
ALEXANDER FORBES, M.D. 1882 - 1965
The
scientific career of Alexander Forbes spanned fifty-six years of active
research and saw the development of the realm of neurophysiology to
which he contributed over 100 publications.
As
a proper Bostonian of his day he could scarcely avoid going to
Harvard. He graduated in 1905. He also received a Master's
Degree in Biology from that institution in 1906 before undertaking a
degree at Harvard Medical School. By the time he received his
M.D. in 1910 he had been infected with the lure of the research
laboratory while he learned the rudiments of electrophysiology with
Professor G. H. Parker. In his fourth year of studies he was
challenged by application of these techniques to problems in inhibition
in the central nervous system reflex pathways under the guidance of
Professor Walter B. Cannon.
After
Forbes graduated Dr. Cannon immediately offered him the position of
Instructor in Physiology, but further suggested he go to Liverpool to
work with Sir Charles Sherrington for two years. In England
Forbes also spent a short time with Keith Lucas and E. D. Adrian in
Cambridge while they were pioneering the biophysics of peripheral nerve.
Forbes
returned to Harvard Medical School "armed with a Sherrington Guillotine
and a Lucas pendulum" (1) and a mind prepared to attempt the process of
grafting the precise biophysical techniques of peripheral nerve studies
to the relatively complex reflexes studies by the Sherrington
school. With Alan Gregg he proceeded to develop electrical
recordings of the myographic results of flexor reflexes. The two papers
which resulted demonstrated electrical recordings of central reflex
phenomena for the first time. (2) The tool of the time was
the string galvanometer. Forbes had one of the first in New England.
However,
the spread of World War I now began to modify everyone's plans.
Forbes who had grown up in a seafaring family was an expert in
off-shore navigation. He enlisted in the Navy. To his
surprise instead of going to sea he was assigned to work with a firm
called Submarine Signal Corp. (later to become Raytheon) where the
priority was development of submarine detection devices, the classical
"pingers" used on submarine chasers.
Postwar
in 1918 Forbes utilized his electrical experience to develop vacuum
tube amplification for the string galvanometer. The result was a
fifty fold increase in sensitivity (to 20 uV/cm at 200 Hz).
This was a remarkable breakthrough! He could now display
spike potentials of propagated impulses with significant detail. The
resulting paper published with Thacher in 1920 pioneered amplification
and recording techniques and provided reliable observations which
Forbes immediately put to use studying the phenomena of reflex
action. (3)
Another
season in England in 1921 working with Adrian led Forbes and Adrian to
write a paper which demonstrated definitively that CNS motor units
followed the all-or-nothing principle. (4) Forbes followed that
the next year with a magnificent review article which established him
as master of the new developments in neurophysiology. (5) To
understand the significance of being able to state that the CNS is made
of nerve fibers that have the same properties as peripheral nerve
fibers one must remember that the leading theory of inhibition at that
time, the McDougall drainage theory of inhibition, postulated that "..
the CNS had channels in which nervous energy moved just like water in a
pipe with many branches, so that if some branches were open there would
be drainage from a reservoir of 'free nervous energy' with a consequent
'inhibition' of flow along other channels." (6) Forbes further
suggested that understanding mechanisms of transmission of nerve to
muscle would shed light on the function of central synapses. On
the strength of that publication he was promoted from Instructor
directly to Associate Professor. He would become full Professor in 1936.
In
subsequent years Forbes went on to develop the idea that reverberating
circuits and delay paths explain to a considerable extent prolonged
excitatory responses. He also had the correct idea on muscle tonus,
postulating that prolonged contraction was due to motor neurons firing
at slow frequencies. These ideas were later verified by on-going
studies by Denny-Brown, Adrian and Bronk when better techniques became
available. Another classic paper was published in 1940 with
Renshaw and Morison on the analysis of field potentials in the CNS. (7)
Although
physiology was his mainstream, Dr. Forbes was also accomplished at
sailing, figure skating, skiing and riding. He piloted his own
airplane and grumbled good naturedly when forced to relinquish night
flying certification after his seventieth birthday. This latter
avocation had led him to spend summers in the mid-thirties perfecting
the techniques of taking oblique photograms from the cockpit of a small
airplane while he mapped the uncharted coast of Labrador. As
World War II approached, although close to 60 years of age, he again
enlisted in the Navy to avail the military of his wealth of
geographical knowledge of the Northern Atlantic coast. His
expertise was vital to the selection of sites for airstrips in Labrador
expediting the transfer of fighter planes from North America to Europe.
When
Dr. Forbes became Emeritus Professor in 1948 he moved from the Harvard
Medical School to the Harvard Biological Laboratories in Cambridge,
Massachusetts. There he and George Wald pursued studies on the
comparative physiology of color vision in laboratory space made
available by Dr. John Welsh. Their work helped to establish that
three separate pigments in the retina of the eye are involved in
perception of color.
Alexander
Forbes died in 1965 at the age of eighty-three. Dr. Forbes is
remembered as a man of enormous accomplishment, and a rare zest and
enthusiasm for life. As one of the founding Trustees of the Grass
Foundation he endowed that organization with 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. Because he was a life-long neighbor of MBL and
collaborator and friend of many of the early MBL community, the Grass
Foundation has named these two annual lectures as an on-going memorial.
July. 1987
1)
Alexander Forbes Memorial, Harvard University Gazette, Vol.
LXI, No. 5 October 16, (1965).
2)
A. Forbes, and A. Gregg, American J. Physiol., 37:
118, (1915) Ibid, 30: 172, (1915).
3) A. Forbes and C. Thacher, American J. Physiol., 52: 409, (1920).
4) E. D. Adrian and A. Forbes, J. Physiol., 456: 301, (1922).
5) A. Forbes, Physiol. Rev., 2: 361, (1922).
6) J. C. Eccles, Perspectives in Biology and Medicine. 13, No. 3, Spring, (1970).
7) B. Renshaw, A. Forbes, R. B. Morison, J. Neurophysiol., 3: 73, (1946).