Historical Roots

The history of science is often discussed in terms of the

scientists who have had a major influence on it. The following
material, which deals with the historical roots of biopsychology,
follows that approach.

William James (Figure 1.2.1). Undoubtedly one of the greatest

minds to have written about psychology, William James pioneered
modern scientific psychology in the United States. He simply
assumed that a knowledge of the physiology and anatomy of the
brain were essential for a full understanding of the mind. In
fact at the very end of Chapter 1 of his great two volume work
"The Principles of Psychology" (2) there is a footnote to the
reader:

Nothing is easier than to familiarize one's self with the

mammalian brain. Get a sheep's head, a small saw, chisel, scalpel
and forceps (all three can best be had from a surgical instrument
maker), and unravel its parts either by the aid of a human
dissecting book, such as "Holden's Manual of Anatomy", or by the
specific directions ad hoc given in such books as Foster and
Langley's "Practical Physiology" (MacMillan) or Morell's
"Comparative Anatomy and Dissection of Mammalia" (Longmans).

Chapters two and three of volume 1 are entitled "Functions of the

Brain" and "General Conditions of Brain-activity". These chapters
provide an elegant summary of what was known about neuroanatomy
and neurophysiology at the time. In addition, many crucial issues
for the field that was to become biopsychology were considered
analytically. It should be a self-imposed requirement for all
biopsychologists to read and reread these two chapters so that
they remain constantly aware of their intellectual debt to this
great thinker.

Ivanovich Pavlov (Figure 1.2.2). Although he always considered

himself a physiologist (he received the Nobel prize for
Physiology or Medicine in 1904), Pavlov made seminal
contributions both to behaviorism and biopsychology. Ironically,
these two approaches to psychology are often seen as antithetical
today. When Pavlov's famous book on conditioned reflexes was
published in English in 1927 (1), it was a major event in the
history of biopsychology. An immediate success, this collection
of Pavlov's lectures was read avidly by scientists in a broad
range of disciplines and thus had a wide sphere of influence.
Much of its popularity among psychologists is attributable to the
proselytizing of J. B. Watson who saw in the conditioned reflex
the basic building block of learned behavior. Indeed, Watsonian
"behaviorism" dominated the field of psychology for many years.
Pavlov's treatment of localization of function in the cerebral
neocortex is still influential in various guises today. If
William James was the first great theorist in biopsychology, then
Pavlov was the first great experimentalist.
Santiago Ramon y Cajal (Figure 1.2.3). The neuron doctrine, the
notion that neurons are individual, separate cells which
communicate with each other across tiny spaces is based on the
brilliant work of the great neuroanatomist, Santiago Ramon y
Cajal, who shared the Nobel prize in Physiology or Medicine with
Camillo Golgi in 1906. Cajal, ironically, used Golgi's staining
method to demonstrate that neurons are individual units
representing the fundamental structure of the nervous system.
Golgi, on the other hand, believed that the nervous system was a
syncitium and that neurons were in physical contact with each
other. These two volatile personalities almost came to blows at
the Nobel presentation ceremony, so strongly did they believe in
their own theories. Time has proven Cajal to be correct, though
modern histological techniques have demonstrated that in certain
classes of synapses direct contact is made between neurons (see
Unit II).

Sir Charles Sherrington (Figure 1.2.4). Much of the work that

Sherrington did would be classified as biopsychology today,
though he is generally classified as a neurophysiologist. His
methods consisted primarily of analyzing defensive flexion
reflexes in the spinal cords of dogs. His brilliant reasoning
about the results of the studies and scientific insights into the
nature of information processing in the spinal cord led him to a
correct interpretation of the neuron doctrine. His introduction
of the concept of "synapse" (to clasp) as the site of interaction
between neurons remains one of the great achievements of modern
neuroscience. Sherrington was awarded the Nobel Prize in
Physiology or Medicine in 1932.

Karl Lashley (Figure 1.2.5). One of the early proponents of the

American school of holism in brain function, Karl Lashley was
strongly influenced in his early career by Shepherd Franz. Franz
had been fighting against the ideas of compartmentalization of
associative and intellectual functions for the first two decades
of the twentieth century. Lashley was a biopsychologist and an
experimentalist. His investigations with Franz revealed that rats
previously trained on a brightness discrimination task showed
little or no impairment on this task after large portions of
their frontal lobes were destroyed.

Many of his contemporaries believed that the frontal lobes were

essential for normal learning and intellectual functioning. The
"engram", the physical trace, i.e., the "memory trace", which
remained in the brain from learning, was thought to reside in
them. Implied in these assumptions was, of course, the notion of
anatomical specificity for various higher order intellectual
functions.

Lashley's theoretical position allowed for some anatomical

localization of function for sensory and motor processes but none
for intellectual ones. This de-emphasis of localization of
function led to his formulation of the principles of
"equipotentiality," and "mass action," which hold that no single
part of the cortex is more specialized for learning than are
other parts. In the years that followed Lashley's work however,
the localization of function hypothesis has gained increasing
popularity. This view of brain function currently dominates the
experimental and theoretical approaches to neuronal organization
within biopsychology.

Donald Hebb (Figure 1.2.6). Pavlov's theory about conditioned

reflexes in learning and Sherrington's studies of synaptic
activity led Donald Hebb, a student of Lashley's, to speculate
about the changes that must take place at the synapse when a
learned association is formed. The essence of his thinking was
that when one neuron repeatedly takes part in activating another
neuron, some growth process or change in metabolism occurs in one
or both cells that increases the ease with which the first neuron
can fire the second (Figure 10.2.5). In a sense, Hebb's work
created a synthesis of the theories of Pavlov, Sherrington and,
to a certain extent, Lashley, and laid the foundation for much of
the work on learning and memory to the present day.

The "Hebbian Synapse" has become the cornerstone of many

biopsychological theories of associative learning. It is usually
invoked to explain the increases in synaptic efficacy that occur
when presynaptic neurons are simultaneously active. This
mechanism is also widely used in connectionist neuronal net
models which often involve computer simulations. The success of
Hebb's model attests to its generality and to its foundation in
empirical evidence, which of course add to its authenticity as a
viable biological mechanism for learning and memory.

Roger W. Sperry (Figure 1.2.7). Specificity of neuronal

connections and their role in regeneration, the analysis of
hemispheric function in 'split brain' patients, and the concept
of the mind as an emergent property of the brain represent three
major contributions of Roger Sperry, who shared the Nobel Prize
in Physiology or Medicine in 1981 with Hubel and Wiesel (see
below). Sperry's work with split brain patients (Figure 11.1.6),
people whose 2 brain halves had been surgically separated in an
attempt to control epileptic seizures, revealed many
specializations of the two hemispheres. Most of his early studies
in this area were conducted with sensory inputs restricted to one
side or the other of the brain. When the 2 hemispheres are
disconnected, communication between them is, of course,
drastically reduced. This hemispheric isolation provided an
important means for Sperry and others to elucidate individual
functions of each brain half.

How developing axons find their way to the correct target in the
brain was another of Sperry's major interests, and he did much of
the definitive research in this area. In amphibians, unlike
mammals, a damaged optic nerve grows back and contacts the
tectum, the major visual area in fish, reptiles, amphibians, and
birds. Sperry found that when the new connections formed, the
animals regained normal vision. In a series of ingenious studies,
Sperry and his students discovered that the regenerating axons
must be following a chemical trail to their "correct" targets (
Figure 4.2.2). Sperry was also very interested in the philosophy
of science and made major contributions to our understanding of
the philosophical issues in biopsychology.

"When subjective values have objective consequences ... they

become part of the content of science ... Science would become
the final determinant of what is right and true, the best source
and authority available in the human brain for finding ultimate
axioms and guideline beliefs to live by, and for reaching an
intimate understanding and rapport with the forces that control
the universe and created man." (5)

David Hubel and Torsten Wiesel (Figure 1.2.8). Hubel and Wiesel
shared the 1981 Nobel prize with Sperry, as a result of their
pioneering work on how the primary visual cortex processes
shapes. They found that most neurons in the visual cortex of
mammals have binocular receptive fields. In other words, the
cells respond to portions of both eyes. In addition, they
demonstrated that these receptive fields tended to be shaped like
a bar or an edge. Such cells may therefore serve as feature
detectors when objects are being viewed. These feature detectors
indicate the presence of a particular feature of an object. There
are three primary categories of cellular feature detectors in the
visual cortex: simple, complex and hypercomplex (Figure 6.2.6) (
Figure 6.2.7) (Figure 6.2.8) (Figure 6.2.9).

Hubel and Wiesel assumed that complex pattern perceptions were

formed from groups of feature detectors acting cooperatively.
This of course raises the important issue of what it is in the
brain that organizes this cooperativity. How does the visual
system use such cell complexes to distinguish your grandmother's
face from that of your psychology professor? At this point we do
not have an answer to that question. In fact, the importance of
the feature detector concept has recently been challenged.
Continuing efforts to investigate perception in the visual system
will, hopefully, provide answers to this riddle.

Rita Levi-Montalcini (Figure 1.2.9).

Rita Levi-Montalcini and Stanley Cohen received the Nobel Prize

in Physiology or Medicine in 1986 for their discovery of Nerve
Growth Factor (NGF). This substance significantly affects the
growth of neurons in the ganglia of the sympathetic nervous
system and dorsal root ganglia. They demonstrated that
administration of NGF to a chicken embryo resulted in the
formation of sympathetic ganglia with many more cells than
normal. The cells were also larger than normal, and showed a
marked proliferation of axonal processes radiating out in all
directions. NGF was first found in a variety of places, including
salivary glands of mice, snake venom, and skin tumors. Later work
showed that it is normally produced by a number of target organs
during development. It is taken up by sympathetic axons which
innervate these organs, and transported back to the cell body,
promoting the continued growth and survival of that cell. When an
axon fails to receive enough NGF it degenerates, and its cell
body dies. NGF was the first identified neurotrophic factor. Many
such factors have since been identified, but it was the work and
insight of Dr. Levi-Montalcini which opened the door to these
findings. More recently, Dr. Levi-Montalcini, inspired by the
writings of Roger Sperry, founded the International Council for
Human Duties, based at the University of Trieste, Italy. This
group, consisting of people from widely based disciplines in the
sciences and humanities, developed a document called "The
Declaration of Human Duties" which is designed as a corollary to
the United Nations "Declaration of Human Rights". Her untiring
efforts as both a scientist and humanitarian rank her as one of
the most unusual and outstanding individuals of the 20th century.