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UW PBIO – A Brief History (1946-2010)

The Department of Physiology and Biophysics
University of Washington School of Medicine
A Brief History

By Marjorie E. Anderson, PhD
(Based in part on an earlier history by Allen M. Scher, PhD)

A review of the many factors that led to this departmental status draws attention to the concurrent development of the department and NIH extramural research and training grants, strong departmental leadership, collegial faculty, and extensive training programs with many predoctoral and postdoctoral trainees.

The following history relies heavily on the annual reports written by each chairman to the dean from 1946 to 1973. Chairmen during that time were Dr. TC Ruch and Dr. HD Patton. Information from those reports is supplemented by the memories of those who were there.

The beginning: Developing a highly-rated department

The responsibility for recruiting chairs to the new School of Medicine fell to Edward Turner, MD, who became the first dean in 1945. His initial choice to chair the Department of Physiology and Biophysics was Robert Pitts, a renal physiologist. Pitts withdrew before he moved to Seattle, however, and one wonders what direction the department would have taken if a renal physiologist had led it instead of Theodore C. Ruch, PhD, a neurophysiologist from Yale. Allen Scher, PhD, who had responsibility for teaching renal physiology in lieu of Pitts, pointed out that it took a long time for the glassware ordered for Pitts to be depleted by the more physically-oriented laboratories of the early members of the department!

TC (Ted) Ruch, PhD (“The Professor”) took the reins as the first chairman of the department in 1946. Ruch had had initial training in psychology at Oregon and Stanford, was a Rhodes scholar at Oxford, and received a PhD in physiology at Yale. He worked with John Fulton and, thus, followed the neurophysiological “lineage” from Sir Charles Sherrington. He stayed on the Yale faculty for 12 years before leaving to build the new department in Seattle.

Ruch’s departmental plan and initial faculty recruitments

Dr. Ruch outlined his plan for the department in the first of the annual reports to the dean (1946). His department was to have three divisions: medical physiology, biophysics, and applied physiology. In addition to Ruch, the initial faculty included three from other departments on campus: Loren Carlson, PhD (temperature regulation and respiratory responses to high altitude), Arthur Martin, PhD, and Julia Skahen, PhD (endocrinology). Two new faculty, Harry D. Patton, MD, PhD (neurophysiology) and Robert Rushmer, MD (cardiovascular physiology), were recruited during the first year. These new appointments strengthened what Ruch called the medical physiology division. Patton had been trained in neurophysiology under Ruch at Yale and came to Seattle via Johns Hopkins, where he had a brief faculty appointment. Rushmer’s medical degree was from Rush Medical College, and he had clinical experience at Mayo. He also came with research experience in the US Air force, where he had been on the faculty of the School of Aviation Medicine prior to his move to Seattle. Rushmer brought a clinically-directed research interest in cardiovascular physiology when he joined the department in 1947.

Ruch’s emphasis on biophysics is interesting, since his own training leaned more toward psychology. In the 1947-48 annual report he notes that biochemistry was largely taking over functional studies that lent themselves to chemical analysis, and anatomists also were more engaged in functional studies. (This was presumably due largely to introduction of the Horseley Clark stereotaxic apparatus, which allowed lesions to be made accurately in subcortical brain structures.) Ruch wrote, “Physiology’s future lies in teaching i) physical aspects of function, ii) reactions of the body to physical environment and iii) certain broad, integrative principles. Therefore, the development of biophysics within a physiology department will clearly place that department at the front of the modern trend” (1947-48 annual report, pg 5). Ruch’s definition of what a biophysics division in the department would do was somewhat different from what current biophysics faculty might envision, however. In the 1947-48 annual report, Ruch says, “One of the duties of biophysics, comparable to research, is to develop and publish descriptions of improved research and teaching apparatus. “This was to be not only for the PBio department, but also for other departments. This must have been a part of Ruch’s efforts to keep funding for electronics in the department, and at the beginning of his review of the biophysics division he says, “The Administration has wisely made Biophysics a subdepartment of Physiology” (1947-48 annual report, pg 5). In fact, the department’s name was Physiology and Biophysics from the beginning.

In the early post-WWII days of the department, it was difficult to secure “young men” for assistant professor or instructor positions due to the “lost generation” of the WWII era. Vahe Amassian, MD, was recruited from England in 1949, however. He had come from the Cambridge school of neurophysiologists (Adrian, Matthews, Rushton), and he occupied a four-year instructor position that Ruch intended to use in a rotating fashion to “bring recently qualified British or European men with physiological experience…. to keep in touch with developments in European laboratories” (1948-49 Annual Report, pg 2).

The problem of recruiting and retaining faculty who will “rise to the top” was a consistent one, in the face of offers from other institutions. Ruch’s annual lament repeated the salary and space inequities, and in the 1949-50 annual report Dr. Ruch pointed out that UW salaries already were not competitive with those of other institutions. Nevertheless, during the same year (1949-50), two new graduates who were just finishing their degrees were recruited as instructors. These were Allen Scher and J. Walter Woodbury.

Allen M. Scher, PhD had completed his PhD at Yale and served in the military. His dissertation dealt with renal circulation, and it was for this reason that he initially took on the role of the department’s renal physiologist when he arrived in 1950.

The first biophysicist hired to the departmental faculty was J. Walter Woodbury, PhD, who came in 1950 with a PhD in medical physiology from the University of Utah. Woodbury had just started his undergraduate studies in physics at Utah when Pearl Harbor was bombed and the US entered WWII. He quickly finished his BS and joined the Radiation Laboratory at MIT as a Staff Member in 1943. The mission of the lab was to test and carry into pre-production radar sets for use on land, sea and air, an important factor in the war. He had some interesting educational experiences tuning the radar sets on various naval vessels on their shakedown cruises. When the war ended, the Radiation lab was quickly shut down, however, and he entered Graduate School at the University of Utah.

Woodbury’s oldest brother talked him into going into physiology, instead of physics, and he earned a Master of Science degree in physiology in 1947. His professors worried that his career would be hampered if he received all three degrees at the University of Utah, but they wanted him to be their first PhD in physiology. They got around this problem by considering his two years at the Radiation Laboratory plus a tour of neuroscience labs in the east as enough outside exposure. His first stop on the tour was at the Department of Physiology at the University of Chicago, where he spent six weeks working with Gilbert Ling. He learned how to pull Ling’s newly perfected micropipette electrodes by heating a short piece of capillary glass tubing in the fringes of a Bunsen burner flame. Woodbury says, “We experimented nearly every night for most of the night, and we gathered enough data to write a paper on the effects of temperature on the resting potential of frog Sartorius muscle.” After he finished his PhD, Ruch recruited him to Seattle. Dr. Ruch was especially impressed by the fact that Woodbury had six full-length publications in national journals during his predoctoral career, and he came from a distinguished academic family (1949-50 Annual Report, pg 1).

In Seattle, Woodbury used his experience in intracellular recording to study the membrane potentials in many different types of cells. His bibliography lists studies of spinal cord (1952), frog sartorius muscle (1953), uterine muscle cells (1954), pregnant uterine muscle (1954), frog ventricular muscle, human heart (1957), and cultured chick embryo heart muscle cells. Especially important was his use of a flexibly mounted ultramicroelectrode to record intracellularly from moving tissues, such as the beating heart (Woodbury and Brady, Science, 1956).

Woodbury and Patton presented some of the first intracellular records taken from mammalian spinal elements, including motoneurons (Cold Spring Harbor Symposium on Quantitative Biology, 1952). This work was little known, however, with the usual credit for the first intracellular recording from spinal motoneurons going to Sir J. C. Eccles. In his book, Creating Modern Neuroscience, The Revolutionary 1950s, Gordon Shepherd says that Dexter Easton, who came to Eccles’ lab in New Zealand after being in Seattle, reported to Eccles that Woodbury and Patton were recording intracellularly from spinal elements. Eccles immediately turned all of his attention to this goal and carried out the series of studies of synaptic transmission in the spinal cord that would lead to a share of the 1963 Nobel prize for physiology and medicine (Shepherd, 2010). This was accomplished with the expertise of John Coombs, who constructed the necessary electronics, especially a high impedance negative feedback amplifier, and of Laurence Brock, who made the glass microelectrodes. Eccles also commandeered some of the equipment that Archie McIntyre had brought back to New Zealand from the Rockefeller University (Shepherd, pg 92). Woodbury and Patton, meanwhile, went on to other things.

After a 4-year search for another biophysicist to join Woodbury, Allan Young was chosen. He had a PhD in physics from the University of Toronto, but he also was well-trained in physiology and had an extensive mathematical background. Young, who came to Seattle initially as a research associate, had interests in respiration and temperature regulation. These were similar to those of Loren Carlson, who came from the UW’s Department of Zoology. Carlson’s interests more closely met Ruch’s vision of an applied physiology division of the department. This included the physiology of exercise, which the war had shown was important to the military.

Of importance in early recruitment efforts was the availability of housing. Housing was scarce in the immediate post-war era, and many of the medical school’s initial faculty lived in the university’s Union Bay housing complex. The Pattons, for example, lived in a two-bedroom unit. Barbara Patton reflected that, “It was fun; we made friends at Union Bay, as well as in the department. When the faculty and their families got together, we all fit in one room”. Union Bay, a camp of one-story huts, was still used for short term housing when Bertil Hille arrived in 1969. The Hilles lived there for two weeks, until the kerosene furnace caught fire and fire engines came!

Faculty loss

Recruitment of new faculty was counterbalanced by the loss (or expected loss) of others. The 1950-51 annual report stated that Dr Rushmer had become superintendent of Research at the Naval Acceleration lab in Johnsville, PA and Professor of Applied Physiology at the University of Pennsylvania. It seems that Rushmer must have changed his mind, however, since he and his activities are cited in each successive annual report, and nothing further is mentioned regarding his departure.

Dr. Loren Carlson, one of the three professorial faculty who came from the UW department of Zoology, was the first of the initial professorial group to leave. In 1959 he accepted the chairmanship of Physiology at the newly-founded Kentucky School of Medicine. His experience as a founding faculty member in the new department at UW, as well as a period of time as Associate Dean, undoubtedly gave him valuable experience in his new position.

Dr. Vahe Amassian left the department in 1956 to accept a professorship at the State University of New York (SUNY) Brooklyn. He had taken a military leave of absence in 1955-1956, and he returned at a time when Ruch said that the departmental “ammunition”, i.e. money for salaries, was “virtually exhausted” (1956-57 annual report, pg 1).

The faculty of the UW PBio department—and the entire medical school—have always been a friendly group of colleagues. Perhaps that –plus Mt. Rainier—are significant reasons why only 10 of the 53 regular and research professorial faculty appointed during its 64-year history (1946-2010), have left for other universities or research institutes.

A list of all of the “teaching” faculty with primary appointments in PBio and some of the joint appointees from other departments can be found in Appendix I.


The initial efforts of the new faculty in a new school of medicine (the first students were admitted in autumn, 1946) were directed toward setting up the teaching program for medical students. Ruch valued good teaching, and he believed that teaching should not be left to student assistants. In the model of Sherrington, Priestly, and Haldane that he had encountered in England, he believed that, “Laboratory teaching is considered not only worthy of the attention of senior staff members, but worthy of their undivided attention” (1946-47 annual report, pg 3). All faculty also attended all of the lectures in the medical student physiology course (they did this at least until 1965), and Dr. Ruch gave junior faculty feedback on their lectures.

Woodbury’s experience was different, however. He says, “After my very first lecture, given in the winter of 1951 on the nerve-muscle synapse, I went to Ted’s office and asked him directly for feedback. He seemed startled, and after a few moments, he mentioned that my voice was a bit high. I agreed with this, but there was not anything that I could do about that. I hasten to add, however, that I snowed the students and probably the faculty by basing my lecture on the then recent classical 1951 paper of Fatt and Katz that clearly established that synaptic transmission is chemical in nature. I speculate that Ted didn’t follow what I said and hesitated to tell me that I had blown the lecture, that I was much too detailed and at too high a level. I didn’t ever ask him again and he never offered any suggestions.”

Ruch’s guiding principles for medical school teaching were that (1) it should emphasize mammalian, human, and clinical physiology, (2) it should emphasize individual instruction, largely through small group classes, and (3) it should include an emphasis on first-hand laboratory experience, using research level equipment. Graduate physiology students and medical students took the same first year physiology courses, and all rotated through three labs on a general topic in a 3-week cycle. Starting in the mid 50’s, groups of four students also spent the entire spring quarter conducting a research project on a single question, the Project Laboratory. Departmental faculty and some from other departments sponsored projects. The results of these projects were written up as scientific papers and presented to the entire class at the annual meeting of the “University of Washington Physiological Society”. Copies of the Project Reports were kept in the departmental library. This first-hand laboratory experience undoubtedly led to the interest in research by at least a few medical students, some of whom (such as Wayne Crill, Marcus Raichle, and Kenneth Casey) pursued extensive research careers, in addition to their clinical practice.

Ruch was very proud of this alternative style of teaching, instead of the cookbook type of experiment. He also encouraged conjoint teaching, such as that done by the anatomy and physiology departments in the areas of endocrinology and neurophysiology. In spite of the emphasis on creative teaching styles, however, Ruch’s frustration with the student behavior is evident in the 1955-56 annual report, in which he says, “Our students are studying courses and not a subject. They are studying to pass examinations and not to learn—they will study only lecture notes if this will suffice for survival.” He continues, “It is almost as though we were teaching students with split personalities, at one time they appear to be mature, eager, and penetrating and at other times they seem to be methodically eating their way through a vast diet of verbal, especially oral foliage without much digestion and little assimilation” (1955-56 Annual report, pg 1). Would the faculty make similar assessments in 2010?

Graduate training

After the first year’s physiology course, graduate training was individualized. In place of advanced lecture courses, masters’ students were required to write a weekly essay based on experimental literature. This was then discussed in a conference with an appropriate faculty member. This method of instruction was flexible and accommodated graduate students with different backgrounds and objectives, as well as medical students who also desired an MS degree. June DeVito, the second recipient of a PhD from the department, relates that Dr. Ruch seemed to have forgotten her, and it seemed as if she wrote essays for several years before she finally was given the go-ahead to finish her dissertation!

The first graduate degree awarded by the department was a Master of Science granted to William Wolfe, MD in June, 1950. The first PhDs in physiology and biophysics were granted to Pai Chin Tsang (1953, TC Ruch, advisor) and June DeVito (1954, TC Ruch, advisor). By the end of the ‘50’s, 8 PhD degrees had been awarded, with topics in environmental physiology, biophysics, neurophysiology, and cardiovascular physiology. Appendix II lists all of the 289 students who have earned PhDs working with PBio faculty between 1953 and 2010 including those whose degree was in PBio and those with degrees in multidepartmental degree programs, such as Physiology/Psychology, Neurobiology and Behavior, Molecular and Cell Biology, and the MD/PhD program, officially named the Medical Scientist Training Program (MSTP).      


Initially, space was limited and scrounged from available sites. A remodeled portion of a building erected for the Alaska-Yukon Exposition (Physiology Hall) housed physiology. In the 1947-48 report, Ruch says that “transformation of a one-time art gallery to chemical laboratory to physiological laboratories has been largely completed” (1947-48 annual report pg 2).

The first phase of the Health Sciences building was completed in 1949, and the Department of Physiology and Biophysics occupied its space there in 1950, just in time to take advantage of available NIH grant funds. By 1957-58, however, Ruch pointed out that the departmental needs had already outgrown the available space, and that plans for new construction must be included in connection with the planned Pharmacy wing.

By the 1961-1962 annual report, Ruch emphasized the space problem by pointing out that the department trained over 33 percent of the graduate students and about 33 percent of the postdoctoral fellows that were trained in all the Basic Science Departments of the School of Medicine and 11 percent of all postdoctoral fellows receiving physiological training in the United States. This commitment to training had not been matched with a commitment of space in which this training could be done.

The shortage of space has been a consistent problem throughout the years, but perhaps the most urgent statement by Ruch of the problem is in the 1962-63 annual report, in which he states, “A men’s toilet-shower room has been converted into an experimental room, with resulting vesicle hardship” (1962-63 annual report, pg. 13).

Electronics shop

Because physiological studies require specialized equipment, Ruch developed a strong electronics shop, which he considered a part of the biophysics section of the department. The staff-built equipment not only for the PBio department, but also for other departments in the medical school. R. S. Bark, the Electronics Engineer who headed this section, assembled photographs, specifications, schematic diagrams, and operating manuals of items built from March, 1946 to November, 1950 in a bound volume kept now in the departmental library. It starts with single channel strain gauge amplifiers and DC amplifiers that would provide signals to recording instruments such as spirometers and direct-writing recorders used in respiratory and cardiovascular studies. The division also designed and built recorders, (such as electric stylus marking oscillographs), stimulators used in neurophysiological studies, power supplies, and ultrasonic generators to be used, initially, in cancer studies. Ultrasound was a modality used heavily by Rushmer in his cardiovascular studies, which later became the basis for numerous clinical applications of ultrasound.

Dr. Ruch noted with particular pleasure in the 1959-60 annual report that the medical student labs were finally equipped with chart recorders, replacing smoked drum kymographs. The result was, “Now the mechanics of recording can be relegated to the background and the student can concentrate upon the physiological issue” (1959-60 annual report, pg. 3). Smoked drum kymographs were still used by Drs. Skahen and Landau for the physiology classes that they taught for nursing students, however.

For those unfamiliar with the smoked drum kymograph, George Brengelmann recalls, “There is the memory of the lab facility used by Skahen and Landau, another reflection of Ruch’s commitment to offering substantial laboratory experiences, I suspect. In my recollections dating back to 1959, the facility included a large stainless steel chamber in which paper strips were coated with carbon from a dense smoke. These were fitted around a cylinder about 9″ in diameter and 18” height rotated by clockwork that had to be kept wound up, a “kymograph”. For longer recordings than permitted by the circumference of the kymograph drum, long loops of smoked paper were made with one end slipped over the kymograph drum and another on a second drum several feet away.” Devices such as levers attached to frog muscles scratched away carbon to reveal the white paper below. Records were preserved by a coat of shellac. Other transducers included capsules for pressure measurement in which a diaphragm that bulged in and out with pressure change was linked to the recording stylus and various other clever gadgets that mechanically amplified faint signals. Memories of the laboratory include spectacular messes involving carbon dust and shellac accidents.”

A special early challenge was presented by the equipment needed for Dr. Carlson’s studies of arctic acclimatization carried out for the US Air Force. Three expeditions to the Arctic, during the winters of 1948-49, 1949-50, and 1950-51, all used equipment designed and built by the PBio electronics division. One of the last pictures in the volume assembled by Bark is of a portable telemetry transmitter system that would be worn on a person’s back and would carry signals from twelve heat loss, temperature, and other body pick-up channels (Fig 2). Ruch and the departmental faculty also recognized the importance of graduate student experience with instrumentation to be used in their research. This was emphasized by a course, given by Drs. Young and Woodbury, on the principles of electricity and electronics.

early amplifier photo
Fig 1. Early amplifier made by the PBio electronics shop
cold weather suit photo
Fig 2. Cold weather suit with telemetered signals

Support for research and research training

Although the teaching mission demanded considerable faculty time and effort in the first few years, Ruch’s reports make it clear that research was a major priority for all of the faculty. By the 1947-48 annual report, research was described on the clinical physiology of the bladder, intracardiac pressure, and the relative pressure of lymph and tissue fluid. By the next annual report (1948-49), the five professorial track faculty all had research publications.

The first grants awarded to the department also were reported in the 1948-49 annual report. Initially, the state was so enthusiastic about supporting the school that biomedical research funds were made available from a tax on liquor by the glass. This money was easily available for a variety of research topics until the legislature targeted the funds to studies of alcoholism.

The medical school at the University of Washington began just as the research grants program of NIH expanded in 1946 from the successful extramural grants program of the National Cancer Institute to a comparable program in the entire NIH. The UW School of Medicine and the Department of Physiology and Biophysics were poised to participate fully in these “golden years” of NIH expansion, and in the 1958-59 annual report, Dr. Ruch recommends that an increase in NIH grant indirect cost allowances be returned to the department to help offset decreases in state support (1958-59 report, pg 6). Ruch says, “It is difficult to escape the conclusion that before long the function of the University will be to supply space, heat, light, janitor service, library and administrative expenses while the faculty through overhead will pay for all research and a large share of teaching expenses” (1958-59 report, pg ?). Sound familiar?

National and international activities

The NIH saw the young, collaborative research-oriented faculty at UW as an ideal group for many of its new programs, and faculty, including Dr. Ruch, were invited to serve on NIH committees that helped to shape NIH programs and policies.

In 1961, Dr. Patton was invited by NIH to join a small group of eminent scholars for a month in Russia. He was appointed to the physiology committee of the National Board of Examiners for several years, and he served on NIH study sections and on the council of the National Institute of Neurological Diseases and Blindness (NINDB).

Allan Young and Robert Rushmer were participants in meetings that had to do with the formation of a Biophysical Society and NIH excursions into the fields of biophysics and biophysical chemistry (1955-56 annual report). By 1954, departmental faculty were represented on five grant-dispersing NIH review committees (pg 2, 1953-54 annual report).

TC Ruch was appointed to committees of NIH and the National Academy of Sciences that were charged with making the country self-sufficient with respect to monkeys for laboratory study of health and disease. These would later pave the way for the National Primate Research Centers, of which Ruch was one of the initial directors.

Training grants

Faculty participation in early review boards and NIH planning committees must have been an important factor when the first training grant opportunities were announced. These followed the increase in research grant funding, and in 1955, the PBio department was awarded the first neurophysiology training grant in the country to a preclinical department from the National Institute on Neurological Diseases and Blindness. It was designed to furnish the opportunity for clinicians to spend one or two years learning the techniques and philosophy basic to neurology and to give basic scientists a chance to learn about the problems of clinical medicine. Ruch lamented that “…clinical departments of Neurology are about as far behind modern recording methods as they are ahead of the smoked drum kymograph” (Annual Report for 1955-56, pg 3). By 1957, NIH had been able to establish only one other training program in neurophysiology. Instead, it expanded the UW program on an experimental basis.

One product of the desired cross training between clinical and basic science programs was the “Neurological Study Unit”, a series of bimonthly conferences that drew students of all years and several departments. Attendance was routinely around 50 or more, and it included white-coated faculty from the East—Neurology and Neurosurgery—and basic scientists from upper campus (Psychology) and from the West—P Bio, Biological Structure, Pharmacology, and Biochemistry. The flyer advertising the topic for the week showed a human who stood under a tree and held the top of his skull in his hand, revealing the brain underneath. It would list a clinical case presentation by a resident or clinical faculty member and discussion of the physiology relevant to the topic, usually by a basic science faculty member. Prominent visiting scholars were included when possible. Arthur Ward, chair of Neurological Surgery, and the head of the division of Neurology, Fred Plum, Gus Swanson, or Phil Swanson, usually sat in the front row and were active participants in the conference. The Neurological Study Unit persisted into the 21st century, and Bill Spain, a neurologist trained in neurophysiology in Wayne Crill’s lab, still managed to organize conferences in the same format.

Orville Smith and Temy Kennedy were two of the first four postdoctoral trainees on the Neurophysiological Training grant, and both stayed as faculty in the department. Dr. Kennedy administered the Neurological Training grant until Dr. Patton retired. A cardiovascular training grant, led by Dr. Rushmer, was started in 1956, and by 1962-63, there were five training grants in the department: neurophysiology, cardiovascular physiology, biophysics, physiology, and physiology-psychology. In addition to these, departmental faculty also participated in training grants in dentistry and nursing. Al Scher remembers one faculty meeting in which Dr. Ruch expressed concern that so much federal research and training money would lead to outside control and influence.

Early Computerization

The departmental faculty realized early that direct input of data into a computer was an important step in advancing quantitative research. Analog-to-digital (A-D) converters were built by Dr. Allan Young and Edmund Brand, as described in the 1961-62 annual report (pg 10). These could be used to transform information from recording instruments to a digital form that could be fed into the University’s IBM 709 computer. A computer-based A-D system for the department had been funded by 1962, and Dr. Ted Kehl was appointed to the faculty to train students and postdoctoral fellows in the programming and other techniques necessary for computer-based quantitative physiological analysis. Drs. Woodbury and Gordon participated in the LINC evaluation program, which was sponsored by NIH.

For many years the LINC was the main computer for training graduate students because it was simple, useful, user friendly and easy to program in machine language. Kehl and his colleagues went on to develop an inexpensive dedicated laboratory computer called the LM2. This custom built laboratory computer became important in the research programs in a number of PBio research labs including those of Drs. Berger, Hille, Scher and Albert Gordon. For quite a few years there was a required hands-on course in which graduate students had to use the LINC. A workable program for performing some useful function was required for a passing grade.

A LINC computer was installed in Dr. Woodbury’s lab by 1964, and by 1965, a Raytheon 440 computer to be used by the entire department and the Primate Center. In fact, Dr. Bertil Hille joined the departmental faculty in 1968 largely because he would have access to a computer with A-D capability. He could use the Raytheon computer in Kehl’s facility all day long—on Saturdays.

International symposia

Even in its first decade, the department sought connections with other scientists nationally and internationally. In addition to the faculty position mentioned earlier as a way to keep in touch with European neuroscience, Dr. Ruch encouraged Dr Woodbury to organize an international symposium entitled: Microneurophysiology of the Synapse, Participants included such stellar scientists as David PC Lloyd, Stephen Kuffler, JC Eccles, Karl Frank, and MGF Fuortes (October 20-23, 1955).

In 1961 the department participated in hosting the international symposium, Nervous Inhibition, organized by Prof. Ernst Florey of the Department of Zoology. This symposium was held at the Friday Harbor laboratories in the San Juan Islands, and it provided the opportunity for UW faculty and trainees to interact with eminent neuroscientists from around the world. One particularly colorful story described Prof. Ragnar Granit arriving by seaplane, instead of by the usual ferry boat. And Suhayl Jabbur, a graduate student at the time from Lebanon, obtained a signed statement from Prof. JC Eccles that Eccles had never made a glass microelectrode. This was of great value to Jabbur, whose advisor, Dr. Towe, insisted that Suhayl make his own glass micropipette electrodes. If Eccles could win the Nobel Prize without learning to make microelectrodes, then Jabbur, from a prominent Lebanese family, was convinced that it was a skill that he could do without!

Other international symposia hosted by the PBio department included:

  • Neural Control of Limb Movement, July, 1986. Eberhard Fetz and Marjorie Anderson, Organizers
  • VI International Motoneuron Meeting, June, 2008. Marc Binder and Randy Powers, Organizers

The textbook

The department’s reputation certainly was enhanced by the faculty participation in writing and editing several editions of the well-known physiology text, Textbook of Physiology, which originally was edited by Howell and published in 1896 by WB Saunders and Co. The publication in 1960 of the 18th edition, now titled Medical Physiology and Biophysics, was the first that included Dr. Ruch as the co-editor and several UW faculty as authors of most of the book’s chapters.

Dr. Woodbury wrote the first textbook explanation of the Hodgkin-Huxley studies of the action potential, and he was asked to write chapters on the H-H quantitative analysis for several other publications, as well. In early 1963, Woodbury and other departmental faculty were asked to put forth a nomination for the Nobel prize. They agreed that Hodgkin and Huxley should be nominated, and Dr. Woodbury wrote the nomination, which the other faculty signed, as well. The 1963 Nobel prize was awarded to Drs Hodgkin, Huxley, and Eccles. Dr. Woodbury is sure that the textbook writings were the reason why they were asked to make a nomination.

The textbook continued for three more editions, with the final 21st edition, published in 1989, edited by five departmental faculty: HD Patton, AF Fuchs, B Hille, A Scher, and R Steiner. With the development of neuroscience and molecular biology departments and graduate programs, however, comprehensive graduate-level physiology courses were rare, and a comprehensive graduate-level text no longer was supported.

Other monographs and textbooks also were published by the faculty, beginning with Carlson’s monograph, Man in the Cold (1953), and Rushmer’s Cardiovascular Dynamics (1955). Table 1 lists monographs and textbooks written or edited by departmental faculty.

Carlson, L. Man in the Cold. 1953.
Rushmer, RF. Cardiovascular Dynamics. Saunders. 1955, 1961.
Ruch, TC. Diseases of Laboratory Animals. Philadelphia: WB Saunders. 1959, 600pp.
Ruch and Fulton. Medical Physiology and Biophysics, 18th Edition. 1960.
Ruch and Patton. Physiology and Biophysics, 19th Edition. 1965.
Ruch and Patton. Physiology and Biophysics, 20th Edition. 1979.
Patton, Fuchs, Hille, Scher, and Steiner. Textbook of Physiology, 21st Edition. 1989.
Stevens, CF. Neurophysiology: A Primer. 1966.
Hille, B. Ion Channels of Excitable Membranes. 1984, 1991, and 2001.
Rowell, LB. Human Circulation: Regulation During Physical Stress. 1986.
Rowell, LB. Human Cardiovascular Control. 1993.
Hlastala, MP. and Berger, AJ. Physiology of Respiration. 1996, 2001.
Rieke, F., Warland, D., de Ruyter van Steveninck, R. and Bialek, W. Spikes: Exploring the Neural Code. 1997.
(Written during postdoctoral years; Published after Rieke’s arrival in Seattle)
Howard, J. Mechanics of Motor Proteins and the Cytoskeleton. 2001.
Binder, MD., Hirokawa, N. and Windhorst, U. Encyclopedia of Neuroscience. Heidelberg: Springer. 2009.
Table 1. Books written and/or edited by PBio faculty

Joint graduate degree programs

Conjoint training in physiology and psychology was a natural for Ruch, who himself had graduate training in both areas. Discussion of a joint degree PhD program between physiology and psychology led to approval for such a degree-granting program by the graduate school in 1960-61. Initially the Joint Physiology/psychology program was administered by Dr. Orville Smith (PBio) and Dr. Montcrieff Smith (Psychology). Doctoral students in the Joint Program took all of the first-year courses and the qualifying exams in both departments and could choose a lab in either department for their dissertation research.

Also in 1960, NIH awarded UW a research training grant to support both pre- and postdoctoral training in physiology/psychology. This was the first training grant of its kind in the country and was managed by Dr. Mitchell Glickstein, a new recruit to the Primate Center. Dr. Glickstein had received a PhD in Psychology from the University of Chicago and did postdoctoral work with Dr. Roger Sperry at the California Institute of Technology and with Dr. Karl Pribram at Stanford University.

PBio has also been a department in which many of the MD-PhD students of the school have done the research training for their PhD. These students begin with the first of the courses for the medical degree, move to several years of primarily research training that culminates in the PhD degree, and then return to complete their clinical training. Although this program takes an extended period of time, students receive fellowship support for both basic science and clinical portions of their education, primarily from an NIH grant.

The MD/PhD program (Medical Scientist Training Program; MSTP) was established at UW in 1971.  PBIO faculty members Wayne Crill and Marc Binder served at times on the steering and admissions committees. A total of 20 students who have participated in the program performed their dissertation research in PBIO laboratories. Those students are identified as in the MSTP (medical scientist training program) in Appendix II.

The faculty of PBio and Zoology departments also developed a plan for a combined training program in General and Comparative Physiology, and this was approved by the graduate school during 1966-67. It was designed to emphasize research on marine species that were plentiful in Puget Sound and that resembled those used by investigators such as Nobel Laureates Hodgkin and Huxley, Professor Florey at UW, and Prof. Stephen Kuffler at Harvard. (In the UW PBio department, these species were used in Dr. Charles Stevens’ lab, and the tanks with “squishies and squeezies” [Dr. Patton’s terms] were included in any visitor tour led by Dr. Patton.)

The proposed Marine Biology center was to be developed on property owned by the university at Manchester, Washington. In contrast to the university’s laboratories at Friday Harbor, the Manchester site on the Kitsap Peninsula could be reached more easily by faculty who had ongoing responsibilities at the Seattle campus. Perhaps the development required was the stumbling block, because the program was never mentioned again in subsequent annual reports. The 1966-67 annual report does acknowledge that “—it is hoped that it will soon be submitted for approval by the Board of Regents” (1966-67 annual report, pg 12). Currently (2010), a small shellfish research laboratory at Manchester is located in facilities provided by the National Marine Fisheries Service. The UW School of Aquatic & Fishery Sciences is conducting studies on oysters, clams, and mussels to assist the shellfish aquaculture industry in the Pacific Northwest, but there are no physiological studies there.

Although a conjoint degree program was never developed between PBio and Bioengineering, several bioengineering students have done their dissertation research in the laboratories of PBio faculty, especially AF Fuchs, M Kushmerick, AM Gordon, and J. Howard.

Joint faculty appointments

The faculty, as well as the training programs, were often inter(multi) departmental. Joint appointments, with full voting rights in both departments, were most often made between clinical and basic science departments. In the best arrangements, this brought basic science techniques and expertise to clinical departments, (often including lab training experiences for clinical residents) and an awareness of clinical problems to the basic science faculty and trainees.


All was not work in Dr. Ruch’s era, nor in those that followed. George Brengelmann remembers Ruch entertaining faculty, students, and staff at his home on Lake Washington for an annual gathering on the day of the hydroplane races, and there were summer expeditions to his property in the San Juan Islands. Ruch enjoyed lively parties, liked to engage his guests in conversation on the many topics of his broad cultural interests, and showed a personal interest in gauging the tolerance of his colleagues for alcohol. On the summer outings to his San Juan retreat, he also just happened to have a few chores in line for which the help of some strong graduate students was duly appreciated.

The first chair transition: Program expansion and excellence amidst competition for resources

Dr. TC Ruch resigned as chairman on November 1, 1964 and turned his attention to his position as Director of the Regional Primate Research Center, which had opened in 1963. Dr. HD Patton, who had been Acting Chair since 1964, was appointed permanent Chairman of Physiology and Biophysics on March 2, 1966.

Dr. Patton took over a department that had achieved nationally recognized excellence. In a national survey, for example, the graduate program was given the top categorical rating, “extremely attractive”, and its ranking among the 106 programs evaluated was 4th, only .022 (of 4) points behind #1 Harvard (pg 3, 1965-66 annual report). In 1970, the graduate program was ranked #2, behind only The Rockefeller University (pg 80, 1970-1971 annual report).

The academic roster in the 1965-66 annual report listed 27 teaching faculty, which included those in professorial and research professorial tracks. These faculty supervised a large cadre of trainees, including 26 postdoctoral fellows and 38 graduate students. Most of these trainees were supported by training grants. Another 10 predoctoral trainees were in the joint Physiology/Psychology graduate program, and at least half of these were working with PBio faculty. Engineering students working with departmental faculty made up the last group of trainees.

Dr. Patton listed 78 major publications by his faculty in his first year as chair, although 47 of these were chapters in the new 19th edition of Physiology and Biophysics, accounting for the spike in publications shown for 1965-66 in Figure 3. The basis for the doubling of publications in 1971 is not clear, but it is clear that his faculty were quantitatively productive scientists, and it is likely that the several training grants that supported pre- and postdoctoral trainees were important contributors.

Fig. 3. PBio publications, students, postdocs, and faculty

Regional (later National) Primate Research Center at the University of Washington

The Regional Primate Research Center was started in 1961, with Dr. Ruch as director and Orville Smith as Assistant Director. Ruch rejected the university’s proposal to house the center in a west Seattle building that had housed a communications center during WW II because it was too far away. He also rejected a proposal to house the monkeys across the street from the clinical building, so that the “smelly monkeys” would not be near patients. Instead, Ruch insisted that the Primate Center- including the “smelly monkeys” be an active part of the campus, and he collaborated with Herschel Roman, the chair of microbiology, to obtain NIH funds for joint construction of a Primate/Cancer wing built as a westward addition to the existing G wing of the Health Sciences building.

Ruch’s insistence that the Primate Center be centrally located on the campus also facilitated his second goal, that all core staff have real faculty appointments in university departments. The G wing and the intersecting H wing housed the PBio faculty and labs, and this made it as easy as possible for Primate Center staff to be faculty in the PBio department. Including Ruch and Smith, seven of the first 12 core staff members hired by 1965 had faculty appointments in PBio. In addition, the Primate Center had a large affiliate program, by which other scientists could use the facilities of the Primate Center, and several affiliates also were PBio faculty.

Ruch’s philosophy was that scientists who were housed in proximity would find common interests and be more likely to collaborate. In fact, the labs were built in “flex style” with “rooms” separated by equipment, not walls and doors. This gave flexibility, but it also meant that the loud-speakers that every neurophysiologist uses to guide the electrode to the proximity of a single cell can be heard in adjacent labs. That, plus serendipity, played a role in the collaborative studies by Eric Luschei and Albert Fuchs that have been fundamental in understanding much of what we know about the neural control of eye position.

The serendipity part happened because this writer lived in a house with several other women during graduate school, and one of my housemates had known Albert’s wife, Susan, when they both worked at Yellowstone National Park. As a consequence, she and Albert came to visit and stayed at our house. When I learned that Albert was a bioengineer/ neurophysiologist who had done his PhD research on monkeys, I took him to see the Primate Center. Eric was there that Saturday, and Albert ended up applying for and getting a core staff job at the Primate Center, again with a faculty appointment in PBio.

The “brain music” heard throughout the Primate Center as electrodes penetrated different parts of the nervous system led to the collaborative early recordings by Albert and Eric from motoneurons and premotor neurons associated with eye movements. Eric was interested in the activity of jaw muscle spindle afferents in the mesencephalic nucleus of the fifth (trigeminal) nerve, a place where muscle spindle afferents could be accessed within the cranium. He did not find neurons with the pattern of activity that he expected during chewing or other movements of the jaw, but he stumbled across many neurons whose firing rates changed when the monkey moved its eyes. Albert recognized that these cells might be carrying the saccade-related activity that he and his graduate mentor, David Robinson, had predicted would be necessary to drive the rapid displacement of the eye in the orbit. Since abducens neurons were in the pontomedullary junction, it would be possible that Luschei’s electrode traversed this region, instead of the jaw-related region. Anatomical verification of the recording position supported this possibility, and the resulting manuscript, one of the first to describe the activity of motoneurons in the brain of a behaving animal, arrived at the offices of the Journal of Neurophysiology on the same day as the manuscript by Fuchs’ mentor, Dr. Robinson, who described the activity of neurons in the oculomotor nucleus that innervates a different set of extraocular muscles. In each case, the motoneurons produce a train of action potentials with an unmistakable sound that changes from one “pure” tone to another as the eyes move to a different position.

National visibility of the faculty

Faculty received recognition from national and international colleagues. This came in the form of invited lectures, awards from professional societies, editorial board positions, and appointment to national review committees, especially NIH study sections and positions on national advisory councils of the National Institutes of Neurological Diseases and Blindness (Patton) and the National Institute of Mental Health (Ruch). In the 1963-64 report written just before Patton changed from acting to permanent chair, he reported that all of the professors in the department served on NIH committees and/or editorial boards of major national journals.

Many of the faculty have received awards, especially from the professional societies. The highest awards have been to Bertil Hille, who came to Seattle because he would have access to computers. The major awards to Bertil have been the 3rd Annual Bristol-Myers Squibb Award for Distinguished Achievement in Neuroscience Research (1990), the 1999 Albert Lasker Award for Basic Medical Research, with Clay Armstrong and Roderick MacKinnon (1999), and the 2001 Gairdner Fnd. International Award with Clay Armstrong and Roderick MacKinnon (2001).

Fig. 4 School of Medicine reception honoring receipt of the 1999 Lasker award by Dr. Bertil Hille, with Drs. Clay Armstrong and Roderick McKinnon. Upper right: Dr. Hille. Clockwise: Dr. Marjorie Anderson, Mrs. Jean Crill, and Dr. Wayne Crill.

Changes in teaching

In 1968 the medical student curriculum underwent a major revision, and the result for medical students was an organ systems approach in which course committees, instead of departments, were responsible for the content, faculty teaching assignments, and course evaluations. Because the resulting courses were not appropriate for graduate students, the department had to develop a separate series of courses for graduate trainees. Considerable time and effort was required of the departmental faculty to develop both the new MD curriculum and the new PBio courses for graduate students. The split was initiated in the autumn of 1968. This also required adjustments in courses for other professional students- nursing, dental, and pharmacy- for which PBio had had responsibility for providing the physiology component of their training.

Patton argued that the new curricula—medical student and graduate student—would require 20 new faculty to maintain the quality of teaching, (!) and if no new space was acquired, this would leave only 500 sq ft/ faculty member. Such a space shortage led to Patton’s most persistent mantra throughout his chairmanship: We need more space!

Regional teaching: WAMI to WWAMI

In response to the need for more primary care physicians in the Northwest and the lack of medical schools in the surrounding states, in 1971 the UW Medical School began a regional medical education program including Washington, Alaska, Montana, and Idaho, the WAMI program. This program involved admitting students to the UW Medical School from these states and having the students from each state complete first year courses at a state university in that state and come to the UW for second year courses. They then had clinical clerkships at sites throughout the four state area. The UW was compensated by each state for the number of students enrolled from that state.

The WAMI program involved Physiology and Biophysics, since they coordinated the teaching of the Physiology course for first year medical students. The department had the responsibility both for teaching the course in Seattle and coordinating the teaching of this course at the five university sites in the four state area. It required working with faculty from the other universities to ensure comparable course content and common tests, and this was accomplished by yearly meetings and visits to the sites. In addition, PBio faculty occasionally needed to provide lecturers to ensure good coverage at each site. To compensate the department for the additional work, the department was provided with one FTE salary. The program started at the University of Alaska in 1971 and was expanded to Idaho and WSU the next year, next to Montana, and finally in 1996 to Wyoming, making it a WWAMI program. Additional expansion of the program to a site at Spokane has further increased the number of medical students trained. This program required substantial extra faculty effort, but it has successfully increased the training of primary care physicians and has expanded medical coverage in the 5-state area, all without starting new medical schools in those other states.

The physiology course itself changed over the years from one on control systems including physiology and some pharmacology to a course with common coverage at each site that emphasizes cell physiology. Albert Gordon coordinated the physiology teaching part of the WAMI (WWAMI) program the first 35 years. Peter Detwiler followed, with a number of other faculty assisting in the teaching. Although there are differences in the cell physiology course at each site that reflect the different instructors, the common tests have shown no consistent differences in student performance between sites.

Creation of the Division of Bioengineering

On July 1, 1968, a new Division of Bioengineering was created, with Dr. Rushmer as Director and Dr. Reid also leaving PBio to join Bioengineering. This was a joint effort of the schools of Medicine and of Engineering, and Dr. Patton opposed the separation of this part of the Department of Physiology and Biophysics. He especially opposed the loss of almost 3,000 square feet of space to Bioengineering (pg 62, Annual reports). Rushmer had been a candidate for the Chair of PBio at the time Dr. Patton was chosen, and considerable tension had developed between them.

Changes in funding for research and training

State funding diminished over the decades, and in 1965, the state contribution per student to maintenance and operations was 1/3 of what it had been in 1951 (1964-65 annual report, pg 17).

Federal funding for research also was in decline when Patton took over as chair. It had grown to the point where universities had become more and more dependent on it, but in the 1966-67 annual report (pg 16), Patton says, “Now that the increased cost of government involvement in Viet Nam is beginning to curtail funds available for such support, the stability of federal support is beginning to be threatened. In addition, Congressional demands for distribution of support on geographical grounds rather than on the basis of merit and feasibility threatens the review system.”

In spite of the decline in available funds, Patton reported in 1968-69 that a new cardiovascular training grant had just been awarded with Allen Scher as the PI, and the four recent faculty appointed in the department (Hille, Stirling, Miller and Luschei) all had been awarded NIH research grants (1968-69 annual report, pg 62). In addition, the training grants in Physiology, Neurophysiology, and joint Physiology/Psychology were renewed, and a new training grant in Oral Physiology was approved for 5 years, directed by Dr. A. C Brown.

The dependence on training grants to fund both graduate students and postdoctoral fellows left a large hollow when, under Secretary of Health, Education, and Welfare Casper Weinberger, training grants were “ruthlessly” cut out (1972-73 annual report, pg c49). Dr. Patton said, “We consider ourselves fortunate that we will be permitted to complete the training of fellows supported by the program before the cut-off date in February 1973, but we are not permitted to take any new trainees.”

In the decades to follow, training grants had to be multidepartmental, and students applied for support after their first year of study. This meant that funds accumulated by the departmental administration or pooled from the indirect costs from research grants funded the students’ first year, and they were supported thereafter by research grants held by their faculty advisors, training grants, if they competed successfully, or fellowships awarded to individual students. Even faculty on full state FTE salaries were urged to request some of their own salary support from their research grants. The funds recaptured from their state salary line were used to fund departmental infrastructure and first year graduate student stipends.

The cardiovascular program project grant (PPG)

In the mid 1980’s, five senior PBio faculty whose research focused on the control and regulation of the cardiovascular system decided to submit to the National Institute of Heart, Lung, and Blood (NIHLB) a proposal for a program project grant with Al Scher serving as the principal investigator (PI). A PPG required that participating PI’s relinquish their separate PI status on their individual grants and incorporate those grants into the PPG. Each investigator continued to control his project’s individual budget and research as before. The advantages included the structure of funding that allowed the whole project to exceed the sum of its parts and provide a significant financial saving, the administrative costs covered all seven projects, equipment costs could be shared, and a surplus of funds on one project could be used by another, if needed. A strong collegial atmosphere and scientific interaction pervaded wherein time, advice, funds, expertise, equipment, technical staff, etc., were shared for the common good.

In part, this atmosphere developed during regularly scheduled meetings, in which participants criticized each other’s experimental approaches positively, negatively, and openly. Larry Rowell says, “Appreciation by all was real, but not always instantaneous. It worked well, and Al Scher was an excellent PI.”

The research areas and directors of the seven projects in the PPG were:

  1. Allen Scher: Program project PI. Control of arterial blood pressure by baroreflexes, myocardial origins of the ECG complex.
  2. Arthur Brown: Circulation to bones and teeth. Temperature regulation.
  3. Eric Feigl: Control of the coronary circulation.
  4. Loring Rowell and George Brengelmann: Control of the peripheral circulation in humans; Role of circulation in human temperature regulation.
  5. Orville Smith (Director of the Primate Center) Central neural control of the circulation.
  6. Curt Wiederhielm: Microcirculation:Control of vasomotion and exchange of materials.
  7. Allan Young: Cutaneous blood flow: measurement by inert gas exchange.

Although other PPG applications were submitted by departmental faculty, this was the only one funded until Dr. Stan Froehner led a group in 2004 that successfully applied for a PPG to study Molecular and Cellular Therapies for Muscular Dystrophy. And some faculty (Albert Gordon) participated in PPGs initiated by faculty in other departments.

Different faculty roles

In addition to Rushmer’s move to chair the new Division of Bioengineering, other faculty took on responsibilities that pulled them away from full-time participation in departmental affairs. Dr. A. C. Brown spent 1969 as a visiting Professor at the University of Malaysia Medical School. (This was the first of several times that Dr. Brown taught abroad, and he continues to do this in retirement.) Dr. Thelma Kennedy was appointed Associate Dean of the Graduate School in 1969. At the time, she was the highest-ranking woman in the university administration, and she continued in that position until 1972. Dr. Robert Van Citters became Associate Dean for research in 1968, and he moved from this position to that of Dean of the School of Medicine in 1970. And Dr. Albert Berger filled the position of Associate Dean for Research and Graduate Education in the School of Medicine from 2001 to 2005.

While the department continued to be especially strong in what might be called Systems Physiology, it is noteworthy that Drs. Hille and Stahl offered a new course, Molecular Physiology, in 1970. This was the first reference to molecular approaches noted in the annual reports. Furthermore, it was reported to be “quite successful!” (1969-70 annual report, unpaginated).

Expanded social activities

Fig. 5 Patton’s boat, the Neuron

Dr. Patton liked to share his love of boating. He took the students on an annual day’s outing on his boat, the Neuron (later replaced by the Neuron II), but for a few weeks in the summer, he and his family were off to the San Juans or the Canadian gulf islands, often in tandem with the Arthur Wards (chair of Neurological Surgery).

A highlight of the year was the annual Holiday Party at the Pattons’ home. It not only brought forth the culinary skills of all who attended, but it also brought forth Dr. Patton’s guitar—and a few bawdy sea tunes. These competed with more traditional Christmas songs, led by his wife, Barbara, preferably accompanied by George Brengelmann.

Fig. 6 Eberhard Fetz (left) and Elsbieta Jankowska, together with other colleagues in the background on Mt Rainier during one of Jankowska’s visits to Seattle from Goteborg, Sweden.

The departmental variety of guitar styles also was showcased in the teaching laboratories. When auditory physiology was a topic and the cochlear microphonics were recorded from a cat’s ear, they were amplified and transmitted to the isolated nerve-muscle lab going on in the adjacent room, and from the audio speakers came ribald sea tunes (H. D. Patton), classical guitar music (Orville Smith), or 60’s folk and protest music (Dave Whitehorn, a graduate student).

Most of the students and postdocs did not come from the west. Neither did Temy Kennedy, the departmental graduate and postdoc program advisor for many years, but she married Rich Berner, a native Seattleite and climber with a wide knowledge of the local mountains. Temy and Rich led many a weekend outing to favorite hiking spots, and students and postdocs who left the northwest (and some of us who did not) took with them some knowledge of the mountains, as well as of science. Other faculty and labs also had traditions of ski outings or hikes, often with visitors to the lab. Dr. Larry Rowell held “Thursday seminars” at Alpental Ski area (with follow-up in the lab on Saturdays). He also jotted notes for his two books on exercise and cardiovascular control while his wife, Colleen, drove the car to and from the ski area.

Fig 7. P Bio Faculty (including some postdocs), 1971

Front Row: HD Patton, Barbara Walike, Louis D’Alecy, Joe Kim, Temy Kennedy, Chuck Stevens, Wayne Crill, Arthur Brown
Second Row: Orville Smith, TC Ruch, Tom Hornbein, Winnie Becker, ______, Eric Feigl, Albert Gordon, Maria Biedenbach
Third Row: J Walter Woodbury, _______, Renella Taylor, Donna Koerker, Allen Scher, Curt Wiederhelm, Bill Stahl, Bertil Hille
Fourth row: Ulrich Buettner, ______, Barbara Landau, Julia Skahen, George Brengelman, Charles Gale
Back row: Albert Fuchs, Eric Luschei, Bill Calvin, Arnie Towe, Ted Kehl, Judith Hildebrandt, Loring Rowell
Fig 8. P Bio Faculty at Pat’s retirement, 1983

Front row: Davida Teller, Joe Miller, Albert Berger, HD Patton, Temy Kennedy, Allen Scher, Donna Koerker, Robert Steiner
Second row: Orville Smith, Bill Stahl, Eric Feigl, Renella Taylor, John Conrad, Winnie Becker, Bertil Hille
Third row: Vahe Amassian, Glen Kerrick, Jack Hildebrandt, Barbara Landau, Phil Swartzkroin, Marjorie Anderson, Wayne Crill, Albert Gordon
Fourth row: George Brengelmann, Erich Luschei, Albert Fuchs, Eb Fetz, Marc Binder, Wolf Almers
Back row: Mike Hlastala, Arnie Towe, Peter Schwindt, Chuck Sterling

The second chair transition: Recruitment of cellular and molecular scientists. Glass at last!

When Wayne E. Crill, M. D. was appointed chair of Physiology and Biophysics in 1983, it was clear that the department lacked significant activity in modern cellular physiology and the molecular techniques used to study cellular function. Many of Crill’s nine regular faculty appointments were aimed at filling this hole. Crill’s philosophy was to appoint the very best candidates available, irrespective of the particular topic of research, and he competed with the best universities in the country to hire Drs. Bothwell, Carlson, Zagotta, Howard, Wordeman, Shadlen, Rieke, Schwindt, and Jagedeesh. The quality of these appointments was reaffirmed, in part, by the fact that three of them (Zagotta, Shadlen, and Rieke) were later chosen as Howard Hughes investigators.

Space was still an issue, especially since the cellular and molecular research studies carried out by the new faculty needed more lab space than had been provided for departing faculty. The I wing, built in 1965 to house the federally-funded Regional Primate and cancer research centers, only provided new space for the new Primate Center investigators. The rest of the PBio faculty remained without additional space. The next space expansion, the J wing, housed the departments of Biochemistry and Genetics, again leaving PBio with no space expansion.

In 1995 the K wing provided the first addition of research space for the School of Medicine since 1964. Michael Whitcomb, MD, the dean at the time, realized that state funds were less and less available for capital construction, and he convinced the University to finance this construction with bonds that would be paid back from the indirect costs that the University gets from grants, some of which had usually been returned to departments. (These bonds are still being paid off in 2010.) A major gift from Bill Gates was awarded to support the development of a new department of Molecular Biotechnology, chaired by Lee Hood, PhD, and the Howard Hughes foundation funded space renovations for its investigators, primarily in Biochemistry and pharmacology, at the top of the J and K wings. None of the K wing was available to P Bio until Bill Zagotta’s appointment as a Hughes investigator. K wing space then became available for the Zagotta and Bothwell labs. When Shadlen was appointed a Hughes investigator, the Hughes foundation remodeled space on the 7th floor of the Primate Center.

Although the additional space associated with Hughes investigators was generally a positive thing, it spatially split the full-time PBio faculty for the first time. Except for labs at the VA hospital for Wayne Crill when he was chief of Neurology there, the move of the Zagotta and Bothwell labs to the K wing was the first time that primary PBio faculty were located outside of contiguous space in the G, H, and I (Primate Center) wings. As the School of Medicine developed new research space at Harborview and South Lake Union, a bigger split occurred. Mark Bothwell moved his lab to South Lake Union.

Implementation of new medical and graduate curricula

Considerable time and effort continued to be spent throughout Dr. Crill’s tenure to develop the curricula for medical and graduate students. At the time Crill stepped down as chair (1999), P Bio faculty offered a series of courses, from which graduate students could choose to cover physiology in a more flexible manner.

Nationally, many basic science departments were rearranged and renamed during this period, and departments and/or degree programs of Neuroscience, Cell Biology, etc became common. At UW, it was realized that some changes needed to be made, but the collaborative style of the faculty here was reflected in efforts to make changes that would include appropriate faculty across campus, but did not leave some faculty with no graduate student programs in which they could participate. To continue to be competitive for the best graduate students, two new interdepartmental degree programs were developed, both run by directors appointed by the graduate school.

Neurobiology and Behavior (NeuB)

Development of the Neurobiology and Behavior graduate program (NeuB) was done in the most expedient manner: expansion of what had been the Physiology/Psychology Joint Program (the Joint program) into a degree-granting neuroscience program with expanded faculty from upper campus (psychology, zoology, computer sciences, engineering, etc) and several basic science and clinical departments of the School of Medicine (PBio, Pharmacology, Biochemistry, and Biological Structure. This change could be done quickly because it did not require that the graduate school approve a new degree-granting authority. The initial directors were Neil Nathanson (Pharmacology) and Albert Fuchs (Physiology and Biophysics).

The Molecular Biology and Cellular Biology degree program (MCB)

The biochemistry department took the lead in developing a multidepartmental degree program that focused on cell and molecular biology. Bertil Hille represented PBio in organizing the new program, which was led by Milton Gordon from Biochemistry.

While the applicants to the multidepartmental programs in Neurobiology and Behavior and in Molecular and Cellular Biology have continued to be strong, applicants to the departmental programs have dwindled. This year (2010), for the first time, Physiology and Biophysics is not actively recruiting for a PBio graduate program. The faculty are primarily focusing on enhancing the biophysics portion of the PBio graduate program.

The Current Era: Strengthening targeted areas, remodeling ancient space

Stanley Froehner, PhD was appointed as the fourth Chair of Physiology and Biophysics in 2000. He came from the University of North Carolina, where he had also chaired the Department of Physiology. Froehner’s PhD was in biochemistry and neurophysiology from CalTech, and he brought biochemical and genetic tools to the study of function and dysfunction (muscular dystrophy) at the neuromuscular synapse.

Dr. Froehner came to Seattle at a time when several faculty had retired, and because Dr. Crill had done a good job of protecting those positions, they were available to fill as a part of Dr. Froehner’s “dowry”. Eight regular faculty (Jane Sullivan, PhD, Fernando Santana, PhD, Sharona Gordon, PhD, Adrienne Fairhall, PhD, Charles [Chip] Asbury, PhD, Andres Barria, PhD, Greg Horwitz, PhD, and Qinghang [Chris] Liu, PhD) were appointed during the first ten years of Dr. Froehner’s chairmanship. It is of interest that three of the eight were women, and this was not completely coincidental. Dr. Froehner recognized that there were outstanding women in the field, and he made a special effort to ensure that they were included as candidates in searches. The three women hired during Dr. Froehner’s first decade as chair (Sullivan, Gordon, and Fairhall) equaled in number the three (Kennedy, Wordeman, and Jagadeesh) who had been appointed to the “regular” faculty during its first 55 years! (This does not include Drs. Skahen and Landau, who had joint appointments with Biological Structure and were hired to teach courses for nursing and other allied health curricula.)

The appointments during Froehner’s era also have brought new techniques and approaches to the department. Dr. Fairhall, for example, is a theoretical mathematician, and she brings strengths in computational neurosciences (Cheap startup lab costs: Good computers and/or pencils!) Dr. Santana brings an interest in membrane channels in the heart, bridging the departmental strengths in membrane physiology with cardiovascular function. Dr. Asbury brings state-of-the-art optical trapping techniques to studies of molecular motor proteins and the nanometer movements that they produce. (And don’t you think that he cares where they place the new campus light rail station!)

Space enhancement

Although space was vacated by retiring faculty, much of it, especially the H wing, was in poor condition. (Dr. Crill, in fact, had taken some of the worst for his lab when he assumed the chairmanship.) Earlier attempts to get outside funds to help fund remodeling of the H wing had been unsuccessful, but Dr. Froehner managed to succeed. The $3.75 million grant that he obtained from the National Center for Research Resources (NCRR) was matched by $4.5 million from the UW. At this time, the School of Medicine had begun to build a second campus at South Lake Union, and that freed up “back fill” space in the main campus building. As a result, Dr. Froehner negotiated an arrangement with Dean Ramsey to add the 3rd floor of the G and H wings to the department’s space. For the first time, the department offices and labs on the main campus were located on three contiguous floors.

In addition, the renovation (which took 5 years from grant submission to final completion) presented a unique opportunity to rearrange the department in a way that made sense scientifically. For example, a biophysics group (Hille, Rieke, Detwiler, Santana and S. Gordon) now occupy contiguous labs. The renovation also included a new 85 seat conference room, a break room and several shared facilities. The ugly black corridor ceilings that resembled an old dirty factory were replaced with brighter light colors and new lighting. Although the renovation was very disruptive (everyone had to move out of their labs for 16 months), the final result was very satisfying. And for the first time since he arrived at UW, Bertil Hille, recipient of the most prestigious awards given to PBio faculty, had a decent lab. (This seems to carry on a UW tradition. The “new” Physics and Astronomy building was built only after Dr. Hans Dehmelt became the University’s first Nobel laureate.)

Major research discoveries and innovations

The preceding semi-sequential description of the development of the Department of Physiology and Biophysics gives a flavor of the “who”, “when”, and a little “what” and “why” of the department. It does little, however, to highlight the major research innovations and discoveries of its faculty and trainees. I have listed a few examples below. You are welcome to send other examples that you think illustrate the scholarly accomplishments of our faculty and trainees.

A few examples include:

Patton and Woodbury– Early intracellular recordings from neurons in the spinal cord. They emphasized three different neural elements: primary afferents with tonic discharge, interneurons, (repetitive firing in response to stimulation of dorsal roots) and motoneurons, with large postsynapticpotentials and fixed antidromic latency of action potentials produced by stimulating the ventral root. The early data were presented concurrently with Eccles’ early intracellular spinal data.

Rushmer, VanCitters, and colleagues– Implanted transducers and associated electronics (including telemetry) in dogs and baboons that allowed measurement of blood flow and pressure in awake, moving animals). These led to non-invasive measurements using ultrasound that have had widespread clinical application.

Luschei and Fuchs– Developed head-mounted apparatus and r-theta adapter to allow an electrode to be introduced to stereotaxic coordinates for recording from deep brain structures in awake, behaving monkeys. This was developed in parallel with the development of a similar apparatus by Evarts and colleagues at NIH and by Mountcastle and colleagues at Johns Hopkins that gave access to cerebral cortex. Luschei’s instrument has been used by many investigators to study brainstem, cerebral, cerebellar, and basal ganglia cells associated with eye, limb, or jaw movement.

Fetz and Finnochio– Showed early (197?) that monkeys can be trained, by operant techniques, to make individual neurons fire, even if their functional neighbors are not. Chet Moritz, Andy Coombs, and Eb Fetz have made significant steps toward putting these techniques to practical application by implanting electrodes for long term recording and behaviorally shaping the neuron’s activity to what would be necessary to drive motoneurons downstream from a lesion to fire in a way that produces a desired movement. It is the kind of “bypass” that could potentially help an individual with spinal cord injury or stroke.

Crill and Schwindt– using a two-electrode voltage clamp, discovered the existence of a persistent inward current in vertebrate neocortical neurons that enables repetitive firing.


This history emphasizes the early years of the PBio department at the University of Washington, when it developed into one of the strongest such departments in the country. I have attempted to give some of the atmosphere in which graduate students were trained, one that emphasized collaboration and encouraged the use of techniques such as the direct use of early computers to acquire and analyze data. Nevertheless, some might argue that a more competitive atmosphere could have led to a consistently higher level of new scientific findings—none of our faculty or former trainees have won the Nobel Prize —yet.

But our former trainees have gone on to provide faculty in many of the top institutions of medical research, they are chairing departments, leading well-funded research programs, and at least one (William Calvin) has spent his career writing books that bring science to the lay public.

TC Ruch’s early insistence that “the development of biophysics within a physiology department will clearly place that department at the front of the modern trend” has proven to be true, and it is of special interest that as departments that carry the name “Physiology” disappear, it is the biophysics component of the department that still makes the department stand out from the neuroscience and cell biology titles commonly labelling departments in biomedical research institutions.


I want to acknowledge the contributions to this effort that are listed below. Any errors or omissions are mine, however. Readers are welcome to send corrections to me for correction of the record.

Marjorie E. Anderson, PhD
Seattle, October, 2010



Scher, Allen. A history of the University of Washington School of Medicine, 
Medical Alumni Association, p 1990.
The initial basis for this summary was the 1987 PBio history written by Dr. Allen Scher for inclusion in Finch, Clement Fulfilling the Dream. A history of the University of Washington School of Medicine, Medical Alumni Association, Pg 59-61, 1990.

Catterall, W. Personal communications.

Crill, Wayne E. Personal communications.

Hille, Bertil. Many personal communications.

Patton, Barbara. Personal communications.

Patton, HD. Annual reports to the Dean, 1964-1973. P Bio departmental records.

Ruch, TC. 
Annual reports to the Dean, 1946-1963. P Bio departmental records.

Woodbury, JW. My life in PBio was exhilaratingAppendix III in this volume 1950 – 1972.