Elio Secondo Raviola, M.D. Ph.D.

Elio Raviola, a scientist and professor emeritus in the Department of Neurobiology at Harvard Medical School passed away on December 23, 2023, at the age of 91. Having received his early training in medicine, neurology and psychiatry in Italy, he immigrated to the US in 1970 where he would become an acclaimed teacher in the medical community, among the world's foremost neuroanatomists and a pioneer in elucidating the ultrastructure of the retina of the eye.

Raviola engaged in research on the nervous system for 66 years, directing a laboratory in the Departments of Anatomy and Neurobiology at Harvard Medical School from 1970 to 2023. From 1972 to 2002 he directed the introductory anatomy course for entering, first-year Harvard medical students, also lecturing extensively in histology and neurobiology to medical and graduate students. He was renowned for his animated, erudite lectures on human anatomy. His excellence as a teacher was documented on several television documentaries about the journey of medical students into the field of medicine. In 1994 he was elected a member of the American Academy of Arts and Sciences.

"His groundbreaking research taught the world how the retina develops and general principles of the form underlying function in the nervous system. His work solved longstanding questions about the role of individual neuronal cell types in the retina, in the mouse, and in primates. Thanks to his decades of intense work, we can assign molecules and genes to specific cell types in the retina, and from my own work I know that genetic access to cell types has revolutionized our ability to understand the building blocks of the nervous system with unprecedented resolution. For more than 60 years, he also taught, provided guidance and mentorship to thousands of Harvard students, postdocs, staff, and faculty. The number of lives that he touched and careers that he impacted may be the largest in the history of Harvard Medical School. I can think of no one who has accomplished so much across so many dimensions of the institutional mission of the school-research, discovery, and education. Many of our faculty have had impact in one of these areas, yet he was singular in his ability to have immense and direct influence across them all," says David Ginty, Chair of the Department of Neurobiology at HMS.

Early childhood in 1930's Asti (Piedmont, Italy)
Elio Secondo Raviola was born on June 15, 1932, in Asti, Italy, where his father managed what would later become the Saclá canning factory. His first language was the Piedmontese dialect, the second was Italian, and later the third was English, which he never formally studied. The surname Raviola (descendant of an old dialect word for "metal worker") and the name Secondo are distinctive of the Piedmont region. He had vivid memories of his childhood, the villages of his ancestors in the countryside (his maternal grandparents came from Celle Enemondo), and bike rides with his father to gather fruit together in the hills outside the town. Asti is the northern capital of Monferrato, known for its rolling hills of grass, poplar woods, hazelnuts, oaks, elms, fruit trees, and vineyards. While today the region is famous for wines (Barolo and Barbaresco, as well as everyday classics like Barbera, Grignolino, Freisa, Nebbiolo Langhe, and Dolcetto) and prized white truffles, historically, it was a landscape of remarkable biodiversity and a unique cultural character which was rooted in his soul. His childhood instilled in him from a young age a deep respect for the beauty of the natural world. The underlying political chaos of Italy between the two wars also informed his appreciation for knowledge, social order, and human understanding. This influence extended to his future evolution as a scientist and intellectual.

In his father, Raviola saw a man of great intelligence, creativity, and skill in building beautiful things with both hands and mind. An expert in managing factories, his father Giuseppe (known as "Pinin," 1901-1980) was also politically active. Both his father and his uncle Carlo, a member of the Arditi Battalion in World War I, became members of the new Italian Communist Party upon its foundation in 1921, later becoming part of the anti-fascist and anti-Nazi resistance movement of the 1930s in Italy. Raviola grew up during Benito Mussolini's dictatorship. This environment instilled in him a sense of workers' rights and values of social justice, which also aligned with the Catholicism of his mother, Luigina Carbone (known as "Luigia," 1903-2000). While his father supervised a clandestine, later partisan, communist brigade from 1922 to 1945, Elio was often tasked as a lookout at the entrance gate during his childhood. Elio attended the Arnaldo Mussolini elementary school on Corso Dante, as well as middle and high school in Asti.

Raviola's joy of discovery, interest in experimentation, physical dexterity, artistic inclination, historical and political awareness, and appreciation for literature, music, and excellent cuisine were cultivated during his childhood. He attributed to his mother "modesty, sincerity, generosity, respect for others, humble intelligence, good taste, richness of feelings, and a spirit of noble sacrifice," as well as her protectiveness, for laying the foundations of his moral education and character. The strictness of his mother also supported his meticulousness as a student from a young age. He saw his maternal grandmother as a woman of remarkable intelligence and honesty, who had formally attended school for a total of six months at the age of 7 before being sent by her father to Turin to learn to be a servant for a wealthy family.

World War II, and High School
During the 1936-39 Spanish Civil war the household was a safe house for Italian fighters returning from Spain (many by foot). The experience of later hiding people at risk in the house from the Nazis during World War II added to the family's sense of insecurity. At the age of 11, during the 1943 Italian Civil War, the country's most tenuous wartime moment, Raviola asked his father's permission to become a partisan fighter, to which his father told him that he was too young. He held vivid memories of German soldiers stationed in front of the house and of American planes bombing the city. Later, American G.I.s moving north to take German-held territory made cigarette smoking fashionable, a habit Elio enjoyed for more than 45 years. The hardship he experienced during the war reinforced his deep disdain for fascism and Nazism, and extremism in general. While not identifying specifically as a communist, late into his life this experience informed his identification as always being "A Man of the Left."

After the war, his parents moved to the alpine town of Morbegno in Valtellina during his high school years, where his father managed a different factory. Elio would take the train to visit his parents at the mountain home during Christmas, Easter, and summer, while during the school year, he stayed with his maternal grandmother in Asti. His parents and grandmother maintained an apartment in Asti, and he stayed there, continuing with high school. About his grandmother, Elio noted, "My grandmother was phenomenal. It's wonderful to be treated as an adult by someone who takes care of you. She had complete trust in me. She knew I knew what I had to do. And I was doing what I had to do." He often recalled the experience of newly felt freedom during his years of high school. This included time with friends and learning to travel, informing a cosmopolitan aspect. His first trip out of Italy was to Allied-occupied Vienna in 1948 at the age of 16 with a high school friend. Soviet soldiers manned the checkpoints as they entered the city. "We went to the Saltzburg Festival. I had never seen a classical opera. Beethoven's Fidelio…To be free, to choose to go where we wanted, to choose a restaurant, to choose anything…To me, it was an incredibly maturing experience." The conductor was Wilhem Furtwängler, chief conductor of the Third Reich. An excellent student, Raviola graduated from Natale Palli High School ("Liceo Scientifico di Asti") in 1951.

As a teenager, Raviola developed a passion for mountain climbing, scaling Punta Rasica, Piz Badile, Ortles, and various peaks in the Valmasino Alps in Italy, without the knowledge of his parents. He noted, "I was happy to be so independent. And in those years, I loved mountain climbing, even with my parents being so against it. In Valtellina, there's a chain of mountains of such pure granite that when you climb, you see the quartz crystals. There's nothing like climbing beautiful granite mountains." At the time hikes and climbs were dangerous, undertaken with only a wool sweater, thick rope, and no other equipment. With friends he would drive to Bagni di Masino and then hike for three hours to reach the huts. Over the years, they became familiar with the hut keepers. His passion for climbing was discovered by his parents when he took a photo of his friend climbing and brought it to a photography studio in the alpine town of Morbegno, which published it in the local newspaper. A worker in the factory showed the photo to his father.

Raviola remarked in 2020, "I liked Asti. I was so attached to the city. Despite the war, my youth was fantastic. My parents were so good to me." He remembered close friends from his childhood in Asti: Stefano Gagliardi, since the very early elementary years; Bartolomeo "Mimmo" Bianchi (1933-2016), a successful agriculturalist; Giovanni Bo (1934-2023), a renowned architect; and Mario Trinchero (1934-2009), a philosopher and professor at the University of Turin. As an only child, his closest living relative in Asti was his cousin, Piera Carbone (1937-2016), with whom he shared a strong bond.

At the age of 90, Raviola described his beginnings: "My life has been an endless, endless fight to improve from the conditions in which I was born. Sometimes I think about what extraordinary people my parents were…The infinite goodness of my mother. The inquisitive spirit of my father. And they were not even aware of it. Both went through only 5th grade. All that they had and overcame, day-to-day, came through incredible effort. And I was born in this little, provincial city that I love, where I grew up. I remember so often going out on the balcony on Corso Torino and seeing the beautiful hills [of Piedmont]. And then, everything went so well for me. I was good in school. I wanted to go to Ghislieri. I went to Pavia. I started an academic career. I came to this country. I found an environment that was my environment…But I missed Asti. I can't do anything about it. Whether it's missing Asti or missing my youth, what I miss are now just glimpses."

University of Pavia and Collegio Ghislieri
While Raviola was expected to work in the factory, his father endorsed his matriculation at age 19 to the University of Pavia (founded in 1361). While at the University of Pavia he was elected to the Collegio Ghislieri in the Foundation Cesare Artom, based on his entrance examination scores. Founded by Pope Pius V in 1567, the Collegio Ghislieri is a prestigious institution linked to the University. Collegio Ghisleri provides housing, scholarships and cultural opportunities and the third-largest private library in Northern Italy. He graduated from the University of Pavia (M.D.) with perfect grades and honors in the School of Medicine and Surgery in 1957 at the age of 25, completing a thesis titled, "Histological and histochemical contribution to the study of the hypothalamic-pituitary neurosecretory system," which was awarded the Veratti prize for the best dissertation for a graduate of the 1956-57 class. For a young man of modest means, the scholarships were essential, and the opportunity to study in Pavia in the fertile political landscape of post-war Italy was transformative.

While at Ghislieri in 1953, as a third-year medical student, Raviola met Giuseppina d'Elia (1935- 1986), a first-year student at Collegio Castiglioni-Brugnatelli, just across the street from Ghislieri. Elio described her as "intelligent, cultured, honest, sincere, passionate, beautiful, frugal, and an indefatigable worker with an enormous sense of duty." Both eventually worked on similar research topics, and Giuseppina became a member of the same Institute of Anatomy at the University. Elio and Giuseppina Raviola were eventually married in Pavia in 1960. Raviola established lasting friendships across a wide range of academic disciplines, including historians, philosophers, and theoretical physicists. He noted that "winning a spot at Ghislieri was wonderful. Those were very interesting years. Very brilliant companions. But I also became part of the University of Pavia and the medical faculty as a student." Some of these friends and colleagues included: Franco Tatò (1932-2022), a renowned industrial executive at Olivetti, Mondadori, Fininvest, and Enel; Fulvio Ghiringhelli, a physician in Pavia; Giovanni Fiori from Brescia, a physician in Milan; and Giorgio Greggia. The impact of these interactions on his further intellectual development cannot be understated. His years at Collegio Ghislieri solidified a deep interdisciplinary curiosity, a cosmopolitan spirit, and a sense of being part of a long historical tradition and heritage of knowledge in northern Italy. For many who knew Elio, he had a vast knowledge base that was part of his engaging character, making him a great conversationalist in any context, with anyone. These qualities, as well as his loyalty to friends, were nurtured at Ghislieri.

Graduate Study and Early Scientific Career
With his medical degree Raviola spent several years engaged in training as a specialist in neurology and psychiatry, managing a unit at the local mental hospital at a time when there were no available psychopharmacologic interventions. Pavia had been the academic home of Camillo Golgi, the pioneer of cellular neuroanatomy who in the late 19th century had discovered a special method of staining, inspiring generations of classical histologists and providing the cellular basis for the rise of modern neuroscience. Raviola learned the tricks of Golgi's 'black reaction' for staining neurons directly from Antonio Pensa (1874 1970), one of his surviving students" (Nature, Vol 444, 16 Nov 2006). He developed a passion for research and the nervous system. He completed his Ph.D. in 1963 at the age of 31 and was an Assistant Professor in the Institute of Human Anatomy from 1958 to 1971.

More specifically, as a graduate student Elio was named "Extraordinary Assistant" in the Institute of Normal Human Anatomy within the Anatomical Institute at the University in 1958, "Deputy Assistant" in 1959, "Assistant In Charge" in 1960, and, at the age of 29, Deputy Assistant in 1961. In September, 1963, at the age of 31, Elio became free Lecturer in Normal Human Anatomy with ministerial decree. Taken together, these stages comprised his progression toward the Italian equivalent of a Ph.D. In conferring this honor the selection board expressed itself thusly: "The scientific production of the candidate is truly commendable…In the practical tests the candidate has demonstrated promptness and profound knowledge. The lessons deserve particular praise for the singular qualities of clarity and expository order and for the properties of language. The Commission unanimously considers the candidate laudably deserving of becoming free lecturer in normal human anatomy."

"Elio Raviola was one of the greatest Ghislerians and graduates of the University of Pavia of all time. His studies have deeply penetrated the functions and structure of the retina, changing the vision we had of this fundamental sense organ and contributed to new treatments for its dysfunctions. He is great man and great scientist who did so much for the development of contemporary neuroscience," says Paolo Mazzarello, Professor of the History of Medicine and Director of the Museum of Natural History at the University of Pavia.

Pavia to Boston, 1965-70
In 1960, at the age of 28, bitten by the thrill of scientific discovery, Raviola found himself frustrated by the pace of his academic progress, and felt that he needed mentorship. Raviola was a free spirit. He had scientific ideas that he wanted to explore. He knew how to work hard, and he felt constrained. He decided to take a chance. He took a train to meet with Professor Rodolfo Amprino (1912-2007) at the University of Bari. Amprino was a student of the famous Italian anatomist Giuseppe Levi (1872-1965). The schools of Golgi and Levi had been intellectually in disagreement with one another at the time, as Levi was an acolyte of the Spanish neuroscientist, pathologist and histologist Santiago Ramòn y Cajal (1852-1934). Golgi and Cajal had received the 1906 Nobel Prize for their investigations on the microscopic structure of the brain. Golgi had believed that everything in the nervous system was a continuous network of communication, while Cajal used the staining techniques developed by Golgi to prove of the existence of synapses: that the nervous system was instead comprised of discrete cells that could communicate. The vestiges of this intellectual rift remained in Italian neuroscience at the time. So, when Elio asked Amprino if he would take him in his department, Amprino suggested that such a change would destroy Raviola's career in Italy. This related to the potential act of switching from one school of thought to another. Amprino told him: "Don't do it." Amprino instead suggested that Elio go to the U.S. for several years for a new experience, and then later transfer to his lab in Bari. Amprino said that he would help him. In 1963 Amprino provided Elio with the address and the name of an Italian scientist working in the US, Rita-Levi Montalcini (1909-2012), who was just starting to publish her findings. Montalcini, whose future discoveries on embryonic nerve cells and nerve growth factors leading to the 1986 Nobel Prize were at the time unknown, was in the Department of Zoology at Washington University in Saint Louis. Her twin sister was living in Rome, and Elio traveled the next year from Solda in the Alps to meet with Rita-Levi during a summer visit from the U.S.

Raviola hoped to spend time in the labs of either George Palade (1912-2008) at the Rockefeller University, or Don Fawcett (1917-2009) at Harvard University, two of the three originators of the field of modern cell biology, along with Keith Porter (1912-1997). While Raviola was an anatomist, Fawcett was a prolific cell biologist from whom Elio felt he could learn new techniques. Through a communication from Montalcini to Fawcett, Fawcett declined the request for Raviola to work in his laboratory because of an academic conflict between Fawcett and Montalcini. Raviola then did something relatively inconceivable at the time. He took a pen, and a very thin sheet of airline paper (like tracing paper, used commonly at the time), writing Fawcett a letter stating a case that he should take Raviola into his laboratory because he hoped to do research with him in electron microscopy, Fawcett's area of technical expertise. This was unusual because such arrangements were normally made through formal channels, by department chairs communicating with department chairs. Fawcett was surprised yet moved. Fawcett invited both Elio and Giuseppina to spend one year as Research and Teaching Fellow, respectively, in Anatomy at Harvard Medical School, from 1965-66. On arrival to Boston by boat, Elio and Giuseppina, now also an anatomist and neurobiologist, were given an empty lab space by Fawcett, who told them to make a list of reagents and other equipment they needed to demonstrate their scientific aptitude in short order. The trial period went well. However, while he was offered a position by Fawcett, Elio and Giuseppina returned to Italy due to her developing an illness, requiring treatment. She recovered in Italy and they were pursued by Fawcett to return to Boston. In 1970 Raviola accepted Fawcett's invitation as Associate Professor of Anatomy. Giuseppina was invited as an Associate in Ophthalmology at Harvard, and in 1972 became Associate Professor of Anatomy at Boston University School of Medicine (and Professor in 1979). The couple led the introductory medical school courses in Anatomy at Harvard and Boston University, respectively, while also developing successful, separate, research careers focused on the retina.

Of his move to Harvard in 1970 Elio has said that "I was so lucky to meet somebody like Don Fawcett, who independently of nationality-of anything [just my skill as a researcher and teacher]-he liked me and encouraged me." Elio's niece by marriage, Anna Milano, recalls that "Among my childhood memories there are Giuseppina and Elio; it was a true celebration when they came to visit us in Vercelli, and I remember well when we went to see them in Pavia on their departure to the United States, ready for a new life. They were my young, beautiful, intelligent aunt and uncle, who I looked upon with admiration, and who I always saw in this way with the passing of the years." Elio remembered the day, October 23, 1970, arriving at Logan Airport and being greeted at Logan Airport by various professors in the department, including Susumu Ito (1919-2015), Jean-Paul Revel (1932-2021), Betty Hay (1927-2007) and others.

Raviola's sense of indebtedness to Fawcett was immeasurable, as was his respect for him as a person, which he summarized in an obituary for Fawcett, with Sus Ito, in 2009: "Don Fawcett's intellect, imagination and vast understanding of zoology and comparative physiology had a unique impact on students, postdoctoral fellows and faculty alike. For him, looking at living things was an adventure that never ceased fascinating him even in the last years of his life. Tolerant of human frailness, endowed with an excellent sense of humor, full of understatement about his gifts and accomplishments, he detested pomposity in all its manifestations" (Journal of Anatomy, 2010;216:1-2).

Scientist at Harvard Medical School, 1970-2023
Raviola became Associate Professor of Anatomy in 1970, Professor of Anatomy in 1974, Bullard Professor of Neuroanatomy and Professor of Ophthalmology in 1989, Bullard Professor of Neurobiology in 1993, and Professor Emeritus in 2013. He dedicated himself to research on the nervous system for sixty-six years. He led a laboratory in the Departments of Anatomy and Neurobiology at Harvard Medical School for fifty-three years, from 1970 to 2023. Elio received numerous offers for leadership roles in other departments early in his Harvard career. These included offers for chairs at Cal Tech, the position of Dean at Washington University in Saint Louis and the University of Pennsylvania, and an invitation from Johns Hopkins as well. He stayed at Harvard because, simply, he didn't want to be a Chair, and was happy in the department at Harvard.

Raviola was an expert of cell biological, electrophysiological, and molecular techniques, solving problems of structure, connectivity and physiology of the retina, the part of the eye that received and processes signals from light. He was interested in how the retina is able to encode information from the visual scene, and then send the encoded information to the brain. To study this complex problem, he employed multiple methods, and applied them to studies in primates, rodents, and rabbits. As he was originally trained in classical anatomical methods, he was able to use these methods in an elegant and rigorous fashion. However, he was not content to use these methods alone, and thus became an expert in the electrophysiology of the retina, as well as some of the newest molecular approaches. Over six decades he contributed to evolving, foundational knowledge in the field of neurobiology, while continuously learning new methods to effectively respond to new scientific questions.

With Ramon Dacheux in the 1980's he conducted novel studies of the way in which the photoreceptors, the cells that receive and process light, interact with secondary neurons in the initial processing of visual information. They studied the functional role of different classes of nerve cells, in part through microscopic analyses of how the cells connect with each other, and in part by recording their electrical responses to light. Throughout the years they made recordings twice weekly of electrical transmissions in rabbit retinal cells. The experiments lasted from early in the morning through most of the following night. After days of wakefulness Dacheux would sleep at Raviola's house. Raviola later wrote: "Very few things in life are as rewarding as sharing the joys and frustrations of scientific discovery with a trusted colleague of great intellectual integrity and the same aspiration for excellence. When the work becomes a silent, graceful, well-choreographed ballet of precise, perfect gestures; when there is little or no need to speak and the collaborators complete each other's sentences because they have become one with their object of study" (Visual Neuroscience, 2007;24:445-447).

With Torsten Wiesel, 1981 Nobel laureate in physiology or medicine, he developed an experimental model of myopia (near-sightedness). They studied how alterations of the visual experience during the postnatal growth of the eye leads to myopia. They discovered that the eye elongation that leads to myopia is mediated by the nervous system, specifically by growth regulating chemicals produced in the retina itself.

On his scientific work Raviola said in 2020 that "I started working on the retina when nothing was known…Throughout my career our knowledge of the retina reached a point where we now know so much. I worked on rods, cones, horizonal cells, rod bipolars and amacrine cells."

Raviola had most recently directed his efforts to characterizing the role of each neuron cell type, focusing on the ways in which amacrine cells in the retina uniquely release the neurotransmitters dopamine and GABA, as well as seeking to understand the specific function of this mechanism in human adaptation to light. Using a multidisciplinary approach he identified all the transcripts present in dopamine amacrine cells, discovered the presence of the common clock-related proteins in those cells (Gustincich et al.), showed that some amacrine cells spontaneously release dopamine and GABA through different mechanisms (Puopolo et al., and Hirasawa et al.), and he described an unique set of synaptic contacts made by dopamine amacrine cells at nodal points of the retinal network needed to optimally shape retinal light adaptation (Contini et al., Masland et al.)

Constance L. Cepko, Bullard Professor of Genetics and Neuroscience at Harvard Medical School remembers when Raviola approached her at the turn of the 1990's about her supporting his learning newer methods in her laboratory. While he was a visiting professor, he behaved as a visiting post-doctoral student: "He came to my lab to learn molecular biology. He kept the most beautiful, neat, and complete lab notebook that I had ever seen. I used it as an example to my students. and he was so courageous, at his august stage in life, to become a novice in a new field. He was driven to this by his desire to understand more about retinal circuitry. He felt he had exhausted the approaches that he had been using, and wanted to try something new, methods that he had come to understand would provide powerful new avenues to achieve his goals. He was able to learn enough to return to his lab and use the new methods to make a transgenic mouse that labelled the dopaminergic neurons that he wanted to understand. It led to a beautiful study of the synapses of that cell type, made possible by his mastery of anatomy and physiology, combined with molecular biology."

Raviola loved and respected his friends and colleagues. He spoke of their scientific successes with the same pride as he did his own. He enjoyed the scientific successes of mentors and colleagues like Weisel, Fawcett, Stephen Kuffler (1913-1980), Ito, David Hubel (1926-2013), Baruj Benacerraf (1920-2011), and others from the very beginning. He would say, "Don was an extraordinary man. Torsten is an extraordinary man. Susumu Ito was an incredible scientist. Like Don and I, Sus made many discoveries. You could make a major discovery in an afternoon, and it's a major discovery. He made many discoveries. In 1965, Sus and Tom Pollard, a second-year medical student at Harvard, discovered how cells move. The biochemical mechanism for muscle contraction had already been discovered. Why an amoeba could walk. Why a cell could translate (the movement of mRNA in a cell's cytoplasm to produce proteins). No one knew anything. Actin and myosin. The basis for cellular motility. Mucosal permeability and how it repairs. But I liked Sus because he was an extraordinary man."

Teaching of Anatomy, 1970-2002
In addition to practicing his science, Raviola was a revered teacher of anatomy for more than thirty years. One of the conditions of his hiring in 1970 by Fawcett was that Raviola would teach Anatomy. He was responsible for the Introduction to Anatomy course for entering, first-year Harvard medical students, also known as "The Body Block." He directed the course for thirty years, from 1972 through 2002, also lecturing extensively in Histology and Neurobiology to both medical and graduate students. His excellence as a teacher was recognized instantly by students and faculty alike. In 1972 Raviola received the Boylston Society Award for excellence in teaching at Harvard Medical School, as well as multiple subsequent pre-clinical teaching awards. He was called by some students "The Italian Master," naturally weaving his cultural and intellectual foundations into his teaching and encouraging personal authenticity from the students themselves. In the dissection room he not only taught the students methods of dissection and anatomical detail, but also reviewed the history of anatomical discovery over the past several thousands of years. He conveyed the importance of humanity and humility, and the sacred calling, for first-year medical students familiarizing themselves with the physical presence of human bodies that had been donated to the medical school. With a remarkable memory, he could quote Cicero, Julius Caesar and Dante verbatim from his high school lessons. This was all tempered with a disarming, warm and kind spirit.

He brought to his teaching a brilliant intellect and quick wit, an encyclopedic knowledge of the history of science and medicine, and a deep understanding and appreciation of human nature and behavior. Raviola believed that teaching required two most essential ingredients: love and generosity. To be a great teacher one also had to be a good scientist, because one's choices and interpretations must be backed by rigorous understanding of the experimental evidence. Great teachers must also know their subject cold. In 1987 Harvard revamped its medical student curriculum to deal with the rapidly expanding scientific knowledge of the prior decades. The curriculum, called the New Pathway, had compressed the duration of courses and the approach for students to learn and integrate the incredible amount of material. This required that faculty members select what knowledge is most relevant for future physicians to learn from their time in the classroom and the labs. It also required skillfully discussing clinical cases with the students in tutorials. Raviola felt that teaching anatomy and histology well required additional unique skills: being a good lecturer, dissector, microscopist, and a wise moderator in student tutorials. Given the volume of information available, this learning process could be particularly stressful for students. Successful teaching therefore also required of professors the humility to recognize the depths of their own ignorance, and a deep respect for the students themselves. Knowledge, commitment to the material, understatement, sense of humor and self-deprecation were the values he cared for most.

Raviola was known in particular for his remarkable lectures, which were a dramatic and artistic performance, cherished by students for decades. He would come to the lecture hall early in the morning and draw detailed, beautiful pictures in color on the black board, using French art chalks. He saw lecturing as a performance art the purpose of which is to transmit just the right amount of information that the class could absorb in one hour, to make the material interesting, to convey a sense of intellectual or practical necessity, and hint at the existence of a world of detail, moving from surface structure to molecular and genetic depth, a whole world that a student could learn through a lifetime of professional practice. Raviola valued teaching anatomy as the occupation of an artist of consummate skill, persuasiveness, and both detail and simplification at the same time. Between 1986 and 2002 Raviola co-led the human anatomy course at Harvard Medical School with Daniel Goodenough. His excellence as a teacher was documented in the television documentary series Chronicle and Nova. Trudy Van Houten, former Director of the Clinical Anatomy Course and Co-director of the Human Body Course at Harvard Medical School, notes that "His lectures were legendary: beautifully organized, original and ingenuous, wonderfully humorous, and full of surprises. They were carefully crafted lessons that included exactly the information students needed, combined with the brilliance and charm necessary to keep students captivated from the first to the last sentence and in memory even decades later. They were, first and foremost, lessons directed at the students who listened to them. His lectures remain unforgettable...I also remember his incredible tact when I was a new anatomy instructor in his anatomy lab at HMS and how generously, and diplomatically, he shared both his knowledge of anatomy and his knowledge of effective anatomical teaching. I also recall, with admiration, how he managed the extraordinary feat of dividing his time, relatively evenly, among eight dissection tables all clamoring insistently for his attention."

"Elio was an amazing scientist and extraordinary person. His aesthetic sensibilities influenced his beautiful anatomical studies, as well as his students and admirers," says Carla Shatz, Professor of Neurobiology at Stanford University and a pioneer in early brain development.

"Elio was a scholar and intellectual of the old school. He had a sense of his discipline's history and tried to convey the same in his lectures, demonstration, and tutorials. His colleagues appreciated these attributes as well," notes James Adelstein, Executive Dean for Academic Programs at Harvard Medical School from 1978-97.

Jim Yong Kim, a co-founder of Partners In Health, former President of Dartmouth College and the World Bank, and one of TIME magazine's "100 Most Influential People in the World"-and a former student of Raviola's-notes that Raviola "was a great man and had such a huge impact on so many. He also just gave us the sense that as students from diverse backgrounds coming into Harvard we were welcome, and we didn't all have to be cut from the same cloth."

Mentorship of Young Scientists, and Service to Harvard and Science in Italy
Raviola had never forgotten how he had been welcomed to Harvard in 1965 by a number of senior colleagues, including the man who would become his best friend, Susumu Ito. A Japanese-American, Ito had been a hero from the famed 442nd Army Infantry Division of World War II, the most decorated unit in U.S. military history, while his parents and sisters had been moved from California to Oklahoma and placed in a federal internment camp. Only learning later about this personal history, it was Ito's grace, kindness and welcoming way that left an indelible impression on the new Harvard faculty member from Italy. Over the decades Raviola returned this spirit in kind. Michelino Puopolo, Associate Professor at Stony Brook University notes that "when I first arrived in Boston in 1998, Elio was like a second father to me." Raviola was a dedicated mentor to many young scientists as well as a trusted advisor to leaders at Harvard Medical School. While serving on medical school committees he also advised departmental, hospital and Harvard Faculty of Arts and Sciences faculty, graduate students and postdoctoral fellows, including on microscopic techniques. Richard Born, Professor of Neurobiology and a former director of the HMS Ph.D. Program in Neuroscience recalls "Elio was such a wonderful man, colleague and scientist. When I first started my lab, we did a project to try to reconstruct electrode tracks in living animals. One of the figures for the resulting paper required a high quality photo-micrograph. I went to Elio for help. He immediately dropped everything and walked me and my student, along with our histology slide, to his microscope room, where he not only took the photo, but gave us a complete lesson on light microscopy and the best color films to use. I think he was happier about our result than we were! Years later, when Kathy Rockland and I teamed up to start a project that involved electron microscopy, she and I sat in his office for about two hours one afternoon while he gave us the entire history of the field, along with detailed technical comments on fixatives and embedding media. He had incredible stories from his youth that he shared with us. He also taught me gross anatomy when I was a first-year medical student. He took a mind-numbingly difficult subject and made it come alive, with clever demonstrations and great humor."

Matthew Lawrence at the St. Kitts Biomedical Research Foundation notes that "He was a source of wise and valued counsel at every important professional step from pursing graduate training, then medical school and ophthalmology, and subsequent commitments to translational science and institution building. Indeed, in the latter, his spirit will long live onward in St. Kitts and our growing team in the U.S. His passion for inquiry and joy in life always elevated the human connections essential to practice, purpose and outcome in biomedical research. My work aspires to the commitments to collaboration, mentorship, and basic science and patient impact that he embodied. I have been privileged to receive his guiding light and long friendship."

Raviola was very active in efforts to strengthen Italian science, including the creation of the Italian Institute of Technology and the establishment of the Giovanni Armenise-Harvard Foundation. In advising the Armenise-Harvard Foundation, which supports basic scientific research at Harvard Medical School and in Italy, he worked in a focused way to help the careers of young, promising Italian scientists. He mentored Italian post-doctoral fellows in his laboratory, including Massimo Contini (University of Florence), Stefano Gustincich (Italian Institute of Technology Genova), Adalberto Merighi (University of Turin), Michelino Puopolo (Stony Brook University), and Enrica Strettoi (Institute of Neuroscience at the Italian National Research Council in Pisa). He cherished engaging with Italian collaborators with independent careers, including Giovanni Berlucchi (University of Verona), Emilio Bizzi (Massachusetts Institute of Technology), Saverio Cinti (University of Ancona), Cesare Montecucco (University of Padua), Enrico Mugnaini (Northwestern University), and Paolo Pinelli (University of Pavia, Catholic University in Rome, University of Milan).

Enrica Strettoi, Director of Research at the Institute of Neuroscience at the Italian National Research Council in Pisa notes that "Elio was my undisputed scientific mentor, the one who believed in me when I was little more than a little girl, who convinced me that I could do it and win a place in the research academy. His enthusiasm, passion for microscopy, intuition of the importance of what you see but which not everyone is able to decipher, they infected me many years ago. I hope I have passed on a bit of all this also to my students, to whom I often turn using his own words…He was so generous, both in a human capacity and scientifically, and was for me my first home in professional terms, the place where I would mentally go when I feel in difficulty or had fear that I was not up to something scientifically that seemed too complex. I was incredibly fortunate to cross paths with him and equally lucky to maintain his respect and esteem. I went to him in very difficult moments as well as in those moments of great professional recognition: his consistency has always been admirable and striking, as he knew how to be himself on every occasion, with wit, clarity, and passion."

He received honorary degrees from the University of Ancona in 1996 and the University of Turin in 2002. In 2002, he was also awarded the Ottorino Rossi Award from the University of Pavia, presented to a scientist who has made an important contribution to research in the field of neurosciences.

Stefano Gustincich, Principal Investigator at the Italian Institute of Technology in Genoa, recalls 30 years of memories when "Elio was a father-like figure nurturing my growth as both a scientist and a human being."

Massimo Contini, Professor at the Università Degli Studi di Firenze notes that "I loved him. It was certainly one of the fundamental meetings of my life. I can't summarize the meaning of this friendship in a letter, we were lucky to have had him in our lives. The memory of his intelligence, his irony and his energy will remain forever."

Saverio Cinti, Professor at the University of Ancona, notes that "I always had the ambition to consider myself his student and his younger brother. I will always remember him with great affection and I will greatly miss his passionate stories of the Pavia period. His words have always been an example and my entire academic life has been influenced by him, especially the great passion for scientific research so expertly pursued and appreciated about him throughout the world."

Cesare Montecucco, Emeritus Professor at the University of Padua notes that "Elio put truly exceptional care and energy into advancing his students...He was a great man. We were different ages, but we came from the same world, and we immediately understood each other's way of seeing life and science."

Raviola served the Departments of Anatomy and later Neurobiology at Harvard under the leadership of five Chairs: Don Fawcett, Betty Hay, Carla Shatz, Michael Greenberg, and David Ginty. He worked in the Departments of Anatomy and Neurobiology from October 1970 to April 2023. Raviola loved being a part of both departments, and he loved and respected his friends and colleagues. He talked about their scientific achievements with as much pride as he did his own. He was deeply engaged in the intellectual life of the Department of Neurobiology and found the department to be "extraordinary." Of the strengths of the department he recently wrote: "A varied approach to neuroscience, total freedom to express oneself, a friendly atmosphere of openness, mutual respect and generosity that leads to rapid circulation of ideas and collaborations, and a Chair concerned about the welfare of the faculty." He saw the mission of the department to be one of "advancing knowledge of the brain by combining rigorous molecular, cellular and behavioral techniques." He greatly appreciated both formally and informally convening with his departmental colleagues in an environment that encouraged the sharing of ideas on a wide range of topics related to science, culture and the personal. During COVID-19, he continued going into the department five days per week for most of the pandemic, when physical return to the medical school was permitted.

Raviola saw his mentorship of students (both medical and research) and colleagues in the department as central to his usefulness as he advanced in age. Into his 80s, he served on the HMS Subcommittee of Professors, the Prizes and Awards Subcommittee, the Honors Committee, the Armenise-Harvard Foundation Junior Faculty Grant Review Committee, the Armenise-Harvard Foundation Scientific Advisory Board, the Armenise-Harvard Foundation Italian Scholarship Advisory Committee on Career Development Awards, the Excellence in Mentoring Award Selection Committee, the Council of Mentors Subcommittee, and he chaired meetings of the Graduate Student Advisory Committees.

Personal Life
Raviola often quoted an old Roman adage: "Est modus in rebus: Everything in proportion. Moderation in all things." He was an avid pianist, working with two teachers over the course of 40 years. Tatiana Yampolsky, a graduate of the Moscow Conservatory where she studied with prominent Soviet pianists Yakov Flier and Dmitry Bashkirov, noted that Raviola "was incredibly devoted to music. Always so passionate and excited during our lessons, we talked endlessly about the topic. Most importantly, I will miss him as a friend, very much." Raviola enjoyed travel, particularly to Italy. He had his favorite places in nature, restaurants, churches and museums where he would enjoy aesthetics and history. This included a tour he developed over decades of lesser-known Caravaggio paintings in churches across Rome, that he would share with friends. He was an enthusiastic reader, intellectually engaged on many topics, including classical music, ancient civilizations, history of art, the U.S. Civil War, and evolving coaching strategies in professional soccer. Raviola was known by most people who knew as being a irreverent and playful, and for emphasizing the importance of having fun. Years after his move to the US, in the 1990s, he became a U.S. citizen.

Raviola noted the difficulty of feeling neither Italian nor American, "a person without a country." Fundamentally identifying himself as a scientist, Raviola had a clear vision for the evolution of science in the U.S. and globally; he connected the evolving scientific landscape to the political environment. He was concerned that "the best young scientists are no longer coming" to the U.S., a research environment which he felt had offered him the freedom and support to manifest his own creative capacities and gifts to the ultimate degree.

Raviola's political education from the childhood experience of being raised in Fascist Italy informed his perspective on current events. He had faith in what he saw as the best aspects of American society: freedom of expression and support of human potential, no matter one's origins. He believed in the importance of American science and industry to the world. He saw the re-emergence of global Fascism over the past decade as an urgent threat to humanity, in part because it fed racism and hate towards the vulnerable: "When the racism reaches a peak, you get Nazis, who appeal to the worst traits of mankind…We're traversing a bad moment for mankind. I really hope that we won't have war. It will be devastating. I don't know whether people realize that a war will mean the end of mankind. It would transform the Earth into a desert...Now it is tolerated to be a racist. [These attitudes] will completely destroy science in this country. Maybe racism is more powerful than we assume. Immigrants are still the great strength of this country."

Americans have developed a greater interest in things Italian over the past several decades. The tradition of Italian intellectualism may be an area of less familiarity. Italian as well as American societies have undergone remarkable social transformation over the decades since World War II. Raviola's life serves as a bridge between these two evolving and shifting cultures. For him, throughout his life in America, what was most important were the opportunities he was given to express himself professionally. To the end, he was grateful for the uniqueness and the gift of his Piedmontese and Italian upbringing, his education in Pavia, the entrée he received to Harvard, and for the open canvas provided by the American scientific landscape of the time.

Raviola gave all of himself to his close friends and family as well. His first wife Giuseppina died in 1986 at the age of 51. His second marriage from 1989 to 2009, to Trude L. Kleinschmidt, ended in divorce. He was an engaged, loving father. Raviola's strong memories of his relationship with his father informed his own generous parenting approach, and the encouragement he gave his son to work hard, take calculated risks, and seek an interesting, productive life. He is survived by his son Giuseppe Raviola, a leader in psychiatry and global mental health at Harvard Medical School, Massachusetts General Hospital and Partners In Health, his daughter-in-law Emily Raviola, and two grandchildren.

He was laid to rest at the Newton Cemetery & Arboretum, Massachusetts (U.S.) on January 2, 2024. A memorial event will be held at the American Academy of Arts and Sciences on April 21, 2024.

Gifts and donations in his honor can be made either to the Department of Neurobiology at Harvard Medical School (specify that the funds are to go to "Trainee initiatives") or to a fund to support the international health care delivery organization Partners In Health.