March 2013
Volume 54, Issue 3
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Lecture  |   March 2013
Introducing Peter Sterling, the 2012 Recipient of the Proctor Medal
Investigative Ophthalmology & Visual Science March 2013, Vol.54, 2266. doi:10.1167/iovs.12-10693
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      Richard H. Masland; Introducing Peter Sterling, the 2012 Recipient of the Proctor Medal. Invest. Ophthalmol. Vis. Sci. 2013;54(3):2266. doi: 10.1167/iovs.12-10693.

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      © ARVO (1962-2015); The Authors (2016-present)

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Introduction
Peter Sterling was born in 1940 in New York, the son of Phillip and Dorothy Sterling, well-known writers and advocates for progressive causes. He was raised and schooled in New York, and attended Cornell University, graduating in 1961. This was at the height of the American civil rights movement, and Peter was involved passionately, protesting and organizing in support of desegregation in the deep South. One of these protests led to his arrest. He paid a fine and was released. Despite this criminal record, he was admitted to medical school. 
He attended New York University Medical School for two years, dropping out because he found neuroanatomy more interesting than medical school—interestingly, a path followed a few years earlier by another noted retinal scientist, John Dowling. The choice of neuroanatomy, not always considered a sexy subject, was prescient, as a recent resurgence has made structural neuroscience one of the hot areas in modern neurobiology. 
Sterling's PhD work was at Western Reserve University with Hans Kuypers, a hard line, old school neuroanatomist. However, Kuypers also was a technical innovator: He was the first to show that single neurons could be labeled by multiple fluorescent markers. There are echoes of his teacher in Peter's later innovations in anatomic technique. His work at Western Reserve was on the fine structure of the spinal cord. 1 After a post-doc in the then-new Department of Neurobiology at Harvard, he moved to the Department of Anatomy at the University of Pennsylvania, where he has remained for the rest of his career. He was appointed Professor of Neuroscience in 1980. 
Peter Sterling has made many scientific contributions, with three perhaps the most significant. The first, and possibly the most noticed by the broad neuroscience community, was his use of 3-dimensional electron microscopic reconstruction of the retina. His advance was to realize that the microcircuits of the retina are small enough that whole circuits can be encompassed in series of thin sections. 2 He developed techniques to implement this, and used the resulting image stacks to work out the relations between ganglion cells and their afferent inputs—from rods and cones all the way down through the retina. 3 This was exciting: it created a concrete template—a visible paradigm for how the retina's microcircuits look. It also was grandfather to an exciting area that now goes under the name of connectomics, which simply is a large scale version, finally possible because of inexpensive computing power, of the analysis Peter pioneered 25 years ago. 
The second contribution came not from direct experimental work, but from his writing on the overall organization of the retina. This is embodied in a long essay, published in Gordon Shepard's collection titled, “The Synaptic Organization of the Brain.” 4 Sterling's chapter, which was updated twice in subsequent editions, is not what we usually call a “review.” Instead, it is a synthesis. In it, Peter dove headlong into explaining WHY things are as they are. This traditionally is called “teleological thinking,” and graduate students are taught to avoid it. Peter embraced it, in the process clarifying everyone's thinking and attracting a new generation of workers to the field. A fine paper by Sterling et al. has the subtitle of “Why ballplayers don't wear shades.” 5 They point out that outfielders do wear long-billed caps, but on even the brightest days rarely wear sunglasses. They explain—in terms of the structure and biophysics of an individual retinal cone—why the outfielder's eye needs to capture every last photon emitted by a rising fly ball. 
Finally, Sterling has created a bridge between theoretical thinking and anatomic structure—between the very top-down and the truly bottom-up. A special focus has been the information-carrying capability of the retina, at both the scale of a ganglion cell collecting synaptic inputs from bipolar cells and the much finer scale of synaptic vesicles fusing to release their neurotransmitter. 6,7 This is structural neuroscience at its best: a reduction of speculative thinking to the firm evidence of actual physical objects. 
Throughout the years, Peter has continued to be engaged in social issues, and has published on subjects as varied as psychosurgery and the overuse of psychoactive drugs. He also has been an engaged teacher and mentor, winning a prize for his teaching of medical students and mentoring many scientists who are now prominent in the field. Last year he participated in the celebration and re-enactment of the original civil rights Freedom Rides. 
Those who know him learn, sometimes to their discomfort, that Peter Sterling is an intense person. He does not do things half-heartedly. Peter embodies the advice of the preacher in Ecclesiastes: “Whatever thine hand finds to do, do it with thy might.” This informs both his concern for social justice and his intellectual life. Sometimes combative, visibly thrilled at the adventure of finding things out, Peter is a fully engaged scientist. Not only his experiments, but his joy and passion have enriched our field. 
References
Sterling P Kuypers HG. Anatomical organization of the brachial spinal cord of the cat. I. The distribution of dorsal root fibers. Brain Res . 1967; 4: 1–15. [CrossRef] [PubMed]
Stevens JK McGuire BA Sterling P. Toward a functional architecture of the retina: serial reconstruction of adjacent ganglion cells. Science . 1980; 207: 317–319. [CrossRef] [PubMed]
McGuire BA Stevens JK Sterling P. Microcircuitry of bipolar cells in cat retina. J Neurosci . 1984; 4: 2920–2938. [PubMed]
Retina Sterling P. In: Shepherd GM ed. The Synaptic Organization of the Brain . New York, NY: Oxford University Press; 1990: 170–213.
Sterling P Cohen E Smith RG Tsukamoto Y. Retinal circuits for daylight: why ballplayers don't wear shades. In: Eeckman FH ed. Analysis and Modeling of Neural Systems . Boston: Kluwer Academic Publishers; 1992: 143–162.
Demb JB Sterling P Freed MA. How retinal ganglion cells prevent synaptic noise from reaching the spike output. J Neurophysiol . 2004; 92: 2510–2519. [CrossRef] [PubMed]
Matthews G Sterling P. Evidence that vesicles undergo compound fusion on the synaptic ribbon. J Neurosci . 2008; 28: 5403–5411. [CrossRef] [PubMed]
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