The Origins of The Grass Foundation

In the fall of 1935, Albert M. Grass and Ellen H. Robinson both came to the Department of Physiology at Harvard Medical School (HMS). This entirely fortuitous confluence of their lives led to their marriage, to a commercial endeavor—the Grass Instrument Company—that would provide equipment of high quality to neuroscientists and other physiologists for over half a century, and finally to the formation of The Grass Foundation, which has benefited the neuroscience community since 1955. The Department of Physiology at Harvard—the seedbed for these accomplishments—had a deep-rooted commitment to providing both financial and moral support to scientists who were at the beginning of their careers. Albert and Ellen clearly benefited from this commitment, for it generated interactions and collaborations that led to and facilitated the success of the Grass Instrument Company and then the Foundation. Thus, the origins of The Grass Foundation must be sought, not only in the conjoined histories and proclivities of Albert M. and Ellen R. Grass, but also in scientific and educational developments that took place in the HMS Department of Physiology between 1906 and 1935, well before Albert and Ellen met there. This essay is an attempt to dissect those tangled threads; it ends with a discussion of The Grass Foundation’s hallmark program—the Grass Fellowship Program at the Marine Biological Laboratory in Woods Hole, Massachusetts—and the impact that this program has had on neuroscience.


Introduction
In the fall of 1935, Albert M. Grass and Ellen H. Robinson both came to the Department of Physiology at Harvard Medical School (HMS). This entirely fortuitous confluence of their lives led to their marriage, to a commercial endeavor-the Grass Instrument Company-that would provide equipment of high quality to neuroscientists and other physiologists for over half a century, and finally to the formation of The Grass Foundation, which has benefited the neuroscience community since 1955.
The Department of Physiology at Harvard-the seedbed for these accomplishments-had a deep-rooted commitment to providing both financial and moral support to scientists who were at the beginning of their careers. Albert and Ellen clearly benefited from this commitment, for it generated interactions and collaborations that led to and facilitated the success of the Grass Instrument Company and then the Foundation.
Thus, the origins of The Grass Foundation must be sought, not only in the conjoined histories and proclivities of Albert M. and Ellen R. Grass, but also in scientific and educational developments that took place in the HMS Department of Physiology between 1906 and 1935, well before Albert and Ellen met there. This essay is an attempt to dissect those tangled threads; it ends with a discussion of The Grass Foundation's hallmark program-the Grass Fellowship Program at the Marine Biological Laboratory in Woods Hole, Massachusetts-and the impact that this program has had on neuroscience.

Albert M. Grass and Ellen H. Robinson
Albert Melvin Grass (1910Grass ( -1992 was born in Quincy, Massachusetts, on September 3, 1910, to Henry J. Grass and Bertha (Martin) Grass. After graduating from Quincy High School, he funded his college education by working at Samson Electric Company testing and installing the amplifiers and systems that provided sound for films (Marshall, 1980;Henry, 1992). He was successful, both academically and financially, and the B.Sc. degree in electrical engineering was duly awarded by MIT in 1934. Albert remained at MIT to work on servo-mechanisms used to simulate earthquakes in the study of strains acting on model water towers and other structures (Marshall, 1980).
In May of 1935, Frederic A. Gibbs-a research fellow in neurology at the HMS Department of Physiology-contracted with Albert Grass to build a 3-channel electroencephalograph (EEG). Albert and his brother Everett worked in the basement of their father's house and finished the project early in the fall of 1935 (Marshall, 1980;Grass, 1984).
This accomplishment, completed in only three months, led to Albert's being hired as a part-time Research Instrument Engineer by the HMS Department of Physiology, and he remained in that position from 1935 until 1943. As part of this job, he continued to improve the EEG (Grass Instrument Co., 1971;Grass, 1984) and its applications (Grass and Gibbs, 1938). In addition, he worked closely with scientists, tailoring equipment to their needs (e.g., square wave stimulators and amplifiers; Forbes and Grass, 1937;Marshall, 1980). Ellen Harriet Robinson (1914-2001 was born in Taun On Upton's recommendation, Ellen decided to immerse herself in "a broader field of brain function" (Marshall, 1995) and began graduate work in the Department of Physiology of HMS in the fall of 1935 (Marshall, 1980(Marshall, , 1995. A special arrangement was made with Radcliffe College so that she could take a course in "Research" (Marshall, 1995), and she was supported by a Porter Fellowship in Physiology (1935)(1936) from the American Physiological Society (Howell and Greene, 1938;Fenn, 1963). Alfred C. Redfield (1890Redfield ( -1983 was her initial sponsor on a project entitled "Auditory Action Potentials" (Fenn, 1963;Brobeck et al., 1987), but her work was ultimately conducted under the sponsorship of Hallowell Davis Fenn, 1963). Ellen collaborated with a number of scientists, including Arthur James "Bill" Derbyshire (Grass, 1980), Edward H. Kemp, and Georges Coppée, a Fellow of the C.R.B. Educational Foundation, Institute de Physiologie, Liège. Her studies of the responses of the brainstem to auditory stimulation led to three publications Kemp et al., 1936.
Ellen Robinson and Albert Grass met in the fall of 1935 and were married on June 28, 1936. Ellen continued as a Ph.D. candidate in the Department of Biology at Harvard University, recording from the auditory cortex of rabbit in the laboratory of the noted physiological psychologist Karl Spencer Lashley (1890 -1958). Soon, however, she decided to devote herself "to motherhood and doing whatever I could to help Albert provide equipment to a growing number of scientists" (Marshall, 1995).

The Development of Electroencephalography at the HMS Department of Physiology and the Formation of the Grass Instrument Company
The report of voltage changes recorded through the cranium of humans by Hans Berger (electroencephalogram, EEG;Berger, 1929) and his observations of EEG variations in patients with epilepsy (Berger, 1932;Gibbs et al., 1936) were the basis for continued EEG studies of epilepsy in the United States.
At the time, Stanley Cobb (1887-1968 and William G. Lennox (1884Lennox ( -1960 were experts on epilepsy (e.g., Cobb, 1922;Lennox and Cobb, 1928;Lennox, 1936;see White, 1984, for Cobb's complete bibliography) carrying out their investigations at Harvard Medical School (Hughes and Stone, 1990). In 1929, Cobb offered Frederic A. Gibbs (1903Gibbs ( -1992, who had just received his M.D. from Johns Hopkins, a fellowship in neuropathology to work on epilepsy. Gibbs worked in the Lennox lab where he met Erna Leonhardt; the couple were married in 1930 and were collaborators thereafter (Hughes and Stone, 1990).
The Gibbses wanted to record EEGs from epileptic patients, and Hallowell Davis' engineer, E. Lovett Garceau (1906-1966, had built amplifiers and a portable EEG that could be used for such recordings Davis, 1934, 1935;Garceau and Forbes, 1934). Encouraged by Cobb, the Gibbses approached Davis to be a collaborator, and Davis enthusiastically endorsed their plan (White, 1984). With the departure of the engineer, Garceau, and the need for EEG machines with more than one channel, Frederic Gibbs sought out advice at the Massachusetts Institute of Technology. There he met Albert Grass (Marshall, 1980;Grass, 1984). With funding from the Macy Foundation, he contracted with Albert Grass in May of 1935 to "build three channels of EEG amplifiers to drive the Western Union Morse Code inkwriting undulator" (Grass, 1984;Hughes and Stone, 1990).
The Gibbses went off for the summer to attend the International Congress of Physiologists in Leningrad and Moscow, and to visit Berger and engineer J. F. Tönnies in Germany. In August of 1935, rather late in the summer, Frederic Gibbs mailed a sketch of Tönnies' neuropolygraph (designed for A. Kornmüller's animal studies) to Albert Grass (Grass, 1984;Hughes and Stone, 1990). By that time, the EEG machine being constructed by Albert and Everett Grass in Quincy must have been well along, for when the Gibbses returned in the fall, it was finished, as mentioned above. This EEG was used by Lennox, Frederic and Erna Gibbs, and Davis in their pioneering investigations, which demonstrated the power of the EEG in the diagnosis of epilepsy (Gibbs et al., 1935(Gibbs et al., , 1936(Gibbs et al., , 1937Brazier, 1968).
The demand for EEG machines increased markedly during the 1940s. To meet this demand, Albert began to manufacture commercial instruments (Marshall, 1987). Thus, the small business that started in a basement in 1935 continued as a partnership between Albert and Ellen Grass, and ultimately developed and grew to become, 10 years later, the Grass Instrument Company. The success of the Company was due to the balance between Albert's engineering skills and Ellen's scientific expertise, which was critical in the proper design of equipment to meet the needs of neurophysiologists ( Fig. 1). Instruments were designed for "convenience, durability and serviceability" (Morison, 1979).
The Grass Instrument Company was never a typical business. In the early years, employees and many of the customers were warm and loyal friends of Albert and Ellen (Morison, 1979), and neurophysiological equipment was loaned to investigators throughout the world, and especially to Grass Fellows and courses at the Marine Biological Laboratory (MBL). To Albert and Ellen, the Company was always meant to contribute "to the development of human knowledge and the search for basic scientific truth." (Grass Instrument Co., 1967). They took great care to ensure that equipment was being properly used for the benefit of the patient. Ricardo Miledi recalls, "When I was in Mexico, I remember on more than one occasion, seeing a letter [from the Grasses] inquiring about doctors that intended to purchase their most advanced EEGs and other equipment. Albert and Ellen were very concerned that their equipment be used wisely for the benefit of the patients and for research, and not merely to extract money from the patients."

Department of Physiology at Harvard Medical School
Albert and Ellen Grass's success was clearly due, in part, to the support they received from established researchers in the Department of Physiology at Harvard Medical School. Walter B. Cannon and Alexander Forbes were especially critical in this regard.
"Speaking personally now," Ellen once said, "Dr. Cannon made very many things possible for Albert and for me. He invested in us at a time when biomedical engineering was indeed in its infancy, and the role for women in science practically negligible" (Grass, 1970).
Walter B. Cannon (1871Cannon ( -1945 served as the George Higginson Professor of Physiology at Harvard Medical School for 36 years (1906 -1942). For this entire period, he was chairman of the Department of Physiology and an Emeritus for his last three years. He made many important contributions to our understanding of how the human body functions: the use of Roentgen rays to investigate gastrointestinal motility (Cannon, 1898;Cannon, 1911;Brooks et al., 1975;Barger, 1981), the effects of emotions on the functional state of the body (Cannon, 1915;Davenport, 1981), the basis of surgical shock (Cannon, 1923), the constancy of the internal environment or homeostasis (Cannon, 1939), autonomic neuro-effector systems (Cannon and Rosenblueth, 1937), and the effects of denervation on various tissues (Cannon and Rosenblueth, 1949).
Walter Cannon spent his undergraduate years at Harvard University and continued on at Harvard Medical School, where the faculty held that interested medical students should be encouraged to conduct original research. Thus, in the first semester of his medical training Cannon and fellow student Albert Moser were encouraged by Henry P. Bowditch (1840Bowditch ( -1911 to conduct a study on deglutition (Benison et al., 1987). Later, as a third-year medical student, Walter Cannon was approached by William Norton Bullard , a neurologist at Boston City Hospital, who offered to fund further research (Taylor, 1931).
These early research experiences clearly had a profound influence on the development of Cannon's scientific philosophies.
Every man active in investigation has more problems in mind that he can work at himself. A part of his service to the world consists in training others by giving to others these problems to work at under his direction. These "others" are ordinarily his students,-young men who have been stimulated by his example. They are not yet established in life, they require remuneration until they have done enough work to warrant their being taken into independent positions. They should receive during these years of training (which are very likely to be productive of good results in research) sufficient compensation to afford comfortable support (Benison et al., 1987).
Cannon is generally considered to have been exemplary in his scientific conduct and his concern for human welfare (Cannon, 1945;Forbes, 1945;Morison, 1945;Davis, 1975). He "saw that the freedom and beneficence of science could be guaranteed only within the framework of a just society, national and international" (Grass, 1970), and was committed to providing promising young scientists, independent of nationality, the opportunity to participate and contribute to the advancement of science (Morison, 1945).
One such scientist was Arturo Rosenblueth (1900 -1970) 1 who came to Harvard from Mexico as a Guggenheim Fellow from 1930 to 1932 to work with Cannon. He quickly became Cannon's "favorite son" and secured a position in the Department of Physiology. Their collaborations continued for the next 14 years (e.g., Rosenblueth, 1937, 1949).
Cannon and Rosenblueth mentored several scientists who would ultimately become founding and early trustees of The 1 The names of the founding and early trustees of the Grass Foundation are printed in bold type in this section of the paper. E-mail: Steven.J.Zottoli@Williams.edu. Grass Foundation (Table 1). Alexander Forbes (1882-1965) as a fourth-year medical student was encouraged by Cannon to become involved in research. After receiving his M.D. degree in 1910 from Harvard Medical School, Forbes studied with C. S. Sherrington (1857Sherrington ( -1952 for two years, and briefly with Lucas in 1912; afterwards, he returned to Harvard and the Department of Physiology (Fenn, 1969, Davis, 1970Eccles, 1970). Forbes added a strong engineering background to the department and was continuously at the forefront of technological advances that he applied to neurophysiological investigations. These included the use of the vacuum tube amplifier in conjunction with a string galvanometer to record action currents in nerve and muscle (Forbes and Thacher, 1920, see also Gasser and Newcomer, 1921;Forbes et al., 1931;Grass, 1984;Frank, 1986;Seyfarth, 1996), the study of reflex activity (Forbes, 1922;Davis, 1975;Seyfarth, 1996), and the use of microelectrodes for extracellular recording from cortical cells (Renshaw et al., 1940;Brazier, 1968).
In fact, Forbes' technical and analytical strengths, along with those of Hallowell Davis (Forbes et al., 1931), complemented the more integrative approaches of Cannon and Rosenblueth (Cannon, 1945). Cannon's encouragement of Forbes as a young medical student could not have affected a more appreciative and capable individual. Alexander Forbes quickly adopted the philosophy of encouraging scientists in his own way. He "anonymously supported others in the department of physiology" (Davis, 1970;Seyfarth, 1996).
One of the many young medical students supported by Forbes was Hallowell Davis (1896Davis ( -1992. He received his B.A. degree in 1918 and the M.D. degree in 1922 from Harvard, worked for a year at Cambridge University in England with Edgar D. Adrian, and then returned to Har-vard in 1923 as an Instructor in the Department of Physiology (Davis, 1991;Galambos, 1998). Some of his studies at Harvard include the all-or-none nature of the nerve impulse (Davis et al., 1926), the use of the EEG in the study of epilepsy (Gibbs et al., 1935), recordings from single units in the "auditory nerve" of cats (Galambos and Davis, 1943; the recordings turned out to be from cell bodies of the cochlear nucleus, Davis, 1975), and the tolerance of the human ear to loud sounds (Davis et al., 1950) (Lindsley, 1934(Lindsley, , 1935a and pioneered the use of the electromyogram in neuromuscular disorders (e.g., Lindsley, 1935bLindsley, , 1936see Lindsley, 1995, for a review).
Robert Morison (1906Morison ( -1986) received an undergraduate degree from Harvard in 1930 and the M.D. in 1935. He was encouraged to pursue research by his mentor, Rosenblueth, during his medical school years. "He [Morison] was a man of great and thoughtful learning but one who, above all, wanted to understand the meaning of life and the significance of science for that fundamental issue. He understood what it was to make a moral vocation of one's intellectual work, an effort that requires not only reading, writing, and thinking, but also something else: the living out, in daily life, of the values and virtues that animate that work." (Callahan, 1987). Morison collaborated with many of those present in the Department of Physiology in the 1930s (e.g., Rosenblueth and Morison, 1934;Rosenblueth et al., 1936) and went on to Rockefeller University and then Cornell (Eisner et al., 1986(Eisner et al., -1987.

The Formation of The Grass Foundation
As the number of requests for financial support of neuroscience endeavors grew, Albert and Ellen recognized that a mechanism must be found to evaluate proposals and make decisions (Morison, 1979). The Grass Charity Trust was formed on December 31, 1948, and charitable disbursements were made after June 27, 1951. This Trust donated most of its assets to The Grass Foundation (The Grass Foundation minutes, 1958), which was formed in 1955 "to assist in advancing knowledge principally in the field of neurophysiology, and including allied fields of medicine and science" (Article 2 Section 1 of The Grass Foundation Constitution and Bylaws).
As we have seen, most of the founding and early trustees of the Foundation were, at some time in the 1930s, members of the Department of Physiology at Harvard Medical School ( Fig. 2; Table 1). This is only fitting, because their commitment to the support of young scientists, their own exemplary performance at the bench, and their concern for human welfare (Morison, 1979) reflect the basic principles that have molded The Grass Foundation. The Foundation currently supports programs within the Society for Neuroscience, at the MBL, and at other institutions. The Grass Fellowship Program at the MBL was one of the first and most important projects of The Grass Foundation, and it continues to flourish.

The Association of Albert and Ellen Grass and The Grass Foundation With the Marine Biological Laboratory at Woods Hole
Albert and Ellen Grass's affinity for the MBL developed over many years and is based on several associations. For example, Alexander Forbes had a natural affection for the Woods Hole area. He spent summers on Naushon Island, which is still owned by the Forbes family, and he was a distinguished investigator at the MBL, publishing research done there with Catharine Thacher (Forbes and Thacher, 1925;Forbes, 1933). Albert Grass was undoubtedly at-tracted to the MBL because, as a center of neurophysiology, it was regularly visited in the summers by scientists who were actively involved in the development of new equipment. With this common interest, Albert developed lasting friendships with several MBL scientists, including Harry Grundfest (1904Grundfest ( -1983 and Stephen Kuffler (1913. Finally, Ellen was drawn to the MBL by her passion for the marine environment and the animals that live there. This passion was particularly evident in her Grass Instrument Company Calendars and the "live displays" presented at the annual meetings of the Society for Neuroscience. Of the initiatives in support of basic science at the MBL, the Grass Fellowship Program most closely embodies the philosophy of the founding trustees who had "a love for the adventure of new ideas, a priority for assisting young investigators, and a program focus to direct its resources to the growth of neurophysiology" (Grass, 1987). Beginning with two fellows in 1951 under the auspices of the Grass Charity Trust, more than 400 young neuroscientists have spent summers at the MBL conducting independent research.
Established in 1959, the Forbes Lectureship is an integral part of the Grass Fellowship Program. Each summer, the Trustees of The Grass Foundation bring one of the world's outstanding neuroscientists to the MBL "to honor the outstanding achievements of Dr. Alexander Forbes as a pioneer and major contributor to the field of neurophysiology, who has always been an inspired teacher of young students." (The Grass Foundation minutes, December 11, 1958). The Forbes Lecturer not only presents two lectures as part of the MBL's Lecture Series, but also shares space with the Fellows in the Grass Laboratory. Forbes inaugurated the series with a pair of lectures, on "The Growth of Neurophysiology" and on "Electrophysiology of Color Vision." Many Grass Fellows have gone on to become leaders in neuroscience (  Miledi, 1953Rosenblueth et al., 1954) when Albert Grass and Steve Kuffler came by to visit Rosenblueth. Informed about the Grass Fellowship Program by the visitors, Miledi and Remolina came to the MBL as fellows in 1955 (Fig. 3).
There was no Grass Laboratory or formal program at that time, and fellows generally worked in separate spaces and did not interact with one another a great deal. Steve Kuffler, Harry Grundfest (1904-1983, and their collaborators were perennial investigators at the MBL, and both were Forbes Lecturers, in 1975 and1979, respectively. They acted as mentors for Miledi and other Grass Fellows in the early years of the program (Zottoli, 1990). Albert and Ellen Grass would also visit the MBL periodically to make sure that the Grass Fellows had what they needed in the way of equipment, space, and support.
While at the MBL, Miledi worked on lobster and crayfish stretch receptors and the squid giant axon (Miledi, 1957). Miledi has gone on to publish over 460 scientific articles, and he is one of two neuroscientists to be chosen more than once by the Trustees of The Grass Foundation as the Forbes Lecturer (the other is Theodore H. Bullock, in 1963 and1991). Miledi presented some of his and Bernard Katz's seminal work on neuromuscular transmission at the MBL in 1964 as the sixth Forbes Lecturer. The topic of his two lectures was "Localization of ACh receptors and cholinesterase in muscle fibres." He returned as the Forbes Lecturer in 1990 (Fig. 3) and delivered two lectures on pioneering work that utilized frog oocytes to study native receptors and express exogenous messenger RNA (Kusano et al., 1977(Kusano et al., , 1982Barnard et al., 1982). The subject of his two lectures was "How to study the brain using frog oocytes." Dr. Miledi has served two terms as a Trustee of The Grass Foundation (19921997-2000. He is currently a Distinguished Professor in the Department of Neurobiology and Behavior at the University of California, Irvine. The philanthropic largess of Albert and Ellen goes well beyond the MBL and has benefited the neuroscience community in many other ways. For example, "individuals known to be sound investigators, working under budgetary or foreign exchange difficulties, often found themselves the recipients of indefinite loans" of neurophysiological equipment through the auspices of the Grass Instrument Company (Morison, 1979). Ricardo Miledi remembers, "Joaquin Remolina and I were once asked to make a list of Grass equipment to be bought for the Department of Physiology at the Institute of Cardiology in Mexico. We made a big list that was sent to the Grass Instrument Company and were looking forward, with great excitement, to the day when the equipment would arrive. Then, to our great consternation, there was a big devaluation of the peso and we were asked to send a new order with a good number of items deleted. Later, when the shipment arrived we were all extremely pleased to see that all the items in our original list had arrived and a few extra ones had been included, as if to compensate for our transient worries. I wonder if any other company exists that would do that?" Albert Grass died in Quincy, Massachusetts, on May 29, 1992, and Ellen died 9 years later on June 14, 2001, also in Quincy. The Grass Foundation that embodies their ideals continues to be committed to providing general support for Table 2 Grass Fellows from 1951Fellows from to 1961 Year Fellow "excellent science." The tireless efforts of Albert and Ellen to monitor the pulse and flow of neuroscience have led to initiatives in support of the field, especially in helping those in need or just starting out. The greatest achievements of the Foundation's initiatives, such as the Grass Fellowship Program at the MBL, have resulted from the ability of the Trustees to listen, hear and respond to needs of scientists in a rapidly changing discipline.