Editors' Note: Alan Light, University of Utah, Salt Lake City, US, remembers the life and work of a pioneering pain researcher. We invite you to share your memories of Ed Perl by posting a comment below.
Dr. Edward R. Perl, nociceptor pioneer and champion of the specificity theory of pain, and a founding member and first president of the Society for Neuroscience, passed away July 15, 2014.
Perl, with Paul Richards Burgess and Paul Bessou, documented in the late 1960s the existence of unique A-delta and C-fiber “nociceptors” by using single unit recordings. “Our routine of testing by gentle mechanical stimulation yielded no activity. After some minutes of fruitless trial, one of us, and we cannot remember whom, picked up a tissue forceps and pinched the skin in the middle of the nerve's receptive field. This evoked a burst of impulses. We looked at each other across the experimental table, both recognizing what we may have seen, and then systematically explored the unit's responsiveness,” said Perl in The History of Neuroscience in Autobiography (Volume 3, Ed. Larry R. Squire). So began the modern era of discovery, characterization, and, ultimately, understanding of the cellular and molecular nature of nociceptors. Perl, along with Burgess and Bessou, documented the important characteristics of A and C nociceptors, including their differential responses to noxious temperatures, and the property of “sensitization,” in which the nociceptors become much more responsive following damage to the tissues they innervate. During this time, Perl coined the name “polymodal nociceptor” to describe C nociceptors that responded to noxious mechanical, thermal, and chemical stimuli.
In the early 1970s, Perl, with Burgess Christensen, went on to show that these nociceptors activated specific neurons in the marginal zone and substantia gelatinosa of the spinal cord in the cat and, as shown later with Takao Kumazawa, in the monkey as well. Later in the 1980s, with Christopher Honda and Siegfried Mense, Perl showed that these neurons activated neurons in specific regions of the thalamus.
During the later 1970s and 1980s, Perl, Alan Light, Dan Trevino, and Miklós Réthelyi, as well as Yasuo Sugiura and Chong Lam Lee, established the anatomical substrates for these pathways by using intracellular labeling of physiologically defined A and C nociceptor primary afferent neurons and similarly identified neurons in the superficial dorsal horn. To determine if these “nociceptors” were, in fact, responsible for the sensation of pain, Perl, with Frithjof Konietzny, Trevino, Light, and Herbert Hensel, further showed that activation of specific nociceptors led to the sensation of pain in humans. These experiments clearly demonstrated the existence of a “labeled line” for pain from nociceptors in the periphery to the thalamus in the brain.
In the early 1990s, Perl, with Jun Sato, Daniel Bossut, and Virginia Shea, showed how damage to the sympathetic-adrenergic system could enhance nociceptor signaling and thus how sympathetically related pain states could be established.
More recently, Perl, with Timothy Grudt, Adam Hantman, Yan Lu, and Jihong Zheng, turned his attention to how the substantia gelatinosa is organized to receive and modulate input from peripheral A- and C-fibers.
Perl received the Bristol-Myers Squibb Award for Distinguished Research on Pain in 1991 and the Ralph W. Gerard Prize in Neuroscience in 1998. He was elected to the American Academy of Arts and Sciences in 1992. In 2000, Perl endowed the University of North Carolina (UNC)-Perl Prize to be given to investigators who have made significant contributions to neuroscience through outstanding or seminal insights. Four of the winners of this prize have gone on to win Nobel Prizes.
Early life and training
Edward Roy Perl was born in 1926 in Chicago, Illinois, US. His parents, John and Blanche Perl, were natives of Hungary and Czechoslovakia, respectively. Perl attended college at the University of Chicago where he enjoyed physics and engineering but was convinced by his physician father to go into the field of medicine. He began a medical training program in the United States Navy Officer Training program as a medical trainee in 1945 at the University of Illinois School of Medicine, graduating with a medical degree in four years, in 1949, at the age of 22. It was during this time that he developed his interest in neuroscience, and met and worked with many of the top scientists in the country in this area of research.
Dr. Perl’s first Science paper, published the same year he obtained his medical degree, was not on pain systems, but on a device to measure cardiac volume in humans. This project became the foundation for today’s impedance cardiography, an important tool in cardiovascular diagnostics. However, he had already developed an interest in neuroscience through his experience in Warren McCulloch’s lab, in which he also coincidently met Walter Pitts, Jerome Lettvin, and Patrick Wall, one of the originators of the gate theory of pain some 17 years later.
“McCulloch and his coterie of associates, Pitts, Lettvin, and Wall, spent most of their time in the unit's library discussing and considering various theoretical approaches to the functional organization of the nervous system. That was heady stuff. I did not understand much of it. However, the gravity of their postulations and the complexity of the situations they were considering were impressive,” wrote Perl in The History of Neuroscience in Autobiography.
In 1948, Perl served as a clerk at Harvard Medical School, Boston, US, where he met Derek Denny-Brown, who influenced him to pursue a career in neurophysiology.
After obtaining his medical degree, Perl did a fellowship with Philip Bard and Vernon Mountcastle at Johns Hopkins, Baltimore, US, where Perl became a neuroscientist and interested in C-afferent fibers, a preoccupation that would encompass the rest of his scientific career.
Perl entered active duty as a Navy physician from 1952-1954, becoming part of a research group that included David Rioch, Robert Galambos, Michael Fuortes, Walle Nauta, and David Whitlock.
A career in neuroscience
In 1954 Perl took his first faculty position at the State University of New York, College of Medicine in Syracuse (now SUNY Upstate Medical University), US. He left for the University of Utah, Salt Lake City, US, in 1957 because of the efforts of Carlton Hunt, who had formed one of the strongest neuroscience-focused departments of physiology in the US. At the University of Utah, Perl collaborated with David Whitlock and published three very influential papers on the spinothalamic pathways and the properties of thalamic neurons that responded to activation of those pathways.
In 1962, Perl began a lifelong connection with European neuroscientists with a year in Toulouse, France (he went to France speaking no French). During this time, his interactions with Paul Bessou, Denise Albe-Fessard, Pierre Buse, Jean-Marie Besson, and Hungarians Janos Szetagothai and Miklós Réthelyi not only added stature to his research efforts, but also aided his research efforts in technical ways, and imbued a lively atmosphere to his laboratory in Utah and North Carolina, since return visits by these dignitaries were frequent for years to come.
In 1963, upon returning to Utah, Perl began his research on C-fibers in earnest, armed with some of the tools necessary to study them. He modified these tools and added to them to create methods that allowed him to study large numbers of physiologically characterized neurons with Paul Richards Burgess and Paul Bessou, and in doing so revolutionized our concept of what nociceptors were and how they behaved in both cats and monkeys.
In 1969, asked by Ralph Gerard to do so, Perl wrote the initial bylaws for a scientific organization that later was called “The Society for Neuroscience.” Perl was elected president of this organization by the founding group of 20, but he chose to take the title of “acting president” because he felt that what was needed was a scientific society, devoted to neuroscience, that was as democratic as possible—one with a leader elected by a representative membership. In 1970, the Society elected Vernon Mountcastle as the first elected Society for Neuroscience president, and the first meeting was held in 1971. Today, the Society has nearly 40,000 members in more than 90 countries and 130 chapters worldwide.
In 1971, Perl moved to Chapel Hill, North Carolina, US, to become chair of the department of physiology at the University of North Carolina School of Medicine for the next 18 years.
In 1989, he resigned as chair following a conversation with a friend. “I asked my mentor and friend, Vernon Mountcastle, over a beer at a meeting in Stockholm on a sunny Swedish day what he thought about resigning the position as chair. He looked at me without a smile and said, ‘It would be the happiest day of your life,’ ” Perl wrote in The History of Neuroscience in Autobiography.
Throughout his outstanding career of over 43 years at the University of North Carolina, Perl continued to solve difficult problems with sometimes controversial results.
Outside of the lab, he enjoyed fine wine, fly fishing in remote areas, sailing along North Carolina’s outer banks, and his wife and family. He has taught many of us a most important lesson, which is how his autobiography ends: “One does not know what tomorrow will bring, but today I still look forward to the pleasure that comes from a successful experiment or that of an evening's sail, the excitement of seeing a trout or salmon rise to a fly, or of the smile of a grandchild. I close this on the way to ask Marjorie [Perl’s wife] for a dinner rendezvous.”
Alan R. Light, PhD, did his postdoctoral work under Perl at the University of North Carolina School of Medicine, Chapel Hill, US; became a faculty member in Perl’s department in 1978; and collaborated with Perl at UNC until 2003. Light is now a professor in the departments of anesthesiology, and neurobiology and anatomy at the University of Utah, Salt Lake City, US.
Further reading:
Myelinated afferent fibres responding specifically to noxious stimulation of the skin.
Burgess PR, Perl ER.
J Physiol. 1967 Jun;190(3):541-62.
Response of cutaneous sensory units with unmyelinated fibers to noxious stimuli.
Bessou P, Perl ER.
J Neurophysiol. 1969 Nov; 32(6):1025-43.
Spinal neurons specifically excited by noxious or thermal stimuli: marginal zone of the dorsal horn.
Christensen BN, Perl ER
J Neurophysiol. 1970 Mar; 33(2):293-307.
Kumazawa T, Perl ER.
J Comp Neurol. 1978 Feb 1;177(3):417-34.
Light AR, Perl ER.
J Comp Neurol. 1979 Jul 15;186(2):133-50.
Light AR, Trevino DL, Perl ER.
J Comp Neurol. 1979 Jul 15;186(2):151-71.
Perl, Edward. "Edward R. Perl." The History of Neuroscience in Autobiography, Volume 3. Ed. Larry R. Squire. San Diego: Academic Press, 2001. pp. 366-413.
Comments
Kenneth Casey, University of Michigan
I am saddened to learn of Ed
I am saddened to learn of Ed Perl's death. Alan Light has composed an excellent memorial tribute. I have only 2 comments to add:
1. To gain a full appreciation of Ed Perl's deep thinking about pain in his later years, one should read his historical summary:
Perl ER. Ideas about pain, a historical view. Nat Rev Neurosci. 2007 Jan. 8(1):71-80.
The last paragraph of this piece shows that Ed was continuing to consider all the evidence in formulating his own ideas about pain; a thoughtful man throughout his life and career.
2. Ed was a formidable runner...and probably overall athlete. He refers (quite modestly) to this in his autobiography. Ed and I ran together on several occasions when we were at meetings together in Ann Arbor or Chapel Hill. Although nearly 10 years my senior, he set a challenging pace. He set the pace in science, too.
Geoffrey Bove, University of New England College of Osteopathic Medicine
The news of Dr. Edward Perl’s
The news of Dr. Edward Perl’s death struck me like the loss of an immediate family member. I did my PhD studies with Dr. Alan Light, directly underneath Dr. Perl’s laboratory, and had the honor of regularly interacting with Dr. Perl. He sat patiently with me one long day as I fumbled with forceps and oscilloscopes, trying to record from those obscure nociceptors by using methods that he pioneered. He was nurturing, saying things like “Let me show you how to isolate that a little better,” or “How can you tell if the electrical and mechanical responses are from the same neuron?” I went home excited to brag about the day to my spouse and anyone else who would listen, sensing that I had just had a somewhat formative experience.
Dr. Perl would often say what was on his mind as well, and with that level of intelligence, the humor can be quite interesting. This might sound harsh to some, but I recall him saying that my first paper was written in “North American Gibberish.” I don’t know why that struck me as funny—maybe because we had already submitted the paper. But more likely, it was because I knew that he had actually read it, and that any feedback (and I did get serious feedback as well) was coming from a man who had an amazing gift for writing, something that he has left on record for us all to enjoy and learn from.
I think that many people misunderstood Ed. He came off as gruff sometimes, but I don't think that he had a malicious cell in his body. After I left the University of North Carolina, he remained available to me. Colleagues sometimes remarked to me about how approachable this scientific icon was. I know that Dr. Perl helped form whatever scientific skills I have, in both tangible and intangible ways. I know that his work has affected all of us in the pain field, and has helped humanity.
Allan Basbaum, University of California
For those of us who have been
For those of us who have been engaged in studies of primary afferents and the spinal cord circuits that transmit signals that generate pain, Ed Perl’s passing is particularly sad. Alan Light’s beautiful piece illustrates the incredible contribution that Ed’s research brought to the pain field, highlighted by his discovery and characterization of the polymodal nociceptor and the lamina I “pain-responsive” neurons.
Here, I would like to remind readers, particularly those who are relatively new to pain research, of one of the more profound and long-standing disagreements (one might even call it a battle) that Ed had with another giant in the pain field, Pat Wall. Ed was a lifelong believer in the specificity of primary afferent fibers and their relationship to labeled lines that transmit “pain” messages. Pat Wall, in contrast, went to his grave railing against specificity and arguing instead for the pattern theory, which postulated that activity across populations of physiologically heterogeneous afferents determines the perception that is eventually generated.
In his autobiography (Squire, 2001), Ed recollected about a 1961 letter that he wrote to Pat Wall about a draft of a Melzack and Wall manuscript that championed pattern theory (Melzack and Wall, 1962). In the letter to Pat, Ed wrote that “their manuscript contained interesting ideas, but that he should be prepared for criticism.” That comment was certainly an accurate harbinger of the heated debate that followed. As a very young postdoc, I experienced firsthand the intensity of the criticism to which Ed referred. I had just completed postdoctoral work with Pat Wall and was visiting the University of North Carolina, where I met Ed. Not surprisingly, we got into a discussion of specificity versus patterning, at which point Ed commented that Pat Wall was brilliant, but, unfortunately, Pat never did a good experiment in his life. Somewhat taken aback and, of course, wanting to defend Pat, I responded to Ed by offering, “How about excitability testing?” I was referring to a powerful and remarkably simple method that Pat developed to track changes in the terminal properties of afferent fibers, at which point Ed replied, "Okay, I’ll give him that one." It was all in good spirit but definitely illustrated the competitiveness that defined their relationship.
They never did resolve their differences. Not surprisingly, recent data indicate that both were right, depending on the part of the pain circuit being studied (see Woolf and Ma, 2007). It is also worth mentioning that Ed and Pat were invited to be on a retrospective IASP workshop several years ago, where there were only good memories and obviously, clear mutual admiration: all gemütlichkeit.
What is important, of course, is that the controversy over specificity versus patterning was one of the most significant forces that drove discoveries in the pain field. From the molecular characterization of nociceptors to the organization of dorsal horn circuitry, and even to the development of imaging methods that highlight variations in the brain patterns evoked by different modalities of pain, Ed’s discoveries will always be remembered.
Reference:
Edward R. Perl. 2001 In: The History of Neuroscience in Autobiography. Ed. Squire, L. R., Acad. Press, San Diego, Vol. 3: 366-413.
Lorne Mendell, Stony Brook University
Ed Perl was a major figure in
Ed Perl was a major figure in the development of the modern science of pain. Most neurobiologists today are not aware of the controversy surrounding the concept of the nociceptor in the 1960s. Although Sherrington had defined nociceptors half a century earlier as sensory fibers responding to damage of the tissue they innervated, their existence as a separate class of sensory fiber had not been established. Some investigators had identified candidate fibers, but they were found only infrequently.
These difficulties were attributed to the belief, based on studies using local anesthesia and electrical stimulation, that nociceptors conduct via small-diameter fibers that would be difficult to record from individually. In the 1960s, Ed and his colleagues succeeded in doing this routinely for both small myelinated A-fibers and unmyelinated C-fibers that formed two separate classes of nociceptive afferents. At the time, these were revolutionary findings with an enormous impact on the field that has persisted until the present day. Ed’s research group also demonstrated the unique projection of nociceptors in the dorsal horn of the spinal cord. This work, which was supplemented by Ed and by many others following up on his seminal findings, demonstrated that the nociceptor is not simply a subset of mechanoreceptors or thermoreceptors with unusually high thresholds, but rather is a separate class of fiber with unique physiological, anatomical, and chemical properties. This knowledge has been crucial for the development of strategies to alleviate pain.
Ed continued productive research up to the end of his life. Most recently, he published important findings on properties of the superficial dorsal horn, which, as demonstrated so convincingly in his earlier work, is where nociceptors terminate. His experiments always addressed important issues in a very straightforward manner, and his superb experimental and technical skill resulted in findings that were reliable and thus extremely important for the field.
Others have written about the disagreements between Ed and Pat Wall concerning specificity versus pattern theory for somatosensation in general and pain in particular. I had been Wall’s graduate student in the 1960s and had worked on spinal mechanisms related to the gate theory. When Ed moved to Chapel Hill to assume the chair of physiology, I was a young assistant professor at Duke, a few miles away. At that time in the early 1970s, we were not particularly close, perhaps in part because of my past association with Pat Wall. Thus, I was particularly thrilled when Ed approached me at a meeting in Paris in the late 1970s to ask whether I would consider joining an NIH Program Project that he was organizing at the University of North Carolina on the effects of peripheral and central injury on the properties of the nociceptive system. Ed was very supportive when I elected to move to Stony Brook in the middle of the initial five-year grant period and encouraged me to remain in the program project grant (PPG). This PPG ran for about 25 years and provided important resources for my work. Chief among these was the opportunity to interact with Ed at a personal level. Although he could be quite challenging in a scientific discussion, which was helpful in focusing my thinking, in fact he was a very approachable and supportive friend. He was always helpful as a sounding board for my ideas, as well as contributing many of his own excellent suggestions. Overall, he was a wonderful colleague. Our research meetings usually ended with a visit to his home for wine and cheese. He and his wife Marge were gracious hosts, and their efforts enhanced our group interactions.
Ed’s passing is a loss for the entire field. He is held in great esteem for his many crucial contributions to pain neuroscience that we all use in considering pain mechanisms. His support was very helpful to me, and I will miss his friendship and his wise counsel.
Sally Lawson, University of Bristol
Ed Perl was at the
Ed Perl was at the International Union of Physiological Sciences (IUPS) meeting in Budapest in 1980 when I first met him. He was a tall, impressive figure whose work I knew well and whom I was very keen to meet. I told him I was about to start intracellular recording in dorsal root ganglion (DRG) neurons. He advised against it, warning me of the difficulties. However, when I explained that I had funding for this study, he immediately offered useful advice. Over the years we met and talked at several meetings, and this led to collaboration.
This is how it happened. He had invited me to be on the advisory board of a doctoral student, which led to regular visits to Chapel Hill, US. On one of these visits, I gave a talk about our DRG work relating electrophysiology and conduction velocity of intracellularly dye-injected neurons to their immunocytochemistry for calcitonin gene-related peptide (CGRP) (with P. McCarthy), using whole DRGs with their peripheral nerves and dorsal roots acutely in vitro. Back in his office after my talk, Ed challenged my certainty about properties of neurons that were immuno-negative for CGRP. His concerns were based on his own dye-injection studies. He had been recording intracellularly fromthese neurons in young guinea pigs every Friday for many years, with the help of Bennie Crepps. Despite intracellular dye injections and immunocytochemistry on some of these neurons, any immunocytochemical patterns in relation to sensory properties had remained elusive, so Ed knew the problems. I explained the records we made of exact locations of dye injections, plus records of locations of sections of dye-injected neurons (camera lucida drawings, photographs) before immunocytochemistry. These enabled us to find the neurons after (non-fluorescent) immunocytochemistry despite loss of fluorescence of the injected dye (see Lawson et al., 1997), and to reject neurons that did not match these records exactly. He accepted my explanation and suggested that we collaborate, bringing together his knowledge and experience of classifying the sensory properties ofneurons and recording those properties in young guinea pigs in vivo and my experience of intracellular recording in these neurons plus the problems of dye-injection followed by immunocytochemistry. Our aims were the same: to relate sensory-receptive properties of DRG neurons to their immunoreactivity for likely nociceptor-related molecules, initially the peptides substance P and CGRP.
From 1990 to 1993 I visited Chapel Hill for six weeks in the fall and spring. It was a great privilege to learn how to identify and classify the sensory properties of these neurons in vivo directly from such a master of sensory receptor-type analysis (see Alan Light’s excellent remembrance above, and Light and Perl, 1993, below for historical perspectives). Our prime focus was on the classification of nociceptors, a field to which Ed, with his co-workers, had contributed so much (Burgess and Perl, 1967; Perl, 1968; Bessou and Perl, 1969; Bessou et al., 1971; Perl et al., 1976). The preparation was guinea pig L6 and S1 DRGs. Some days, but not others, we easily recorded from C-fiber nociceptors. D hair units were few in these DRGs, but occasionally we recorded several in a day. Ed called these groups of similar neurons recorded on the same day the “Cell du Jour.” We never knew the cause, but differing electrode properties was a possibility. From this work, helped by Bennie Crepps and Juping Bao, we published papers on the sensory properties of DRG neurons with immunoreactivity for substance P (Lawson et al., 1997) and CGRP (Lawson et al., 2002). When writing these papers, and describing C-fiber units responding to noxious mechanical and heat stimuli, Ed was firm that the term “C polymodal nociceptor” should be applied only to those with a cutaneous receptive field as previously defined (Bessou and Perl, 1969). Those with deeper receptive fields we called “C mechanoheat units,” leaving open the possibility that they may differ from those with cutaneous receptive fields.
This precision exemplified an important aspect of Ed’s approach to science. Direct observation and accurate terminology were key. Recently, he suggested the term “pathological” pain as more precise than the commonly used term “chronic” pain. He was right, of course. The variable time scales of the chronicity make the term imprecise, especially in the context of preclinical research. The word “pathological,” meaning abnormal or disease related, is more appropriate.
Once this methodology was established in my lab in Bristol, UK, Ed stopped those experiments in his lab; he had passed that baton to me. In turn, I made sure that, whenever possible, my group and I provided training in these techniques to those wishing to learn them. Ed now focused on other important issues, such as connectivity of the dorsal horn, on which he did such exquisite work. My experience working with him, and our combined methodologies, formed a solid basis upon which, for the next 20 years, much of the research in my group relied and developed. This research included determining the sensory receptive properties and electrophysiological properties of DRG neurons 1) expressing Nav1.7, Nav1.8, Nav1.9, TrkA, IB4-binding, HCN1, HCN2, and TREK2, and 2) showing altered properties and spontaneous firing in models of pathological pain.
Ed and his wife Marge were generously hospitable. I stayed with them on multiple occasions. They fed me lovely food, with bread made by Ed, and we talked and talked. Sometimes they took me to their place at Oriental on the east coast of North Carolina where initially they had a wooden shed for tools and a shower, and later they had built a lovely hexagonal wooden house on stilts that overlooked the river where their boat was moored alongside their wooden jetty. You could look out on trees, reeds, herons, their boat, the river, and occasional passing boats. Their boat was a nonesuch design, which they called Dernier (the last [boat]). A single large sail with a wishbone boom rather like that of a windsurfer, only very much larger. We would sail, and sometimes we (Ed, Marge and I) stayed overnight on the boat at anchor, and Ed would cook. There was always a glass or two of wine. We even raced Dernier in the local annual Oriental sailing race. Dernier proved too large, and they downsized to a smaller (still quite large) nonesuch and called it Dernier2, the second last [boat]! They eventually contented themselves with their lovely house overlooking the river.
Ed and Marge also visited Bristol for Physiological Society meetings. This enabled me to entertain them a little and show them around Bristol. Ed joined in experiments with my group, demonstrating methods of defining certain sensory properties of DRG neurons even more precisely.
I count Ed and Marge as very dear friends. We always met up for a meal when at scientific meetings, and I dropped in to Chapel Hill to visit them whenever I had the chance.
Sometimes he would give me one of his lab’s manuscripts to comment on. My last visit was two and a half years ago with my husband, Tim Biscoe. Ed and I spent some hours talking science. He gave me good advice on a paper (which I followed with the help of Cristian Acosta). Then, the four of us went out for a meal together—a lovely evening. Sadly, Ed was injured in a fall the next day and was taken to a rehabilitation center to recuperate.
Ed was the best kind of friend and mentor—caring, supportive, interested, but remaining always capable of being constructively critical and intellectually challenging. I miss and shall miss him. I will continue to keep in touch with my friend Marge, who, it always seemed to me, was the light of Ed’s life.
Reference
1. Alan Light and Edward Perl, "Peripheral Sensory Systems," in Peripheral Neuropathy Edition 3, ed. PJ Dyke and PK Thomas, 1993, 149-165.
Mark Zylka, University of North Carolina at Chapel Hill
Perl Memorial Lecture to
Perl Memorial Lecture to honor the life and work of Dr. Edward Perl, January 23, 2015.
As nicely described by Dr. Light above, Dr. Edward Perl spent the majority of his scientific career here at UNC. I started my lab at UNC in 2006, with Ed as my immediate lab neighbor and colleague. Like so many others, I benefitted greatly from his wisdom in the areas of pain and neuroscience, and miss him greatly. He was an inspiration and a true pioneer.
I also wanted to let everyone know that our department will be holding a memorial lecture in his honor on January 23, 2015. Several of Dr. Perl’s former colleagues will be in attendance and anyone who wishes to come is welcome.
Dr. Thomas Jessell, a former Perl-UNC Prize recipient, will be giving the keynote lecture, entitled “Perl, Pain and Proprioception in Perspective.”
I will give a brief introduction, covering Dr. Perl’s life and major scientific contributions.
If you are interested in attending, please register for this free event here (so that we know how many people to expect): http://www.med.unc.edu/cellbiophysio
As you may know, Perl endowed a prize, known as the “Perl-UNC Neuroscience Prize,” that carries a $20,000 award and is given to recognize a seminal achievement in neuroscience. Six former Perl Prize recipients have gone on to win a Nobel Prize. Should you be interested, the Perl family and Department of Cell Biology & Physiology kindly suggest a contribution to the Perl Neuroscience Prize Fund. Please visit http://www.med.unc.edu/neuroscience to make your donation (select “Make a gift”).