Cell Assemblies and Phase Sequences

According to Hebb, the neural interconnections in a newborn’s brain are essentially random. It is
experience that causes this network of neurons to become organized and provide a means of
effectively interacting with the environment. Hebb speculated that every environmental object
we experience fires a complex package of neurons, called a cell assembly. When we look at a
pencil, for example, our attention shifts from the point, to the shaft, to the eraser. Each shift of
attention causes different neurons to fire, and, at first, these neurons fire independently of the
others. Eventually, however, because the neurons stimulated by the presence of a pencil fire
either simultaneously or in close succession, they become a neurological package corresponding
to the experience of a pencil. According to Hebb, it is reverberating neural activity that allows
neurons that were temporarily separated to become associated. For example, the neurons
activated by observing a pencil’s point become associated with the neurons activated by
observing a pencil’s eraser, although the observations do not occur at exactly the same time.
Hebb believed that neural activity caused by stimulation continued for a short time after the
stimulation ceases (reverberating neural activity), thus allowing the development of successive
neural associations. Once a cell assembly exists, it can be fired by internal or external stimulation
or by a combination of the two. When a cell assembly fires, we experience the thought of the
environmental object or event to which the assembly corresponds. For Hebb, the cell assembly
was the neurological basis of a thought or an idea. In this way, Hebb explained why
environmental objects do not need to be present for us to think about them. Just as the various
neurons stimulated by an object become neurologically interrelated to form a cell assembly, so
do cell assemblies become neurologically interrelated to form phase sequences. Hebb (1959)
defined a phase sequence as “a temporally integrated series of assembly activities; it amounts to
one current in the stream of thought”. Like a cell assembly, a phase sequence can be fired by
internal or external stimulation or by a combination of the two; when one or more assemblies in
a phase sequence fire, the entire phase sequence tends to fire. When the entire phase sequence
fires, a stream of thought—a series of ideas arranged in some logical order—is experienced.
According to Hebb, childhood learning involves the slow buildup of cell assemblies and phase
sequences, and this kind of learning can be explained using associationistic terminology. Adult
learning, however, is characterized by insight and creativity and involves the rearrangement of
already existing cell assemblies and phase sequences. Although childhood learning can be
explained in terms of associationistic principles, adult learning is better explained in terms of
Gestalt principles. As we will see in the next chapter, Hebb’s contention that neurons that are
active together become associated came to be called Hebb’s rule and was instrumental in the
development of a powerful and influential form of artificial intelligence (AI), connectionism (see
Rumelhart, McClelland, & the PDP Research Group, 1986). Beyond these works, Hebb also
authored many other signal publications in psychobiology. For example, in 1946 he published an
article summarizing his research on the nature of fear. In 1949 he described the results of a study
in which animals were reared in either an enriched or an impoverished sensory environment. He
found that animals reared in an enriched sensory environment were relatively better learners as
adults. In a series of experiments run under his supervision, the effects of sensory deprivation on
cognitive processes were examined (for example, see Heron, 1957). In 1955 Hebb reported
research showing the relationship between levels of activity in the small brain structure, called
the reticular activating system (RAS), and cognitive and behavioral performance. The
examination of this relationship was called arousal theory. It was while they were doing research
on arousal theory in Hebb’s laboratory that James Olds and Peter Milner discovered
reinforcement centers in the brain (Olds & Milner, 1954). Buchtel (1982) provides an excellent
sample of Hebb’s influential articles on topics in psychobiology, and a complete list of Hebb’s
more than 80 publications is provided.

ROGER W. SPERRY
Roger Wolcott Sperry (1913–1994) was born in Hartford, Connecticut. He received his BA in
English from Oberlin College in 1935 and his PhD in zoology from the University of Chicago in
1941, where he learned neurosurgical techniques from the eminent neuroembryologist Paul
Weiss. After receiving his doctorate, Sperry studied with Lashley at the Yerkes Laboratories in
Florida (1942–1946). In 1946 he returned to the University of Chicago first as an assistant
professor of anatomy and then, in 1952, as assistant professor of psychology. In 1954 Sperry
moved to the California Institute of Technology in Pasadena (Caltech) as the prestigious Hixon
Professor of Psychobiology.

The Split-Brain
At Caltech, Sperry pursued his interest in the routes by which information is transferred from one
side of the cerebral cortex to the other. In a now-famous series of experiments, Sperry and his
colleagues discovered two possible routes for such interhemispheric transfer—the corpus
callosum (a large mass of fibers that connects the two halves of the cortex) and the optic chiasm.
The optic chiasm is the point in the optic nerve where information coming from one eye is
projected to the side of the cortex opposite to that eye. Sperry taught cats and monkeys to learn a
visual discrimination with a patch over one eye. He then tested for transfer by switching the
patch to the other eye and found complete interocular transfer. Sperry then began his search for
the mechanism by which information is transferred from one side of the cortex to the other. He
found that ablating either the corpus callosum or the optic chiasm alone or together after training
did not interfere with transfer. He also found that ablating either the corpus callosum or the optic
chiasm before training did not interfere with transfer. However, he found that ablating both the
corpus callosum and the optic chiasm before training eliminated interhemispheric transfer. Thus,
ablating the corpus callosum and the optic chiasm had in essence created two separate brains
with no exchange of information between them. For example, when an animal’s brain was split
in the manner just described and it was taught to make a visual discrimination with a patch over
one eye, it had no recollection of that learning when tested with the other eye (Sperry, 1961,
1964). A brain that has had its corpus callosum and its optic chiasm ablated is referred to as a
split-brain preparation. Sperry and his colleagues, Joseph Bogen and Philip Vogel, discovered
that humans suffering from severe drug-resistant, intractable epilepsy could benefit from having
their brains split in the manner described above. Presumably, with split-brain preparation, a
seizure begun in one hemisphere would not have a mechanism available to spread its influence to
the other hemisphere and thus increase its intensity. In many cases, patients treated in this way
improved enough to leave the hospital. In everyday living, these “split-brain” patients showed
almost no abnormality in spite of their radical surgery. Sperry and his colleagues developed a
number of tests that made it possible to study the function of each cerebral hemisphere
independently of the other. Although Paul Broca and others had provided information indicating
hemispheric specificity as early as 1831 and speculation concerning hemispheric specificity was
quite popular toward the end of the 19th century, information concerning hemispheric specificity
remained extremely limited. The additional knowledge provided by Sperry and his colleagues
was dramatic. They found that each hemisphere had its own characteristic range of cognition,
memory, emotion, and consciousness. Under Sperry’s leadership, research on the “left brain” and
the “right brain” became very popular. Unfortunately, some speculations concerning hemispheric
specificity began to exceed the facts. For example, it was speculated that some people are right-
brain dominated and others left-brain dominated and that tests could be devised that reveal this
domination. It was also speculated that educational practices could be employed to specifically
enhance either right- or left-brain functions. The belief that the two cortical hemisphere can be
educated independently goes back at least as far as BrownSequard (1874a, 1874b) and in one
form or another, has been entertained ever since. Jerre Levy, another one-time colleague of
Sperry, attempted to set the record straight in her article “Right Brain, Left Brain: Fact and
Fiction” (1985). In this article, Levy emphasizes the point that in people with normal brains, the
contributions of the two hemispheres to thought and behavior are inseparable. Levy concludes,
“The popular myths are misinterpretations and wishes, not the observations of scientists. Normal
people have not half a brain nor two brains but one gloriously differentiated brain, with each
hemisphere contributing its specialized abilities.… We have a single brain that generates a single
mental life” (1985, p. 44). At least as early as Fechner, there had been philosophical speculation
about how the two hemispheres related to conscious experience. Sperry’s work certainly
energized such questions. Sperry himself had a lifelong interest in the mind–body (brain)
problem and how that problem relates to human values, and many of his publications, especially
his later ones, reflected those interests. Sperry believed that consciousness emerges from brain
processes and, once emerged, has a causal relationship to behavior. Thus, Sperry was an
interactionist concerning the mind–body relationship. He believed that by correlating mental
events directly to brain processes, he avoided dualism. In his lifetime, Sperry published almost
300 articles in the most prestigious journals, and many of those articles were widely translated
(Puente, 1995). Among the many honors received by Sperry were the Karl Lashley Award of the
American Philosophical Society (1976); the Wolf Prize in Medicine (1979); the Ralph Gerard
Award from the Society of Neuroscience (1979); the Nobel Prize (1981; shared with Harvard
neuroscientists David Hubel and Torsten Wiesel); and the Lifetime Achievement Award from
the APA (1993). Sperry died in 1994, in Pasadena, California, at the age of 80, from a
degenerative neuromuscular disorder. At this point we have now mentioned the Nobel Prize in
physiology or medicine several times (including Pavlov’s award). As seen, research related to
psychology has won this prestigious award on occasion. Additional examples include Egas
Moniz (1949) for his work on the lobotomy, and Georg von Bekesy (1961) for his work on
hearing. In the previous chapter we noted the works of Sartre and Camus, who won the Nobel
Prize in literature for their existential writings. The Nobel Prize in Economics has also been
awarded to two cognitive scientists, Herbert Simon (1978) and Daniel Kahneman (2002), both of
whom we will cover in Chapter 19. Freud was nominated 11 times, but never won (and the
award is not given posthumously). For more on psychology and the Nobel Prize, including
Munsterberg’s attempt to have Wundt receive the award, see Benjamin (2003).