chromosomesis illustrated in stepse-i.

318. Drets ME, Corbella E., Panzera F, & Folle GA (1983) C-banding and non-homologous associations. II. The
"parachute" Xyp sex bivalent and the behavior of heterochromatic segments in Epilachna paenulata.
Chromosoma, 88, 249-255.

(3). Experimental Mutagenesis

Ever since the research of Fieser and Ardao (1955) we know that the Gonyleptidine secretion produced by
Acanthopachylus aracnids discovered by Estable consists of three dimethyl p-benzoquinones whose main
component is 2,3-dimethyl -2-5-ciclohedian-1-4dione.
In subsequent work, the experimentswere extended to study the action of these compounds by treating
chromosomes from human leukocytes and bone marrow mouse cells with the three components of Gonyleptidinein
a continuous way or in pulses, confirming that the these agents possess clastogenic properties. The clastogenic
action is so intense thatis capable of producing different chromosomal aberrations completely comparable to the
ones obtained by treatingAlliumonion and Lymantria dispar (grasshopper)cells.

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The aracnids secreting Gonyleptidine live in colonies of about 50 to 300 individuals. Usually their living
areas are completely free of microorganisms and other animals.In some cases, they share the same ecological niche
with another opilionidPachyloides thorelli thorelli. The formation of these associations depends, apparentlyof the
time of year, so they are probably also affected by temperature and humidity, as this species can live freely or
associated in colonies or sub-colonies or buried in the ground during winter.

Figure 59.a: TheAcanthopachylus secreting Gonyleptidine. The arrow indicates a secretion drop. b, Colony of
Acantopachylus as is found in Nature (S.R. Capocasale photos)
The animal uses the secretion as a defensive weapon as other opilionids, black beetles and termites which
secrete similar benzoquinones. The extraordinary case is of the bombardier beetles described by Eisner et al.
(1977) that are animals capable of projecting at distance their defensive secretion against their enemies.
In the case of Acanthopachylus, is evident that the continuos dispersion inthe environment of pbenzoquinoes vapors in the ecological niche where the animal lives could contribute, along the time, to an increase
of the spontaneous mutation rate of other living beings inhabiting near these areas. For this reason, we
hyphotetized that this continuos secretion could originate a chemical warlike playing an additional role in the
mutation and evolutionary pocessses occurring in Nature [319].
In certain experimental conditions, the 2,3-dimethyl p-benzoquinone can also induce an extreme
chromosome shortening resembling chromosome fragmentation. This observation would suggest of practical
importance when evaluation the mutagenic action of new synthetic substances [320].
319. Drets ME, Folle GA & Aznarez A. (1982): Clastogen Action of a Dimethyl p-Benzoquinone of Animal
Origin. Mutation Res, 102: 159-172

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320. Drets M E (1983): Extreme chromosome shortening induced in human lymphocytes by 2,3 dimethyl pbenzoquinone resembling fragmentation. Brief Communication. Environmental Mutagenesis, 5(6), 923-927.

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Triple colophon
(1) On the great hope of Sez in the scientific future of Uruguay.
RADIO INTERVIEW TO PROFESSOR SEZ
Official Radioelectric Broadcasting Service of Uruguay, (SODRE).
A Radio Interviewwas made toProfessor Francisco Alberto Sez by science journalist Mr. Fermn Rivero,
Director of the Program "Scientific Horizons", on his return to Uruguay after stayingfor one year at the
Department of Zoology, Columbia University, New York, USA, where he carried out research as Visiting
Researcher with Professor Arthur Pollister and with Torbjrn Caspersson Professor in the Institute for Cell
Research and Genetics of the Karolinska Institute in Stockholm, Sweden, between 1951-1952.
This program was broadcast by CX6 Official Radioelectric Broadcasting Service (SODRE) of Uruguay,
between the months of May/June, 1952, approximately. The original recording was made using a wax disk by Dr.
C. Mndez Bauer. Later on the recording was transcribed on magnetic tape (1976) and the text recovered by the
author (1986).
REPORT CONTENTS

Reporter:
Professor Sez was attracted since he was young by the enchantement offered by Biology becoming in this
area as an original investigator, earning awards and being designed member of European and American scientific
societies. The solid scientific reputation of Professor Saez is reflected by the fact that he was elected VicePresident of the Seventh International Congress of Genetics held in Edinburgh in 1939. He has turned his
preferential attention to the so-called atoms of biology, the chromosomes, which contain entities, in an infinitely
small world, the whole secret of life.
During his stay in the United States, he had an outstanding participation at the Annual Meeting of the
American Society of Genetics and Zoology, in which he submitted an important research paper. In that country, he
was appointed member of several scientific societies. In Europe, he gave lectures and seminars in some of the
centers he visited, admiring the fruitful work carried out there, principally in Sweden,with Professor Caspersson
of the Karolinska Institute in Stockholm, and in Italy
We remember at this time, that Professor Caspersson honored "Scientific Horizons" during his stay in
Uruguay, during the Symposium of Cellular Physiologyin 1950.
As the journey of Professor Sez produceda very fruitful work, of that noble activity of the intellect and
dignity that always distinguished our territory and that his trip was not beingof vanity, as happen withthose

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scholarships, many times favored by mediocrity because they are mediocrities, let him talk us something we,
observed and studied the ideas and techniques with great masters of Science.

Professor Sez:

I thank you very much, my friend Rivero, for your invitation and the honor of given meto collaborate in this
second stage of the"Scientific Horizons " program. During my stay in the United States and Europe, I worked on
new methods in microspectrophotometry, in order to improve our understanding of certain intranuclear
components such as the nucleic acids, thatplayan important role in the processes of inheritance and cell
reproduction.
It is notable howit has been possibleto penetrate in the infinitely small and mysterious environment constituted by
the cell nucleus by means of spectrographic analysis. By this procedure, we can be explore within cell regions, of
even of one ten thousandth of a square millimeter. These methods rely on the fact that many organic compounds
have a characteristic absorption spectrum to either visible light or infrared or ultraviolet light. In a word , they are
substances capable of absorbing light at certain wavelengths of the electromagnetic spectrum
To what is due this importance of nucleic acids ?
The significance of these studies is that we must understand deeply the behaviour of living matter such as nucleic
acid compounds. This substance is important since it integrates these tiny particles called genes governing the
heredity of organisms, cell and viruses. The cancer problem is also lluminatedopening new paths with unlimited
perspectivesthat bring us clearer to its understanding . Hence the enormous interest aroused by nucleic acids,
which form the substrate needed to keep the balance of life.
Could you tell us, Professor Saez which results you obtained from yor research?
It would be long to explain, dear friend Rivero, within the limitation imposed by this transmission, but in brief I
will say that I managed to discover by microspectrophotometric study the differences between chromatin which
carries the hereditary substance and the inert chromatin that is not acting in inheritance but playing a major role in
controlling the development and reproduction of the cell. I could thus detect that both types of chromatin differed
by having different light absorption curves. This discovery was communicated at the Annual Joint Meeting of the
Society of Biology from the United State held last year at Cornell University in the state New York. That was
attended by about two thousand researchers. In order to learn the methods of the famous Swedish School I also
worked some time in the Karolinska Institute of Cell Research in Stockholm with Professor Caspersson with
ultraviolet spectrophotometry techniques, obtaining confirmatory results of what I did in the United States.

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What impression do you bring on the development and promotion of science in those countries?
The panorama of the scientific research is admirable in the U.S. and in European countries such as England,
Belgium, Switzerland and Sweden. In the latter country the government, together with private aid, continues to
contribute large amounts of money to the progress of science. As an example I will mention the Karolinska
Institute for Cell Research, in Stockholm which I would call the "Palace of Spectrophotometry" not existing in the
world anything similar. Not only its instrumentation and facilities they have are impressivebut also the way they
have of helping an investigator. I had two electrical engineers at my disposal for the operation of their complicated
instrument,a microspectrograph, which I used to make my observations.
In which European country did you find further development in the field of biology?
Excluding Sweden, which is, as I said a moment ago, one of the most advanced countries, I have the impression
that Italy is, after the world war, one of more vigorously recovering countries. Germany also, of course, but the
resources have not reached yet to the level which requiredby scientific work. In this sense, the National Research
Councils, dedicated to promote the development of scientific research creating scholarships and grants to
laboratorieswhere a serious and continuing work is doneconstitute a powerful fact becauseof the multiple benefits
and incentives for research they produce. England, France, Belgium and some other countries, including
Brazil,have these organizations that have clear concepts that the progress of a nation that depends on its scientific
research institutions.
Do you plan to continue this work with us?
Since my arrival, it was my immediate concern to restart work at the Department of Cytogenetics in my position of
the Institute for Biological Research directed, as you know, by Professor Clemente Estable. The Rockefeller
Foundation has donated 60,000 dollars to our laboratory for instrumental acquisition, which will broaden horizons
of work since the methods used in modern biology have become more complex and expensive, requiring wellequipped laboratories and libraries.
What are your current plans?
To be brief, I will mention one. Some time ago, Professor Estable discovered that the secretion of a gland of a
small arachnid called Gonyleptes had antibiotic properties. Interested by this surprising fact, I thought what would
be the effect of this substance on cell reproduction. Soon I obtained evidences that it was capable of producing
aberrations and curious disturbances that could alter the cell division mechanism as well as the hereditary process

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that suffers by the action of this extraordinary substance which has been named by Estable Gonileptidine. In these
experiments we have the collaboration, with great efficacy of my research assistants Claudio Solari and
MaximoDrets, being full ofyoung plethoric enthusiasm and vocation for original investigations.
I am going to pose one last question regarding the scientific work in ourcountry: what is your opinion?
I have strong faith and great hope in the scientific future of Uruguay but,before we can cross the circle imposed on
us by our position with respect to the Northern Hemisphere, and to stand on the same level in the universal concert
of Science, it is absolutely necessary to send young men, already trained, to the United States and Europe, to enter
in contact with the great researchers and penetrate the modern techniques. Thus, I have the certainty that, in the
short term, we will have seriously prepared researchers, holders of rigorous scientific and profound theoretical
basis methods, who in a continuing effort, will bring the best of themselves in the pursuit of those wonderful
mysteries that perpetually cause the avid curiosity of the real researcher.
You have listened toProfessor Francisco Alberto Sez, Head, Department of Cytogenetics of the
Research Institute of Biological Sciences, Montevideo, directed by Professor Clemente Estable.

2 The Teacher's eternal message to youth

The following is a fragment of the conference given by Prof. Sez in the Faculty of Phyisical, Natural an
Exact Sciences of the University of Buenos Aires in October, 1935, with the auspices of the Student Center of
Natural Sciences Doctorate [145].
Young people listening me, very soon you will be a naturalist, a new man devoted to scientific work
thatwill be enriched to the event of our own efforts in our intellectual environment.
I Imagine you, plethoric of enthusiasm, anointed by the sacred flame of a irresistible scientific vocation
irresistible yearning to unleash the dream that you hide in your thoughtful forehead.
The nature of our soil is awaiting you with endless secrets, remembering that the most trascedentalto
pulling back the veils of nature is the method of attack.
The time has come to renew old molds that preceded us. Their work methods, although worthy of respect,
should be transformed in full, updating them to the prevailing thought.
We live in a new era that astonishes by the boldness of its techniques and you, who are the future, will be
responsible for furthering them. Only by working with high-minded goals you will develop our science and our
country."

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(3) The deep insight of two great cytogeneticists:

A) The gene, invisible wonder of Biology.
Francisco A. Saez (1956)
"The gene has a formidable potential. When it exerts its action carries within itself the creative power.
Reproduction always originates an identical gene, so that each of them is multiplied each time the cell is
reproduced. This extraordinary property is its most transcendent feature. The gene may well be considered then,
as the physical basis of life; as a unit of life. If the cell is a colony of genes, we can imagine that the first signs of
life were dominated by simple naked genes, free. Most notable is that these particles, whose size is below one
millionth of a millimeter and that gathered together those oftoday ishumanity might not reach the volume of an
aspirin tablet, hold in the deepest mystery the of theirnature intimate constitution. But we should be carefulbefore
assuming that with the gene we resolved the last structure of life. Science has discovered a solar system in the
atom. This teaches us to be circumspect in such problems thus who knows if the tiny gene does not hide an even
infinitely smallerworld "[194].

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B)Leon Inaugurale.
Professeur Jrme Lejeune
Chaire de Gntique Fondamentale.
Facult de Mdecine de Paris.(10 Mars 1965)
Des insaississables prformations dEmpdocle aux esprits animauz de Descartes; des molcules organiques de
Bufon aux squences de bases des acides nucliques, une hypothseinluctable reparat, lancinante.
Quel que soit le nom quon lui donne, quelles que soient les qualits quon lui prte, ou la structure quon lui donne,
ou la structure quon lui suppose, quelque chose doit exister aux confins de la matire et de la forme, quiles transcende
toutes deux et pourtant les unit.
Virus formative, moule intrieur, ou codage des macromolcules, ne sont que les tranformations progressives dune
ide toujours la mme; ide qui nous parat invitable, peut-tre parce quelle est rellement inscrite au profound de nous
mmes; ide que je ne saurais mieux rsumer quen paraphrasant le dbut dn trs vieux livre.
Au commencement il y a un message
Ce message est dans la vie
Et ce message est la vie

Figure 60. -Prof. Jrme Lejeune talking with Prof Sez on problems
of the human cytogenetic karyotypes at the Faculty of Physics,

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Natural and Exact Sciences.Buenos Aires, 1965.

AnUnforgettableEpilogue:
For Maximo Drets:
It has well been said that the purest glory of a teacher is to forge wise men that exceed him. It is my
greatest hope that you continue, dear Maximo the path that leads to the full achievement of your ideals.
All achievements of science were born of a longing, a sublime aspiration often branded as chimerical, that
persistent effort converted into an amazing reality.
Sure that it shall be this way, I foresee the most fruitful and luminous path in your research career.
With all my love, Francisco A. SezJanuary 1970-

Figure 61.-Cover and back of the volume published by UNESCO in honor to Dr. Francisco A. Sez on his 70th
anniversary with dedication (224).

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Beak ML & Beak W (1988): Evolution by polyploidy in Amphibia: new insights. Cytogenet Cell Gene. 80; 2833
Beak ML & Beak W& Rabello-Gay MN (1965):Cytological evidence of constant tetraploidy in bisexual frog
Odontophrynus americanus. Chromosoma, 19; 188-193.
Benavente R, & Folle G. (Eds) (2010):Chromosomes, Genomes and Beyond. Cytogenetics and Genome Res;128,
(1-3): 1188.
Caspersson T (1950):A Universal Ultramicrospectrograph for the Optical Range. Exp Cell Res; 595-598.
Caspersson T., Jacobsson F., Lomakka G., Svensson, G. Sfstrm R.(1953):A high resolution ultramicrospectrophotometer for large-scale biological work Exp Cell Res.; 560563.
Caspersson T, Farber S, Foley GE, Kudynowski J, Modest EJ, Simonsson E, Wagh U, Zech L.(1968):Chemical
differentiation along metaphase chromosomes. Exp Cell Res. 49(1): 219222. [
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fluorescent agents. Chromosoma, 30: 215-227.
Chial H (2008):Cytogenetic methods in diagnosing genetic disorders. Nature Education 1(1); 34
Darlington CD (1932): Recent advances in cytology. 1a ed., J. & A: ChurchillLtd., London, 559 pp. (& folle G2a.
ed P.Blakistons son & Co., Inc.,Philadelphia, 671 pp., 1937).
De Winiwarter H, Oguma K (1926): Nouvelles recherches sur la spermatognse. humaine Arch Biol 36: 99-166.
Eisner T, Jones TH, Anehnsley DJ, Tschinkel WR, Silberglied RE & Meinwald J (1977):Chemistry of defensive
secretions of bombardier beetles (Brachinini, Metriini, Ozaenini, Paussini).J Insect Physiol, 23: 1383-1386.
Estable C, Ardao MI, Pradines-Brasil N, Fieser LF (1955):Gonyleptidine. J Am Chem Soc, 77(18):4942.
Gall JG & Pardue ML (1969):Formation and detection of RNA-DNA hybrid molecules in cytological preparations.
PNAS 63: (2) 378-383.
Galgano M (1933a): Studi intorno al comportamento della cromatina nella spermatogenesi di Rana sculenta L. I.
Il nmero e la forma dei cromosomi nel processo normale. Arch Ital Anat Embriol, 31: 1-86.
Galgano M (1933b): Evoluzione degli spermatociti di I ordine e cromosomi pseudosessuali in alcune specien di
anfibi. Arch Ital Anat Embriol, 32(1): 171-200.

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Garber E (1944): Spontaneous Alterations of chromosome morphology in Nothoscordum fragrans. Amer J Bot
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Innes WT (1954): Exotic equarium fishes. Sex changes in fishes.Innes Pub. Co.,Philadelphia, pp. 496-497.
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129

THREE GREAT CYTOGENETICISTS OF ALL TIMES

FRANCISCO ALBERTO SAEZ

Photo:1968
CLARENCE E. MCCLUNG URUGUAY (S.A.)
Photo: 1930
(U.S.A.)

CYRIL DEANDARLINGTON
Photo: 1920
ENGLAND

130

NOTICES AND ACKNOWLEDGEMENTS

This book is an English version of the Spanish book published in 2013 by DIRAC, the Editorial Office of the
Faculty of Sciences of the University of the Republic, Uruguay. The present edition has carefully re-written
including more historical information about the Institute of Biological Sciences Clemente Estable, adding
additional figures and new footnotes. Special emphasis was given to the scientific close relationhip existing with
the Brazilian geneticists.
The present English version of the book on Prof. Francisco A. Sez (Life and Research Work) has been donated by
the Author to the Brazilian Society of Genetics (Sociedade Brasileira de Gentica) so its distribution is free of
charge to anybody.
The chapter on Sez family was partially based upon autobiographical documents given by Prof. Sez to the
author, as well through information, and documents about the family structurekindly provided by his daughter
Prof. Graciela Saez, Director of the Historical Institute of Morn, Buenos Aires, which allowed the construction of
the "Sez Familygenealogy ".
The bibliography of Sez publications was partly collected by Blanca Modernell Ferreira, (2nd Ed., IIBCE, 1979)
and considerably expanded by the author.
The story "The actor on stage" was partly based on the obituary written by Professor Juan H. Hunziker,
(Argentina) Fig. 27; Footnote 23 (288).
The three images of metaphases illustrating the cover were taken from the paper of Sez FA, Rojas P & De
Robertis E [31].
Other recent contributions of members of the Department of Cytogenetics appear in the book Cytogenetic and
Genome Research. Benavente & Folle (Eds.,2010). (seeAdditional References, p.121).
I express my sincere thanks to Professors Francisco Salzano (Brazil), Eddy De Robertis (USA), and Nstor O.
Bianchi (Argentina) for their interesting anecdotes on Prof. Saez.Besides, Professor Nstor O. Bianchi (Guidance
for young researchers) and (Cyrcadian Rhythm and Sez Forgetfulness), and Professor Eddy. De Robertis. (How I
learned to respect the wisdom) kindly translated into English their own stories about Prof. Sez, a valuable
collaboration greatly appreciated.
I wish to express my deepest gratitude to my son, Comp. Eng. Gustavo A. Drets, who patiently helped me to
overcome many problems faced with the computer and the word processing system. Without his competent and
kind cooperation this book would have been impossible.
I am alsodeeply indebted to Professor Eddy M. de Robertis for his generous and detailed correction of the English
translation of this version.
I thank Mr. Hctor Valverde for his valuable advice about the process of sex reversal observable during breeding
aquarium fish species.

131

Finally, the author thanks Mr. Gabriel Santoro, Head of the Editorial DIRAC of the Faculty of Sciences of
Uruguay,for hiseditorial workof the book,originally published in Spanish,thatwas additionally enriched by a
number of footnotes on severalsubjects of the text.

132

SUBJECT

PAGE

Memorial
2
Prologue, to the master
3
The origins biographical shades
4
Genealogy of the family
9
IIBCE scientific diversity
13
His great friend: Estable
19
About vocation
22
Saez & LA friends
23
The eminent teacher
28
Generating prestige
30
The geneticist translator
31
A compromised scientist
31
Anecdotes & facets of Saez
33
Recruiting first Uruguayan disciples
33
The inspired researcher
35
The absent-minded professor
35
Remembering Sez
36
How Ilearned to respect the wisdom
37
Guidance for young researchers
39
Cyrcadian rhythm & Saez forgetfulness 40
The actor on stage
42
Scientific contributions
44
Cytological methods for chromosomes 44
Orthoptera
45
Amphibians
50
Polyploidy in amphibians
53
Plants
55
Mammals
58
Native animals
59
Arachnids
60
Experimental studies
60
Citotaxonomy
61
Sex chromatin
62
Experimental mutagenesis
62
Cytospectrophotometry
63
Honors and Awards
69
Disciples and coworkers76
Scientific publications
77
Book Translations
84
Reviews of scientific publications
85
Articles on Prof. Sez
90
Papers from other authors reviewed
91
Obituary
96
Addenda Scientific legacy
97
(1)Microphotometry
97
B Software for microphotometry
99
CChromosome banding
101
The story of C-banding discovery
102
DMicrophotometry& banding
105
F Features of subtelomeric regions 107
A Selective endoreduplication
109
2)H uman & Insect cytogenetics
110
BAutosomes end to-end associations
110
C Parachute chromosomes
110
3) Experimental mutagenesis
112
Radio Interview
114

FIGURE NoPAGE

FOOTNOTE No.
1- Inst. Chemitry.
6
2 Cajal
6
3 Ostwald
6
4 Pi Suer
7
5 Lehmann Nitsche 8
6- Torres
8
7 - Estable
9
8- Cordero
11
9 - Miller
14
10 Pern
21
11 Houssay
21
12 Leloir
21
13 McClung
22
14 Belloni
25
15 Mc Clintoc
28
16 Loedel
29
17- Sabato
29
18 Lanuza
30
19 Muller
32
20 - Larraaga
34
21 - Morgan
38
22 - Ginza
41
23 Hunziker
43
24 Darlington
46
25 - Pollister
64
26 Caspersson
66
27 Stern
75
28 Flemming
104
-

The deep insight

118

1 - Genealogy Saez Family 9

2 Papico Boy
10
3 - Saez 1921
10
4 - Saez 1928
12
5 IIBCE Former Laboratories13
6 Cordero, Sosa, Saez , Herter
13
7 Harry Miller, Saez, Estable
15
8 Gallery Brazilian Geneticists
16
9 IIBCE Builidings
17
10 Cell Sorter&Electronic Microscope 17
11 IIBCE personnel
18
12 Indoors garden
19
13 Saez & Estable
20
14 Saez tiendo en vasos Coplin
20
15 - Saez & McClung
23
16 Cordeiro & colleagues
24
17 Saez in congress 24
18 Saez, Freire Maia, MED
25
19 Saez , Brncic & Personnel
26
20 - Saez & Brncic, Beak, Cantu 27
21 Saez &Mc Clintoc
27
22 Postgrade Course F. Medicine28
23 Cover Book Cytology
30
24 Hans Muller 32
25 Show subte
33
26 4 authors
42
27 Hunziker
43
28 Camera Lucida
44
29 Darlington
46
30 Heterocromatinization
47
31 Batovi Hill
49
32 Saez & MED observing Drosophila
50
33 Meiosis Bufo
51
34 Poliploidy in Amphibia
53
35 Sorghum almun
57
36 .- Mulita & Carpincho
59
37 Caspersson Photo64
38 Caspersson & Pollister Cytophotometers 65
39 Saez, Lison, Schreiber
67
40 Award Lucio Cherny
73
41 Saez & Stern
75
42 Saez et al. retuming from Estanzuela 76
43 Saez with a large optical microscope84
44 Saez Anniversary
95
45 Last photo
96
46 Image Discriminator
98
47 Nuclei diagram 99
48 Bandscan
100
49 Zeiss MP01 system 101
50 Banding pioneers
102
51 C-banding map
104
52 G bands
105
53 Graphic scanned chromosomes
106
54 Graphic density patterns108
55 t-Sister chromatid exchanges
108
56 - Endoreduplication
110
57 Non homologous association
111
58Parachute chromosomes
112

Leon Inaugurale
Epilogue
Additional References

119
120
121

59- Gonyleptes
60 Lejeune & Saez
61 UNESCO book celebrating Sez

The teachers message

117

113
119
120

PAGE