The Problem of Serial Order

in Behavior
K. S. LASHLEY
Harvard University and the Yerkes Laboratories
of Primate Biology

The previous speakers have approached our common problem by

considering the properties of the elementary units of which we believe
the cerebral structure to be built up. They have considered the kinds
of neural integration or behavior which can be anticipated from those
properties. The remaining members of the symposium have in their
research been concerned chiefly with the analysis of complex behavior,
seeking to derive general principles of neural integration from the in-
finitely complex products of that integration. Our common meeting
ground is the faith to which we all subscribe, I believe, that the
phenomena of behavior and of mind are ultimately describable in the
concepts of the mathematical and physical sciences. In my discussion
here, I have deliberately turned to the opposite extreme from the neu-
ron and have chosen as a topic, one aspect of the most complex type
of behavior that I know; the logical and orderly arrangement of
thought and action. Our discussion so far has dealt chiefly with the
conditions of input and of immediate switching in the nervous mecha-
nism, without explicit consideration of what is already going on within
the system.
My principal thesis today will be that the input is never into a
quiescent or static system, but always into a system which is already
actively excited and organized. In the intact organism, behavior is the
result of interaction of this background of excitation with input from
any designated stimulus. Only when we can state the general charac-
teristics of this background of excitation, can we understand the effects
of a given input.
The unpronounceable Cree Indian word "kekawewechetushekamik-
owanowow" is analyzed by Chamberlain (7) into the verbal root,
tusheka, "to remain," and the various particles which modify it as fol-
112
 K. S. Lashley 113
lows: ke(la)wow, the first and last syllables, indicating second person
plural; ka, a prefix of the future tense; we, a sort of imperative mode
expressing a wish; weche, indicating conjunction of subject and ob-
ject; mik, a suffix bringing the verb into agreement with a third person
subject and second person object; and owan, a suffix indicating that the
subject is inanimate and the object animate. A literal translation: "You
will I wish together remain he-you it-man you" or, freely, "may it remain
with you." This difference in structure between Cree and English
illustrates an outstanding characteristic of verbal behavior; the occur-
rence of predetermined, orderly sequences of action which are
unique for each language. In English the adjective precedes, in
French it follows the noun which it modifies. In English the move-
ment or action of the subject is expressed as early as possible after
the subject; in German the expression of action may be postponed until
all qualifying thoughts have been expressed. In a sentence discussing
this subject, Pick (20) introduces fifty-five words between the subject
and the principal verb. Each Chinese word, and to a lesser extent,
each English word, stands as an unchanging unit. In the highly inflec-
tive languages, such as Sioux, the form of almost every word in the
sentence may be altered, according to some attribute of the subject,
as when two objects rather than one or several are discussed.
The study of comparative grammar is not the most direct approach
to the physiology of the cerebral cortex, yet Fournie (10) has written,
"Speech is the only window through which the physiologist can view
the cerebral life." Certainly language presents in a most striking form
the integrative functions that are characteristic of the cerebral cortex
and that reach their highest development in human thought processes.
Temporal integration is not found exclusively in language; the coor-
dination of leg movements in insects, the song of birds, the control of
trotting and pacing in a gaited horse, the rat running the maze, the
architect designing a house, and the carpenter sawing a board present
a problem of sequences of action which cannot be explained in terms
of successions of external stimuli.
ASSOCIATIVE CHAIN THEORIES
In spite of the ubiquity of the problem, there have been almost no
attempts to develop physiological theories to meet it. In fact, except
among a relatively small group of students of aphasia, who have had to
face questions of agrammatism, the problem has been largely ignored.
It is not even mentioned in recent textbooks on neurophysiology or
physiological psychology, nor is there any significant body of experi-
mental studies bearing upon the problem. The spinal animal scarcely
114 The Problem of Serial Order in Behavior
exhibits serial activity, so the physiologist may be excused for over-
looking the phenomenon. On the other hand, psychologists have been
concerned chiefly with the question of whether or not the organizing
processes displayed in serial action are conscious, and very little with
the organization itself. I have chosen to discuss the problem of tem-
poral integration here, not with the expectation of offering a satis-
factory physiological theory to account for it, but because it seems to
me to be both the most important and also the most neglected problem
of cerebral physiology. Temporally integrated actions do occur even
among insects, but they do not reach any degree of complexity until
the appearance of the cerebral cortex. They are especially characteris-
tic of human behavior and contribute as much as does any single
factor to the superiority of man's intelligence. A clearer formulation
of the physiological problems which they raise should be of value, even
though a solution of the problems is not yet in sight.
I shall consider first some of the questions raised by the structure of
language, then turn to other forms of serial action for indications of
the nature of the nervous mechanisms involved.
To the best of my knowledge, the only strictly physiological theory
that has been explicitly formulated to account for temporal integration
is that which postulates chains of reflexes, in which the performance of
each element of the series provides excitation of the next. This concep-
tion underlay the "motor theories" of thinking which were advocated
by several psychologists early in this century. Watson (26) sought to
identify thought with inaudible movements of the vocal organs, linked
together in associative chains. The peripheral chain theory of language
was developed in greatest detail by Washburn (25). She distinguished
what she called "successive movement systems" and, although she drew
her examples from memorized series of nonsense syllables, her implica-
tion was that such series are typical of all language behavior. She de-
fined a movement system as "a combination of movements so linked
together that the stimulus furnished by the actual performance of
certain movements is required to bring about other movements." She
described speech as a succession of vocal acts in which the kinesthetic
impulses from each movement serve as a unique stimulus for the next
in the series (25, pages 11 ff.). Attempts to confirm these peripheral
theories by mechanical (Thorsen, 23) or electrical (Max, 19) recording
of muscular tensions have given no valid evidence in support of them.
It should be noted that, at the time when the theories were proposed,
it was generally believed that conduction in the nervous system is al-
ways downstream from sense organ to muscle, and that muscular con-
traction must always follow promptly on stimulation. The existence of
 K. S. Lashley 115
reverberatory circuits which could maintain central activity was
scarcely suspected.
The introspective psychology which objected to such peripheral
theories did not explicitly formulate an alternative neurological theory,
but there is implicit in it a view that verbal thought is a simple chain
of central processes in which each element serves to arouse the next
by direct association. Titchener, for example, maintained that the
meaning of a word (or of an auditory image in his system) consists
of the chain of associations which it arouses; that it has no meaning
until such a sequence has occurred. From this it must be inferred that
he was thinking in terms of a simple associative chain, since no other
relating process is suggested.
OBJECTIONS TO THE ASSOCIATIVE CHAIN THEORY
A consideration of the structure of the sentence and of other motor
sequences will show, I believe, that such interpretations of temporal
organization are untenable and that there are, behind the overtly ex-
pressed sequences, a multiplicity of integrative processes which can
only be inferred from the final results of their activity. There is an
extensive controversial literature dealing with this inferred integrative
activity. Pick (20) devotes almost his entire book, Die agrammatischen
Sprachstorungen, to reviewing discussions of the subject. Most of this
literature deals with the question of whether or not the integrative
processes are conscious. Much of this is irrelevant to the present topic,
but the advocates of so-called imageless thought did present a great
deal of material indicative of the complexity of the problem of thought
structure. From this, and other evidence which I shall present, I believe
that the production of speech involves the interaction of at least three,
possibly four, major neurological systems which are interrelated but
somewhat independently variable.
Let us start the analysis of the process with the enunciation of the
word. Pronunciation of the word "right" consists first of retraction and
elevation of the tongue, expiration of air and activation of the vocal
cords; second, depression of the tongue and jaw; third, elevation of
the tongue to touch the dental ridge, stopping of vocalization, and
forceful expiration of air with depression of the tongue and jaw. These
movements have no intrinsic order of association. Pronunciation of the
word "tire" involves the same motor elements in reverse order. Such
movements occur in all permutations. The order must therefore be
imposed upon the motor elements by some organization other than
direct associative connections between them. So, for the individual
movements in writing or typing the word, finger strokes occur in all
116 The Problem of Serial Order in Behavior
sorts of combinations. No single letter invariably follows g, and
whether gh, ga, or gw is written depends upon a set for a larger unit
of action, the word.
Words stand in relation to the sentence as letters do to the word; the
words themselves have no intrinsic temporal "valence." The word
"right," for example, is noun, adjective, adverb, and verb, and has
four spellings and at least ten meanings. In such a sentence as "The
mill-wright 011 my right thinks it right that some conventional rite
should symbolize the right of every man to write as he pleases," word
arrangement is obviously not due to any direct associations of the
word "right" itself with other words, but to meanings which are deter-
mined by some broader relations.
It has been found in studies of memorization of nonsense syllables
that each syllable in the series has associations, not only with adjacent
words in the series, but also with more remote words. The words in the
sentence have, of course, associations with more remote words as well
as with adjacent ones. However, the combination of such direct asso-
ciations will not account for grammatical structure. The different posi-
tions of the word "right" in the illustrative sentence are determined by
the meanings which the positions in relation to other words denote,
but those meanings are given by other associations than those with the
words in the spoken sentence. The word can take its position only when
the particular one of its ten meanings becomes dominant. This domi-
nance is not inherent in the words themselves.
From such considerations, it is certain that any theory of gram-
matical form which ascribes it to direct associative linkage of the
words of the sentence overlooks the essential structure of speech. The
individual items of the temporal series do not in themselves have a
temporal "valence" in their associative connections with other elements.
The order is imposed by some other agent.
This is true not only of language, but of all skilled movements or
successions of movement. In the gaits of a horse, trotting, pacing, and
single footing involve essentially the same pattern of muscular con-
traction in the individual legs. The gait is imposed by some mechanism
in addition to the direct relations of reciprocal innervation among the
sensory-motor centers of the legs. The order in which the fingers of the
musician fall on the keys or fingerboard is determined by the signature
of the composition; this gives a set which is not inherent in the associa-
tion of the individual movements.
 K. S. Lashley 117

THE DETERMINING TENDENCY

What then determines the order? The answer which seems most in
accord with common sense is that the intention to act or the idea to
be expressed determines the sequence. There are, however, serious
difficulties for this solution. There is not much agreement among psy-
chologists concerning the nature of the idea. The structuralist school,
under the leadership of Titchener, held that the idea consists of mental
images, often the auditory images of words, and the meanings are
nothing but sequences of such images. Describing the role of images
in his lecturing, Titchener wrote (24), "When there is any difficulty
in exposition, a point to be argued pro and con or a conclusion to be
brought out from the convergence of several lines of proof, I hear my
own voice speaking just ahead of me." What solution of the lecture
problem for the lazy man! He need not think but only listen to his
own inner voice; to the chain of associated auditory images. A be-
haviorist colleague once remarked to me that he had reached a stage
where he could arise before an audience, turn his mouth loose, and
go to sleep. He believed in the peripheral chain theory of language.
(This clearly demonstrates the superiority of behavioristic over intro-
spective psychology. The behaviorist does not even have to listen to his
own inner voice.)
Seriously, such positions offer no solution for the problem of tem-
poral integration. Titchener finds his grammar ready made and does
not even raise the question of the origin of the succession of images.
The chain-reflex theory, while definite, is untenable.
The third view of the nature of the idea was developed by a group
known as the "Wiirzburg School" (see Boring, 4); exponents of image-
less thought. It held that some organization precedes any expression
that can be discovered by introspective or objective means. Thought is
neither muscular contraction nor image, but can only be inferred as
a "determining tendency." At most, it is discovered as a vague feeling
of pregnancy, of being about to have an idea, a Bewustseinslage. It is
not identical with the words which are spoken, for quite frequently no
word can be recalled which satisfactorily expresses the thought, and
we search a dictionary of synonyms until a word or phrase is found
which does seem appropriate.
In his discussion of the relation of thought to speech, Pick (20)
accepts this point of view, but he asserts further that the set or the
idea does not have a temporal order; that all of its elements are
COtemporal. Evidence in support of this conclusion comes, for ex-
ample, from translation of one language into another which has a
118 The Problem of Serial Order in Behavior
different sentence structure. I read a German sentence, pronouncing
the German words with no thought of their English equivalents. I then
give a free translation in English, without remembering a single word
of the German text. Somewhere between the reading and free transla-
tion, the German sentence is condensed, the word order reversed, and
expanded again into the different temporal order of English. According
to Epstein (9), the polyglot shifts readily from one language to an-
other, expressing the same thought in either, without literal transla-
tion. The readiness with which the form of expression of an idea can
be changed, the facility with which different word orders may be
utilized to express the same thought, thus is further evidence that the
temporal integration is not inherent in the preliminary organization
of the idea.
THE SCHEMA OF ORDER
The remaining alternative is that the mechanism which determines
the serial activation of the motor units is relatively independent, both
of the motor units and of the thought structure. Supporting evidence
for this may be found in the mistakes of order, the slips and inter-
ferences which occur in writing and speaking. For some time I have
kept records of errors in typing. A frequent error is the misplacing or
the doubling of a letter. These is typed t-h-s-e-s, look as 1-o-k-k, ill as
i-i-1. Sometimes the set to repeat may be displaced by several words.
The order is dissociated from the idea. Earlier, in preparing this
paper, I wrote the phrase, "maintain central activities." I typed min,
omitting the a, canceled this out and started again; ama. The impulse
to insert the a now dominated the order. I struck out the a and com-
pleted the phrase, only to find that I had now also dropped the a from
activities. This example suggests something of the complexity of the
forces which are at play in the determination of serial order and the
way in which conflicting impulses may distort the order, although the
primary determining tendency, the idea, remains the same.
The polyglot, who has become proficient in a secondary language,
who thinks in it and even dreams in it, may still tend to use the gram-
matical structure of his native tongue. If, as in French, that tongue
applies gender to inanimate things, the English pronouns referring to
them may take the gender of the French equivalents, though the
French nouns are not thought. The German postponement of the verb
or the Magyar use of the past infinitive may be incorporated in the
new language. In such cases, the structuring seems to be dissociated
both from the content and from the simple associative connections
of the words themselves.
 K. S. Lashley 119
The ease with which a new structure may be imposed on words is
illustrated by the quickness with which children learn hog Latin. The
form which I learned involved transposing the initial sound of each
word to the end of the word and adding a long a. Thus—at-thay an-
may oes-gay own-day e-thay eet-stray. Some children become very
facile at such inversions of words, and re-structure new words without
hesitation. From such considerations it seems to follow that syntax is
not inherent in the words employed or in the idea to be expressed. It is
a generalized pattern imposed upon the specific acts as they occur.

"PRIMING" OF EXPRESSIVE UNITS

There are indications that, prior to the internal or overt enunciation
of the sentence, an aggregate of word units is partially activated or
readied. Evidence for this comes also from "contaminations" of speech
and writing. The most frequent typing errors are those of anticipation;
the inclusion in the word being typed of some part of a word or word
structure which should properly occur later in the sentence. It may be
only a letter. Thus I wrote, wrapid writing, carrying the w from the
second word to the first. Not infrequently words are introduced which
should occur much later in the sentence, often five or six words in
advance.
In oral speech, Spoonerisms illustrate the same kind of contamina-
tion. The Spoonerism is most frequently an inversion of subject and
object: "Let us always remember that waste makes haste." But it may
be only a transposition of parts of the words: "Our queer old dean"
for "our dear old queen." The frequency with which such contamina-
tions occur is increased by haste, by distraction, by emotional ten-
sion, or by uncertainty and conflict as to the best form of expression.
In some types of aphasia the tendency to disordered arrangement of
words is greatly increased, and, in extreme cases, the attempt to speak
results in a word hash with complete loss of grammatical organization.
Professor Spooner, after whom such slips are named, was probably
suffering from a mild form of aphasia. In these contaminations, it is as
if the aggregate of words were in a state of partial excitation, held in
check by the requirements of grammatical structure, but ready to
activate the final common path, if the effectiveness of this check is in
any way interfered with.
In his Psychopathology of Even/day Life, Freud has given numer-
ous examples of similar contaminations of action outside the sphere of
language. We do not need to accept his theories of censorship and
suppression lo account lor such slips. They are of the same order as
120 The Problem of Serial Order in Behavior
misplacements in typing and represent contaminations of co-existing,
determining tendencies to action.
Such contaminations might be ascribed to differences in the relative
strength of associative bonds between the elements of the act, and
thus not evidence for pre-excitation of the elements or for simultaneous
pre-excitation. However, the understanding of speech involves essen-
tially the same problems as the production of speech and definitely de-
mands the postulation of an after-effect or after-discharge of the sen-
sory components for a significant time following stimulation. Thus, in
the spoken sentence, "Rapid righting with his uninjured hand saved
from loss the contents of the capsized canoe," the associations which
give meaning to righting are not activated for at least 3 to 5 seconds
after hearing the word.** I shall refer later to other evidence for such
long after-discharge of sensory excitations. The fact of continued acti-
vation or after-discharge of receptive elements and their integration
during this activation justifies the assumption of a similar process
during motor organization. The processes of comprehension and
production of speech have too much in common to depend on wholly
different mechanisms.
INTERNAL AND OVERT SPEECH
One other point with respect to the organization of speech: The
earlier literature on aphasia emphasized the distinction of internal and
overt speech. The aphemia of Broca and the pure motor aphasia of
Wernicke and later writers were held to be a loss of the ability to
enunciate without loss of ability to think in words and without paraly-
sis of the organs of speech. The brain insult was assumed to affect
only the transition from the thought to the enunciation of the word.
We may doubt the existence of instances of such "pure" defects and
question the reliability of the early clinical examinations in view of
the more careful analyses that have been made since 1917, but the dis-
tinction of internal and overt speech is still valid and the transition
still unexplained. Watson interpreted internal speech as inaudible
movements of the vocal organs, and Jacobsen (15) and Max (19) have
given evidence of changes in muscular tonus during verbal thinking or
thought of movement. This is far from proving that the motor dis-
charge is essential for the internal formation of words, however.
I once devised an instrument to record small movements of the
0
Dr. Lashley ingeniously laid the groundwork for this three paragraphs earlier,
when he mentions "wrapid writing." The audience all heard, "Rapid writing
with his uninjured hand," etc. "Capsized canoe" required a complete and amusing
about-face. EDITOR.
 K. S. Lashley 121
tongue. Within the limits of its sensitivity, it showed that in silent
thinking the tongue usually drops to the back of the mouth and shows
no detectable movement. Verbal problems, such as the correct squaring
of three-place numbers, could be carried out with no trace of overt
movement. If, however, I urged the subject to hurry or if I slapped his
face, his tongue came forward and showed movements corresponding
to the syllabification of internal speech or of the computation he was
performing. This I interpret as indicating that internal speech may be
carried out wholly by processes within the nervous system, with some
unessential discharge upon the final common path for vocal move-
ments. Facilitation of the motor path, either by increased emotional
tension or by "voluntary" reinforcement, increases its excitability until
the same central circuits whose activity constitutes internal speech are
able to excite the overt movements. This aspect of the language func-
tion is irrelevant to the problem of syntax or serial order, but is im-
portant as illustrating a further point in the dynamics of the cerebrum.
Many activities seem to require for their performance both a specific
patterning and also a general facilitation, a rise in dynamic level.
There are, I think, indications that hemiplegia and motor aphasia are
primarily expressions of a low level of facilitation rather than a loss of
specific integrative connections which are involved in the use of lan-
guage or in the patterning of our movements. A monkey, for example,
after ablation of the precentral gyrus may seem unable to use the arm
at all, but if emotional excitement is raised above a certain level, the
arm is freely used. As soon as the excitement dies down, the arm is
again hemiplegic. I have seen something of the same sort in a human
hemiplegic. The problem of the availability of memories, which was
raised earlier in the discussion here, may find a partial solution in such
fluctuations in dynamic level. In many of the organic amnesias the pat-
tern of integration seems to be retained but can be reactivated only by
an abnormally intense sensory or central reinforcement.
GENERALITY OF THE PROBLEM OF SYNTAX
I have devoted so much time to discussion of the problem of syntax,
not only because language is one of the most important products of
Iniman cerebral action, but also because the problems raised by the
organization of language seem to me to be characteristic of almost all
olhcr cerebral activity. There is a series of hierarchies of organization;
I he order of vocal movements in pronouncing the word, the order of
words in the sentence, the order of sentences in the paragraph, the
rational order of paragraphs in a discourse. Not only speech, but all
skilled acts set-in to involve the saint: problems of serial ordering, even
 122 The Problem of Serial Order in Behavior
down to the temporal coordination of muscular contractions in such a
movement as reaching and grasping. Analysis of the nervous mecha-
nisms underlying order in the more primitive acts may contribute ul-
timately to the solution even of the physiology of logic.
It is possible to designate, that is, to point to specific examples of,
the phenomena of the syntax of movement that require explanation,
although those phenomena cannot be clearly defined. A real definition
would be a long step toward solution of the problem. There are at least
three sets of events to be accounted for. First, the activation of the
expressive elements (the individual words or adaptive acts) which do
not contain the temporal relations. Second, the determining tendency,
the set, or idea. This masquerades under many names in contemporary
psychology, but is, in every case, an inference from the restriction of
behavior within definite limits. Third, the syntax of the act, which can
be described as an habitual order or mode of relating the expressive
elements; a generalized pattern or schema of integration which may be
imposed upon a wide range and a wide variety of specific acts. This is
the essential problem of serial order; the existence of generalized
schemata of action which determine the sequence of specific acts, acts
which in themselves or in their associations seem to have no temporal
valence.
I shall turn now to other phenomena of movement which may be
more readily phrased in physiological terms and which may suggest
some of the mechanisms underlying serial order.
DURATION AND INTENSITY OF NERVOUS DISCHARGE
A consideration of the control of extent and rate of movement sup-
ports the view that sensory factors play a minor part in regulating the
intensity and duration of nervous discharge; that a series of movements
is not a chain of sensory-motor reactions. The theory of control of
movement which was dominant at the turn of the century assumed
that, after a movement is initiated, it is continued until stopped by
sensations of movement and position, which indicate that the limb has
reached the desired position. This theory was opposed by a good bit ol
indirect evidence, such as that accuracy of movement is increased
rather than diminished with speed. I had opportunity to study a
patient who had a complete anesthesia for movements of the knee
joint, as a result of a gunshot wound of the cord (16). In spite of the
anesthesia, he was able to control the extent and speed of movements
of flexion and extension of the knee quite as accurately as can a normal
person.
The performance of very quick movements also indicates their in-
 K. S. Lashley 123
dependence of current control. "Whip-snapping" movements of the
hand can be regulated accurately in extent, yet the entire movement,
from initiation to completion, requires less than the reaction time for
tactile or kinesthetic stimulation of the arm, which is about one-eighth
of a second, even when no discrimination is involved. Such facts force
the conclusion that an effector mechanism can be pre-set or primed
to discharge at a given intensity or for a given duration, in inde-
pendence of any sensory controls.
CENTRAL CONTROL OF MOTOR PATTERNS
This independence of sensory controls is true not only of intensity
and duration of contraction of a synergic muscle group but is true
also of the initiation and timing of contraction of the different muscles
in a complex movement. The hand may describe a circular movement
involving coordinated contractions of the muscles of the shoulder,
elbow, and wrist in about y10 second, and the stopping of movement
at a given position, of course, is only a small fraction of that time. The
finger strokes of a musician may reach sixteen per second in passages
which call for a definite and changing order of successive finger move-
ments. The succession of movements is too quick even for visual reac-
tion time. In rapid sight reading it is impossible to read the individual
notes of an arpeggio. The notes must be seen in groups, and it is
actually easier to read chords seen simultaneously and to translate
them into temporal sequence than to read successive notes in the
arpeggio as usually written.
Sensory control of movement seems to be ruled out in such acts.
They require the postulation of some central nervous mechanism which
fires with predetermined intensity and duration or activates different
muscles in predetermined order. This mechanism might be represented
by a chain of effector neurons, linked together by internuncials to
produce successive delays in firing. In some systems the order of action
may be determined by such a leader or pace-setter. Buddenbrock (6)
has shown for the stick insect, and Bethe (3) for a number of animals
from the centipede to the dog, that removal of one or more legs results
in a spontaneous change in the order of stepping. Thus, for the insects,
the normal order is alternate stepping of the first pair of legs with right
lirst, left second, right third leg advancing together. With removal of
the left first leg, the right first and left second alternate and the order
brromes right first, left third, right third stepping together, with left
.scc-oiid and right second advancing together, instead of alternately.
These investigators were interested in spontaneity of reorganization,
rather than in the mechanism of coordination, and did not propose any
124 The Problem of Serial Order in Behavior
theory for the latter. They did show, however, that it is necessary to
remove the leg completely to get the change in pattern of movement;
sensory impulses from a limb stump would prevent it. Such coordina-
tion might be explained, perhaps, by a combination of loss of excit-
ability in the centers of the absent limb, by the excitation of the re-
maining anterior center as a leader or pace-setter, and the spread of
alternate waves of inhibition and excitation from the more anterior to
the more posterior limb centers. The spontaneous change in coordina-
tion shows, however, that the coordination is not due to the action of
predetermined anatomic paths but is the result of the current physio-
logical state of the various limb centers.
Such an hypothesis implies also the assumption of a polarization of
conduction along the neuraxis, with the order of excitation determined
by the spatial arrangement of the centers of the legs. I see no other
possibility of accounting for the facts. The examples of circular move-
ment and of finger coordination, involving temporal integration of
movements, seem to call for a similar hypothesis. They might be
ascribed to an habitual linkage of the movements through a simple
chain of internuncials but for two facts. First, such series are usually
reversible at any point or can be started from any point. This would
require the assumption of a second set of internuncials habituated to
conduct in the opposite direction, and this in turn leads to the further
assumption of a polarization of conduction. Second, such patterns of
coordinated movement may often be transferred directly to other
motor systems than the ones practiced. In such transfer, as to the left
hand for writing, an analysis of the movements shows that there is not
a reduplication of the muscular patterns on the two sides, but a re-
production of movements in relation to the space coordinates of the
body. Try upside-down mirror writing with the left hand and with
eyes closed for evidence of this. The associative linkage is not of
specific movements but of directions of movement. An analysis of sys-
tems of space coordinates suggests mechanisms which may contribute
to production of such series of movements in a spatial pattern.
SPACE COORDINATE SYSTEMS
The work of Sherrington, Magnus, and others on postural tonus and
reflexes has defined one level of spatial integration rather fully, yet it is
doubtful if these studies have revealed the effective neural mechanism.
The work has shown that the tonic discharge to every muscle in the
postural system is influenced by afferent impulses from every other
muscle, toward increased or decreased activity, according to its syner-
 K. S. Lashley 125
gic or antergic action. To these influences are added vestibular and
cerebellar effects. Diagrammatically these mutual influences of the
muscular system may be represented by separate reflex circuits from
each receptor to every muscle, as Sherrington (21, p. 148) has done.
But no neuro-anatomist would, I am sure, maintain that such separate
circuits or paths exist. What the experiments on posture actually show
is a correlation of sensory stimulation and of tonic changes in a net-
work of neurons whose interconnections are still undefined. The reac-
tions isolated experimentally have the characteristics of simple directly
conducted reflexes, but their combination results in patterns of move-
ment and posture which have definite relations to the axes of the body
and to gravity.
This postural system is based on excitations from proprioceptors.
The distance receptors impose an additional set of space coordinates
upon the postural system, which in turn continually modifies the co-
ordinates of the distance receptors. The dropped cat rights itself, if
either the eyes or the vestibular senses are intact, but not in the ab-
sence of both. The direction of movement on the retina imposes a
directional orientation on the postural system. Conversely, the gravita-
tional system imposes an orientation on the visual field. Upright objects
such as trees or the corners of a room appear upright, at no matter
what angle the head is inclined. Derangement of the vestibular system
can disturb the distance orientation or the orientation of the receptors,
as in the apparent swaying of the vertical as a result of the after-images
of motion following hours of rocking in a small boat.
There are other, still more generalized systems of space coordinates.
We usually keep track of the compass points or of some more definite
index of direction by a temporal summation of the turns made in
walking, though not always with success. Finally, there is a still more
plastic system in which the concepts of spatial relations can be volun-
tarily reversed, as when one plays blindfold chess alternately from
either side of the board.
Explanation of these activities, these complex interactions, in terms
of simple isolated interconnections of all of the sensory and motor
elements involved seems quite improbable on anatomic grounds and
is ruled out by results of our experiments on sectioning of the spinal
cord. Ingebritzen (14) studied rats with double hemisection of the
cord; one-half of the cord cut at the second, the other at the fifth
cervical segment. In the best case only a small strand of the spino-
ccrcbellar tract of one side remained intact. These rats were able to
balance in walking, oriented to visual stimuli, scratched with the right
126 The Problem of Serial Order in Behavior
or left hind foot according to the side of the face stimulated, were able
to run mazes correctly, and even learned to rise on the hind feet and
push down a lever with the forepaws in opening a box.
The alternative to the isolated-path theory of the space coordinates
is that the various impulses which modify postural tonus are poured
into a continuous network of neurons, where their summated action
results in a sort of polarization of the entire system. I shall consider
later the integrative properties of such a net. For the moment I wish
to emphasize only the existence of these systems of space coordinates.
Their influences pervade the motor system so that every gross move-
ment of limbs or body is made with reference to the space system.
The perceptions from the distance receptors, vision, hearing, and
touch are also constantly modified and referred to the same space
coordinates. The stimulus is there, in a definite place; it has definite
relation to the position of the body, and it shifts with respect to the
sense organ but not with respect to the general orientation, with
changes in body posture.
Memories of objects usually give them position in the space system,
and even more abstract concepts may have definite spatial reference.
Thus, for many people, the cardinal numbers have definite positions on
a spiral or other complicated figure. What, if anything, such space
characters can contribute to temporal integration is an open question.
They provide a possible basis for some serial actions through interac-
tion of postural and timing mechanisms.
RHYTHMIC ACTION
The simplest of the timing mechanisms are those controlling rhyth-
mic activity. T. Graham Brown (5) first showed by his studies of de-
afferented preparations that the rhythmic movements of respiration
and progression are independent of peripheral stimulation and are
maintained by a central nervous mechanism of reciprocal innervation.
He suggested that this mechanism of reciprocal innervation, rather
than the simple reflex, is the unit of organization of the whole nervous
system. He thus foreshadowed, in a way, the conception of reverbera-
tory circuits which is coming to play so large a part in neurological
theory today. Hoist (13) has recently shown that the rhythmic move-
ment of the dorsal fin of fishes is a compound of two superimposed
rhythms, that of its own innervation and that of the pectoral fins. These
two rhythms are centrally maintained.
Musical rhythms seem to be an elaboration of the same sort of thing.
The time or beat is started and maintained at some definite rate, say
160 per minute. This rate is then imposed upon various activities. The
 K. S. Lashley 127
fingers of the musician fall in multiples of the basic rate. If the leader
of a quartet speeds up the time or retards, all the movements of the
players change in rate accordingly. Not only the time of initiation but
also the rate of movement is affected. The violinist, in a passage re-
quiring the whole bow, will draw the bow from frog to tip at a uniform
rate for the required number of beats, whether the tempo is fast or
slow. With practiced violinists, the rate of movement is extremely
accurate and comes out on the beat at the exact tip of the bow.
Superimposed on this primary rhythm is a secondary one of empha-
sis, giving the character of 3/4, 4/4, 6/4, or other time. The mechanism
of these rhythms can be simply conceived as the spread of excitation
from some centers organized for reciprocal innervation; as a combina-
tion of the principles of Brown and of Hoist. There are, however, still
more complicated rhythms in all music. That of the melodic line is
most uniform. In much music, the melodic progression changes in
2, 4, or some multiple of 4 measures. In improvisation, the performer
keeps no count of measures, yet comes out almost invariably in a reso-
lution to the tonic of the key after some multiple of eight measures.
Here a generalized pattern is impressed on the sequence, but it is a
simpler pattern than that of grammatical structure. It only requires the
recurrence of a pattern at certain rhythmic intervals; a pick-up of a
specific pattern after so many timed intervals.
There are, in addition, still less regular rhythms of phrasing and
emphasis. Parallels to these can be found in speech. The skilled ex-
temporaneous speaker rounds his phrases and speaks with a definite
though not regular rhythm.
The rhythms tend to spread to almost every other concurrent ac-
tivity. One falls into step with a band, tends to breathe, and even to
speak in time with the rhythm. The all pervasiveness of the rhythmic
discharge is shown by the great difficulty of learning to maintain two
rhythms at once, as in three against four with the two hands. The
points to be emphasized here are the widespread effects of a rhythmic
discharge indicating the involvement of almost the entire effector sys-
tem, the concurrent action of different rhythmic systems, and the im-
position of the rate upon both the initiation and speed of movement.
Consideration of rhythmic activity and of spatial orientation forces
the conclusion, I believe, that there exist in the nervous organization,
elaborate systems of interrelated neurons capable of imposing certain
types of integration upon a large number of widely spaced effector
c l r m r n l s : i i i llic one case transmitting temporally spaced wave's of
facilitative excitation to all effector elements; in the other imparting
u directional p o l a r i / a l i o n to both receptor and effector elements. These
128 The Problem of Serial Order in Behavior
systems are in constant action. They form a sort of substratum upon
which other activity is built. They contribute to every perception
and to every integrated movement.
INTERACTION OF TEMPORAL AND SPATIAL SYSTEMS
Integration ascribed to the spatial distribution of excitations in the
nervous system has been much more intensively studied than the
temporal aspects of nervous activity. Theories of integration are based
almost exclusively upon space properties, time entering only in theories
of facilitation, inhibition, and after-discharge. In cerebral functions,
however, it is difficult to distinguish between spatial and temporal
functions. The eye is the only organ that gives simultaneous informa-
tion concerning space in any detail. The shape of an object impressed
on the skin can scarcely be detected from simultaneous pressure, but
the same shape can readily be distinguished by touch when traced on
the skin with a moving point or when explored by tactile scanning.
The temporal sequence is readily translated into a spatial concept.
Even for vision it might be questioned whether simultaneous stimula-
tion gives rise directly to space concepts. The visual object is generally
surveyed by eye movements, and its form is a reconstruction from such
a series of excitations. Even with tachistoscopic exposures, the after-
discharge permits a temporal survey, and, with visual fixation, shifts of
attention provide an effective scanning.
Since memory traces are, we believe, in large part static and persist
simultaneously, it must be assumed that they are spatially differen-
tiated. Nevertheless, reproductive memory appears almost invariably
as a temporal sequence, either as a succession of words or of acts.
Even descriptions of visual imagery (the supposed simultaneous re-
productive memory in sensory terms) are generally descriptions of
sequences, of temporal reconstructions from very fragmentary and
questionable visual elements. Spatial and temporal order thus appear
to be almost completely interchangeable in cerebral action. The trans-
lation from the spatial distribution of memory traces to temporal
sequence seems to be a fundamental aspect of the problem of serial
order.
I spoke earlier of the probability of a partial activation or priming
of aggregates of words before the sentence is actually formulated from
them. There is a great deal of evidence for such preliminary facilitation
of patterns of action in studies of reaction time and of word association.
Reaction time, in general, is reduced by preliminary warning or by
instructions which allow the subject to prepare for the specific act
required. In controlled association experiments, the subject is in-
 K. S. Lashley 129
structed to respond to the stimulus word by a word having a certain
type of relation to it, such as the opposite or a part of which the
stimulus is the whole; black-white, apple-seed. The result is an attitude
or set which causes that particular category to dominate the associative
reaction. Whether such preliminary reinforcement is to be ascribed to
accumulation of excitatory state, as defined by Sherrington (21), or to
some other physiological process, the facts of behavior assure that it
is a genuine phenomenon and plays a decisive role in determining the
character of the response.
Once the existence of such states of partial activation is recognized,
their possible role in temporal integration must be considered. There
are indications that one neural system may be held in this state of
partial excitation while it is scanned by another. Here is an example.
A series of four to six numbers is heard: 3-7-2-9-4. This is within the
attention or memory span and is almost certainly not remembered in
the sense in which one's telephone number is remembered, for memory
of it is immediately wiped out by a succeeding series of numbers.
While it is retained in this unstable way, subject to retroactive
inhibition, the order of the numbers can be reasserted: 3-7-2-9-4,
3-2-7-9^, 4-9-2-7-3, and the like. It is as if, in this case, a rhythmic
alternation can suppress alternate items, or a direction of arousal can
be applied to the partially excited system. Another example which
illustrates even more clearly the spatial characteristics of many mem-
ory traces is the method of comultiplication, used in rapid mental cal-
culation. In attempts to play a melody backward, we have a further
illustration. I find that I can do it only by visualizing the music
spatially and then reading it backward. I cannot auditorily transform
even "Yankee Doodle" into its inverse without some such process, but
it is possible to get a spatial representation of the melody and then to
scan the spatial representation. The scanning of a spatial arrangement
seems definitely to determine, in such cases, the order of procedure.
Two assumptions are implied by this. First, the assumption is that the
memory traces are associated, not only with other memory traces, but
also with the system of space coordinates. By this I do not mean that
the engram has a definite location in the brain; our experiments show
conclusively that such is not the case. Rather, when the memory trace
is formed it is integrated with directional characters of the space
system, which give it position in reference to other associated traces.
Second, the assumption is that these space characters of the memory
trace can be scanned by some other level of the coordinating system
and so transformed into succession.
This is as far as I have been able to go toward a theory of
130 The Problem of Serial Order in Behavior
serial order in action. Obviously, it is inadequate. The assumptions
concerning spatial representation and temporal representation may
even beg the question, since no one can say whether spatial or temporal
order is primary. Furthermore, such determining tendencies as the
relation of attribute to object, which gives the order of adjective and
noun, do not seem to be analyzable into any sort of spatial structure
or for that matter, into any consistent relationship. I have tried a
number of assumptions concerning the selective mechanism of gram-
matical form (spatial relations, the relative intensity or prominence
of different words in the idea, and so on) but I have never been able
to make an hypothesis which was consistent with any large number
of sentence structures. Nevertheless, the indications which I have
cited, that elements of the sentence are readied or partially activated
before the order is imposed upon them in expression, suggest that
some scanning mechanism must be at play in regulating their tem-
poral sequence. The real problem, however, is the nature of the se-
lective mechanism by which the particular acts are picked out in this
scanning process, and to this problem I have no answer.
Such speculations concerning temporal and spatial systems do
little more than illustrate a point of view concerning nervous organiza-
tion which is, I believe, more consistent both with .what is known of
the histology and elementary physiology of the brain and also with
behavior phenomena than are the more widely current theories of
simple associative chains of reactions.
Nearly forty years ago Becher (2, page 243) wrote: "There is no
physiological hypothesis which can explain the origin and relations
of temporal forms in mental life; indeed, there is no hypothesis
which even foreshadows the possibility of such an explanation." The
situation is little better today, but I do feel that changing conceptions
of the fundamental organization of the nervous system offer more hope
for a solution of such problems than did the physiological knowledge
available when Becher wrote. However, we are still very far from
being able to form an explicit explanation of temporal structure.
THE FUNDAMENTAL MECHANISM OF INTEGRATION
Neurological theory has been dominated by the belief that the
neurons of the central nervous system are in an inactive or resting
state for the greater part of the time; that they are linked in relatively
isolated conditioned reflex arcs and that they are activated only when
the particular reactions for which they are specifically associated are
called out. Such a view is incompatible both with the widespread
effects of stimulation which can be demonstrated by changes in tonus
 K. S. Lashley 131
and also with recent evidence from electrical recording of nervous
activity. It is now practically certain that all the cells of the cere-
brospinal axis are being continually bombarded by-nerve impulses from
various sources and are firing regularly, probably even during sleep.
The nervous activity which they in turn elicit depends upon the cur-
rent physiological state of the neurons with which they are connected.
It is probably not far from the truth to say that every nerve cell of
the cerebral cortex is involved in thousands of different reactions.
The cortex must be regarded as a great network of reverberatory cir-
cuits, constantly active. A new stimulus, reaching such a system,
does not excite an isolated reflex path but must produce widespread
changes in the pattern of excitation throughout a whole system of
already interacting neurons.
The facts of cerebral structure support such a view. The cortex
is composed chiefly of neurons with short axons. LeGros Clark (8)
has found for the striate area of the monkey that Marchi degeneration
extends for only a short distance from a point of injury. In the striate
area of the rat, I have never been able to trace degeneration beyond
three or four cell diameters from the margin of a lesion, and I
believe that this lack of long transcortical fibers is true of other
areas as well as of the visual cortex. Visual perception reveals close
integration of different parts of the striate areas in spite of the absence
of long association fibers. In the visual cortex of the rat there are only
19 neurons for each afferent fiber. To produce the animal's visual
acuity, all of the afferent fibers must be firing continually. There are
approximately 34,000 cell bodies in the lateral geniculate nucleus of the
rat, and the minimum number of visual units necessary to produce
the visual acuity of the rat is actually above this figure. (The acuity
is determined by direct experimental tests.) These figures should be
of interest in relation to the numerical values cited by Dr. von
Neumann. The number of cells in the visual cortex of the rat is
only about 106, and in some of my experiments where I have removed
the greater part of the visual cortex the capacity for discrimination
of visual forms has been retained when no more than 20,000 cells of
the visual cortex remain. There is also evidence that no part of the
cerebral cortex except the visual areas is essential for visual perception
and memory.

DR. LORENTE DE N6: What is the number of afferents in the optic

nerve?
on. LASHLEY: There are 290,000 afferents in the optic nerve of the
rat, and the figure is reduced to 34,000 in the lateral geniculate. The
132 The Problem of Serial Order in Behavior
actual numbers are 9,000,000 myoids, 290,000 ganglion cells, and
34,000 cells in the lateral geniculate. That may include cells with short
axons also. There are about 125,000 cells in each of the five layers of
the cortex. These figures are for one eye and hemisphere.
DR. VON NEUMANN: In the human being the corresponding number is
about 125,000,000 for the first, isn't it?
DR. LASHLEY: I know of no figure for that level.
DR. LORENTE DE NO: It hasn't been analyzed in any way.
DR. VON NEUMANN: The optic nerve corresponds to the second one?
DR. LASHLEY: Yes. The axons of the ganglion cells pass through
the optic nerves. There is an average of 300 visual cells firing into
each central pathway. There are fewer than 5 cells in the receptive
layer of the visual cortex of the rat for each afferent fiber of the optic
radiation and only 19 cells per afferent fiber in the entire visual
cortex. Since the visual acuity of the rat requires that all of the
34,000 cells of the radiation be firing constantly, it seems certain that
all of the neurons within the striate areas, the visual cortex, must
be firing constantly. There is a good bit of evidence that all of the
integrative functions of vision are carried out within the striate areas.
In the rat, I have removed, from one or another animal, practically
every other part of the isocortex without disturbing visual perception
or memory. With monkeys I have removed the supposed visual asso-
ciative areas without producing any significant loss of visual functions.

These facts lead to the conclusion that the same cells in the visual cor-
tex participate in a great variety of activities. Practically all of the cells
of the area must be fired by every visual stimulation, and these same
cells must be the ones which retain the visual memories. The con-
clusion follows that differential responses depend upon the pattern
of cells which are excited in combination. The visual cortex is a network
of cells of short axon without long interconnections between its parts
or with other cortical areas. Its integrative functions are an expression
of the properties of such a network.
The same conception must be applied to other cortical areas. There
are, of course, long association tracts in the cortex, such as the corpus
callosum, the superior longitudinal fasciculus, and the temporo-frontal
tracts. Once, 26 years ago, I suggested facetiously that these might be
only skeletal structures, since I could find no function for them. No
important functions of these tracts have yet been demonstrated. Section
of the corpus callosum produces only a slight slowing of reaction
time, ipsilateral as well as contralateral (Akelaitis, 1); section of
occipito-frontal fibers produces, perhaps, a temporary disturbance
 K. S. Lashley 133
of visual attention but no other symptoms. The integrative functions
seem to be carried out as well without as with the main associative
tracts. The major integrative functions must, therefore, be carried
out by the network of cells of short axon. The properties of such
networks of cells must be analyzed before the mechanisms of the
cerebral cortex can be understood. Something can be inferred from
the characteristics of excitability of cells and their arrangement in
recurrent loops. If, as seems a necessary conclusion from the histology
of the striate area, all of the cells of the network are subject to constant
excitation and are firing whenever they recover from the refractory
state, then mutual interference of circuits will produce complicated
patterns throughout the area, patterns which will stabilize in the
absence of differential stimulation, as is perhaps indicated by the
regularity of the alpha rhythm. Any new afferent impulses reaching the
area can only produce a reorganization of the existing pattern. What
happens at any particular point in the system, as at an efferent neuron,
is the statistical outcome of the interaction of myriads of neurons,
not of the transmission of impulses over a restricted path, of which
that efferent cell forms a link. It is possible to isolate parts of the
system by operative means or by anesthetics and so to get a one-to-one
relation of stimulus locus and responding muscles, from which the
reflex mechanism has been inferred. As Goldstein (12) has pointed
out, however, the parts isolated in the reflex are influenced by a multi-
plicity of effects in the intact organism of which there is little or
no trace in the isolated preparation.
I can best illustrate this conception of nervous action by picturing
the brain as the surface of a lake. The prevailing breeze carries
small ripples in its direction, the basic polarity of the system. Vary-
ing gusts set up crossing systems of waves, which do not destroy
the first ripples, but modify their form, a second level in the system of
space coordinates. A tossing log with its own period of submersion
sends out periodic bursts of ripples, a temporal rhythm. The bow
wave of a speeding boat momentarily sweeps over the surface, seems
to obliterate the smaller waves yet leaves them unchanged by its
passing, the transient effect of a strong stimulus. Wave motion is not
an adequate analogy because the medium which conveys the waves
is uniform, whereas the nerve cells have their individual characteristics
of transmission which at every point may alter the character of the
transmitted pattern.
The great number of axon terminations on every nerve cell has not
been considered in theories of integration. It implies, of course, that the
cell can be fired by impulses from a variety of sources. But it also
134 The Problem of Serial Order in Behavior
suggests another possibility, more fruitful for understanding of integra-
tive processes. A nerve impulse arriving over a single axon terminal
may not fire the cell but may modify its excitability to impulses from
other sources. In an elaborate system of neurons such subthreshold
effects might establish a pattern of facilitation which would determine
the combination of cells fired by subsequent excitations. The space
coordinate system and various types of set or priming may be pictured
as patterns of subthreshold facilitation pervading the network of
neurons which is activated by the more specific external stimulus.
Such a view of the mechanism of nervous action certainly does
not simplify the problems nor does it as yet provide any clue to the
structuring that constitutes the set or determining tendency, or to the
nature of such relations as are implied in the attribute-object, oppo-
sites, or other abstract concepts. A few relations seem reducible to
spatial terms, part-whole, for example, but even for these there is
no clear conception of the neural basis of their space properties. These
considerations do not, I believe, contradict fundamentally the basic
conceptions that have been formulated by Dr. McCulloch. They do,
however, indicate a direction of necessary elaboration. The nets active
in rhythmic and spatial organization are apparently almost coextensive
with the nervous system. The analysis must be extended to the proper-
ties of such nets; the way in which they are broken up into reactive
patterns in the spread of excitation, to give, for example, directional
propagation or its equivalent. I strongly suspect that many phenomena
of generalization, both sensory and conceptual, are products, not of
simple switching, but of interaction of complex patterns of organization
within such systems.
SUMMARY
The problems of the syntax of action are far removed from anything
which we can study by direct physiological methods today, yet in
attempting to formulate a physiology of the cerebral cortex we cannot
ignore them. Serial order is typical of the problems raised by cerebral
activity; few, if any, of the problems are simpler or promise easier
solution. We can, perhaps, postpone the fatal day when we must face
them, by saying that they are too complex for present analysis, but
there is danger here of constructing a false picture of those processes
that we believe to be simpler. I am coming more and more to the
conviction that the rudiments of every human behavioral mechanism
will be found far down in the evolutionary scale and also represented
even in primitive activities of the nervous system. If there exist, in
human cerebral action, processes which seem fundamentally different
 K. S. Lashley 135
or inexplicable in terms of our present construct of the elementary
physiology of integration, then it is probable t'hat that construct is
incomplete or mistaken, even for the levels of behavior to which it is
applied.
In spite of its present inadequacy, I feel that the point of view
which I have sketched here holds some promise of a better understand-
ing of cerebral integration. Attempts to express cerebral function in
terms of the concepts of the reflex arc, or of associated chains of neu-
rons, seem to me doomed to failure because they start with the assump-
tion of a static nervous system. Every bit of evidence available indi-
cates a dynamic, constantly active system, or, rather, a composite of
many interacting systems, which I have tried to illustrate at a primi-
tive level by rhythm and the space coordinates. Only when methods
of analysis of such systems have been devised will there be progress
toward understanding of the physiology of the cerebral cortex.

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DISCUSSION
DR. KLTJVER: In 1914, there appeared a book which, in the opinion
of the author, marked an epoch—since during the remaining years of
his life he celebrated the day the manuscript was completed instead of
his birthday. The author was von Monakow, and the title of the book
was Die Lokalisation im Grosshirn. There is a remarkable connection
between von Monakow's monumental contribution to neurology in
1914 and Dr. Lashley's presentation today. In my opinion, this is the
first time since 1914 that a neurological thinker has presented such a
trenchant analysis of the role of the time factor in behavior. If you
study von Monakow's book you find that the central concept of his
work is the concept of a "chronogenic localization," a concept which
has been almost completely ignored during the intervening decades.
It is not possible today to enter into a discussion of the numerous
problems raised by Dr. Lashley's presentation. As regards the relation
of thinking to temporal organization, we are, it seems to me, confronted
with a certain dilemma. Ideas, concepts, and meanings themselves
have no reference to time and space, and, yet, the expression, formula-
tion, and identification of ideas are processes proceeding in time and
occurring in space. It is the relation of ideas to temporal and spatial
factors which is of interest to the physiological psychologist. Even the
 Discussion 137
fact that the verbal expression of an idea is linked up with a temporal
sequence of acoustic events raises intricate questions. I got an inkling
of some of the difficulties on a visit to Vogt's Institute for Brain Re-
search, where an investigator utilized the methods of experimental
phonetics in studying the speech utterances of neurological and psy-
chiatric patients. He found, on the one hand, that the curves he had
obtained could not be interpreted without considering the acoustic
events "carriers of meanings" and, on the other hand, that certain
forms of expression survived speeding up and slowing down and even
the complete reversal of the temporal sequence. I do not know
whether any further attempts have been made to utilize the techniques
of experimental phonetics.
I should like to comment on only one other point. It is true that the
same idea may be expressed in various languages by utilizing a dif-
ferent temporal order of words, just as the same perceptual relation
may be recognized on the basis of different relata; but it is also true
that the structure of language may prevent us from expressing ideas
and thinking along certain lines. The fact that the language I am using
now does not possess a word for a rough, wet object undergoing a
color change may have the consequence that I overlook such an object
or that I do not identify, remember, and think about it. Consequently,
such an item may not or cannot enter a temporal sequence utilized in
expressing an idea. In this connection Allers' experiments on "word
nearness" and "word remoteness" are of particular interest. According
to Allers, every attempt to express an idea in words induces a tendency
towards selecting items in terms of word nearness. Word nearness and
word remoteness may become decisive in determining the temporal
sequence and in expressing or not expressing an idea. The line of
demarcation between word-near and word-remote elements is subject
to marked variations and even varies from individual to individual. No
doubt, the sequence of phenomena calling for an expression in words,
the sequence of words, and the sequence of ideas are sequences of a
different order. The great value of Dr. Lashley's analysis lies in the
fact that it exhibits the significant factors involved in the expression of
ideas as well as in other instances of serial ordering, and that it
utilizes such factors effectively in formulating mechanisms of cerebral
activity.
DR. KOHLER: Dr. Lashley rightly said that human orientation in space
is mostly achieved in a sequence of events. We move our eyes, or our
attention wanders, and so forth, as we inform ourselves about our
environment. Curiously enough, however, while the process of becom-
ing so informed occurs in time, and consists of successive acts, the
138 The Problem of Serial Order in Behavior
result appears almost independent of this temporally extended origin.
Apart from objective changes, which we may ignore in this connec-
tion, the space in which we live, and the objects around us, appear for
the most part perfectly stationary. Dr. Lashley agrees, of course, with
this observation. I wonder whether he can tell us how in this case the
history of becoming informed disappears so completely from the final
information.
DB. GERARD: I tried yesterday, in a brief summary, to remain strictly
non-partisan while indicating the various viewpoints that had been
presented and which were still to come. Actually, I find it impossible
to think through or even towards the complexities of behavior if
restricted to atomic units traveling along atomic fibers. Even the work
we did on the frog brain, which showed a potential field present in
which moving waves of electrical activity could travel and cross com-
plete anatomical cuts, seems inexplicable at that level. My own inclina-
tion is to think of the discrete elements and the patterns of their
activity as involved in the more particular performances of the nerv-
ous system, in the immediate sequences of action, as emphasized, but
also as active in the more integrated behavior of the whole of the
nervous system. This doesn't mean that, in any way, I am disagreeing
with Dr. Lashley's final comment. One can hope, with some confidence,
that all the properties of the most complex nervous system will prove
to be adumbrated in the properties of the simplest element and con-
nection when we know those properties sufficiently well.
I would like to raise one particular point. Some years ago, Culler
and his colleagues reported an observation which may have been
elaborated or may have been refuted; I haven't been following that
literature. If it is correct, it seems very relevant to this whole problem,
and I'd like Dr. Lashley's reaction to it. In dogs, conditioned in some
particular way, a circumscribed region of the cortex, not in the area
either of the receptor or of the effector, became electrically excitable
and produced the conditioned response. When the conditioned reflex
was extinguished, that area became inexcitable. There was, thus, a
sharp localization of conditioning in that particular case, one of the
few striking exceptions to the more general pattern of mass activity.
What is the present status of such work?
DR. HALSTEAD: I couldn't help thinking, as I listened to this excellent
example of what I would call the higher mental processes, that I could
have prevented all this by putting a lesion in a particular part of one
nervous system—the anterior portion of Dr. Lashley's prefrontal areas
If instead I had put a lesion in area 19 of Dr. Lashley, certain im-
portant elements in his presentation would undoubtedly have been
 Discussion 139
lacking. Yet we could have removed probably each lobe of the brain
individually without disturbing the basic form or organization.
I have been greatly impressed with the case that Dr. Lashley has
made for non-specific, non-mosaic representation. I would like to
emphasize, and if I am incorrect I hope that he will point it out, that
his view is not incompatible with progressive specialization of function.
The language functions that he talked about are not disturbed, except
when particular areas in the brain are involved. The visual functions
that he described are not disturbed, except when particular regions of
the brain are involved. The important thing is that, within the region,
there seems to be the possibility for equivalence to arise.
DR. LORENTE BE NO: It is typical of Dr. Lashley that he places on the
board a series of figures that no anatomist had ever determined.
Dr. Lashley had to determine them himself in order to go on with his
work.
I don't think that I have yet fully appreciated the value of Dr. Lash-
ley's talk. I will have to read it and study it thoroughly, but, while I
was listening, there was going through my head a mental picture
of the future development of a number of experiments that I intend to
perform—suggested to me by Dr. Lashley's speech. One of the prob-
lems that Dr. Lashley has suggested to me is very accessible of ex-
perimentation in the study and treatment of things of purely central
origin. There is one very good situation in which we meet discharges
of very long duration, which can be created in the absence of every
peripheral influence. You know that stimulation of the labyrinth pro-
duces a rhythmical movement of the eye and establishes the fact that
this rhythm is not dependent upon the feedback from the eye muscles
for its production. Impulses can be recorded directly from the nerves
to the eye muscles, and therefore any sensory feedback is eliminated.
What the labyrinth sends to the nervous system is a continuous stream
of impulses, not interrupted rhythmically. The interruptions are pro-
duced in the nervous system, and, furthermore, can be produced when
the labyrinths are extirpated. Thus in the usual vector nystagmus, if
one labyrinth is extirpated, the spontaneous nystagmus will last some
10 or 12 or 15 days after that, and then the spontaneous nystagmus
ceases. If the second labyrinth is then extirpated, the nystagmus re-
appears and lasts for 7 or 8 days. This is a purely central phenomenon
—there are no impulses of any kind coming from the periphery into the
centers, and the nystagmus can be recorded from the motor nerves
after the extirpation of the eye. Consequently, it is describable as
rhythmical activity, and is the type of rhythm that I'm going to in-
vestigate as soon as 1 get to it.
140 The Problem of Serial Order in Behavior
I would also like to make a remark in reference to Dr. Gerard's
statement about neural elements in cerebral physiology. We have been,
for many years, considering only the transmission of impulses as the
natural mode. Now, this potential field of yours is another mechanism
by which the nerve cells act upon others. Sometimes neurons act upon
others by the transmission of impulses, and sometimes by the potential
field of the currents that they establish.
DR. WEISS: The great value of Dr. Lashley's presentation lies in the
fact that it places rigorous limitations upon the free flight of our fancy
in designing models of the nervous system, for no model of the nervous
system can be true unless it incorporates the properties here described
for the real nervous system. You will recognize that our current models
are far short of satisfying this condition.
Dr. Lashley's theses receive crucial support from the student of
development. The embryologist has long been up against a tremendous
task. While the physiologist and psychologist deal with the ready-
made machine of the nervous system and can add to it as many proper-
ties as he thinks necessary, the embryologist must explain just how
such an immensely intricate, yet orderly, thing can develop. These
studies are still in their infancy, but a few things have already come
out which gibe completely with the conclusions that Dr. Lashley has
presented here; for instance, the relative autonomy of structured pat-
terns of activity, and the hierarchical principle of their organization. As
I said previously, the nervous system is not one big monotonic pool
whose elements can be freely recombined in any number of groupings,
thereby giving an infinite variety of nervous responses. This used to be
the old idea of the associationists, and it is utterly incompatible with
what we have learned about the development of the nervous system
and its functions in animals. I wouldn't have spoken here if Dr. Lash-
ley hadn't said that he is convinced that whatever happens in the brain
of man is foreshadowed, and, in principle, is the same as what happens
in the lower organisms. Therefore, assuming that premise, I would now
like to add a few comments.
First, we know from the lower organisms that the working of the
central nervous system is a hierarchic affair in which functions at the
higher levels do not deal directly with the ultimate structural units,
such as neurons or motor units, but operate by activating lower pat-
terns that have their own relatively autonomous structural unity. The
same is then true for the sensory input, which does not project itself
down to the last final path of motor neurons, but operates by affecting,
distorting, and somehow modifying the pre-existing, preformed pat-
terns of central coordination, which, in turn, then confer their distor-
 Discussion 141
tions upon the lower patterns of effection and so on. The final output
is then the outcome of this hierarchical passing down of distortions and
modifications of intrinsically preformed patterns of excitation, which
are in no way replicas of the input. The structure of the input does not
produce the structure of the output, but merely modifies intrinsic nerv-
ous activities that have a structural organization of their own. This
has been proved by observation and experiment. Coghill has shown
that the motor patterns of the animal develop prior to the development
of sensory innervation. I have shown, as others have, that the removal
of the sensory innervation does not abolish the coordination of motor
activities. Moreover, coordinated motor functions of limbs and other
parts develop even if these parts have been experimentally prevented
from ever becoming innervated by sensory fibers. Therefore, the sen-
sory pathway can have nothing to do with the structure of the motor
response. There are still some authors who try to save the old associa-
tionist idea that actually the input shapes the structure of the output.
I think that they are fighting a losing fight, and I think that today's
discussion ought to have given them the coup de grace. The essential
independence of the structure of motor activity is dramatically demon-
strated when one exchanges and reverses the limbs of animals and then
finds them crawling backwards whenever they aim to crawl forwards,
and vice versa. Many of you have seen my film where this had been
done in the developed animal, but the same operations have been done
in embryos and these animals have then functioned in reverse from the
very beginning. What more spectacular expression can there be of the
intrinsic primacy of the motor patterns of behavior for which the
external input acts only as a selective trigger?
The autonomous development of central functions raises a further
question: Are the response patterns preformed as merely static condi-
tions, or are they of dynamic nature, that is, properties of automatic
central activities? Lashley has favored the latter view, and, again, ex-
periments in lower animals furnish corroborative evidence. Intrinsic
automatic rhythms have been shown, for instance, by Adrian in the
brain stem of the goldfish and in insect ganglia, by Prosser in other
arthropods, by Bremer and by von Hoist in the spinal cord, and by
Bethe in jellyfish. I have shown experimentally that any group of bul-
bar or spinal nerve cells taken from vertebrates, if deprived of their
structural bonds of restraining influences and allowed to undergo a
certain degree of degradation, will display permanent automatic,
rhythmic, synchronized activity of remarkable regularity. Rhythmic
activity, therefore, seems a basic property of pools of nervous elements.
W l i r l l i c i I l i i s ; i u ( o i i i a ( i s i i > is generated by neuron switch works or
142 The Problem of Serial Order in Behavior
rather by the rhythmic waxing and waning of metabolic activity ex-
pressing itself in the electric field, as suggested by Dr. Gerard, is a
secondary question. The principal point is that the rhythm is not some-
thing generated through an input rhythm, but is itself a primary
rhythm which may be released and even speeded up or retarded by the
input, but is not derived from the input. So we have experimental
evidence that autonomy of pattern, rhythmic automatism, and hier-
archical organization are primary attributes of even the simplest nerv-
ous systems, and I think that this unifies our view of the nervous sys-
tem. I conclude with expressing my hope that today's discussion will
really mark a turning point in the building of neurological theories so
that more regard will be given to all the actual facts that we do know
about nervous systems.
DR. MCCULLOCH: There is a paper by Wiener, written when he was
working with Bertrand Russell, in which he described the space of im-
plications. It is a peculiarly degenerate sort of space. Now I, like many
other men, find it difficult to think in motion. When I think about mo-
tion, I freeze it in a four-space. If you take Wiener's degenerate space
of implication and make yourself a four-space image of it, you can
visualize these rhythms as fixed forms. You can practically superimpose
large numbers of these forms easily in the mind instead of battling with
two or three. That is the first point, and it is merely a technical device.
Second, you remember that I said that on the way down from the
cerebral cortex I was utterly unhappy about our knowledge of how the
cortex did this or that on account of our ignorance of the lower mech-
anisms. I want to say a word about two rhythmical affairs that we are
now working on. If, in the intact animal, you stimulate the nucleus
caudatus, the animal just slumps. If, however, you cut bilaterally the
cerebral peduncles, through which all impulses coming from the cere-
brum have to go to reach the hindbrain and lower structures, and
if you then stimulate the nucleus caudatus, you get beautiful automat-
ically associated rhythmical movements. One, for example, resembles
a cat fishing in the gold-fish bowl. Another resembles feline boxing.
There is always some part of the body, arm, leg, or face, leading in
each of these movements. The sequence of each movement belonging
to this group starts in the axial system, then goes to the axio-appen-
dicular, and thence to the appendicular, moving ever outward. I have
never seen any evidence of somatotopic localization in the input to
the nucleus caudatus. Yet here, in its output, on coming down any one
puncture, stimulation at superior positions gives responses beginning in
the tail and hindleg, deeper, in the forelegs, and near the very bottom,
in the face. From the bottom you get complicated lapping and chew-
 Discussion 143

ing movements just as you do from the amygdala. You get similar
movements from the putamen under these circumstances and always
this representational reversal, the lowest part being represented high-
est. Now, if you go down their descending systems, whether you go
down the ansa lenticularis or go into the bunch of fibers that pass
into the subthalamus and turns down, you can follow right down,
stimulating them with similar results. Let me remind you that your
stimulus is just a series of impulses, say 40 or 30 per second. The fre-
quency of the movements increases with the voltage of stimulation, to
a maximum less than 10 per second. You continue to get these rhyth-
mical movements as you proceed downstream until you reach the
pneumotaxic center, when, instead of other movements, panting super-
venes. Downstream of the pneumotaxic center, just as you get only
torsion to the right or else to the left, so you get only inhalation or else
exhalation, or only facilitation or else relaxation. Now the interesting
thing is that these rhythmical movements increase in frequency as you
increase the voltage of stimulation up to a limiting value of nearly 10
per second. Above that, you never get them, for hypertonus begins to
come into the background and stops the movement.
This system, whatever else it does, takes a stream of impulses from
the basal ganglia, which is merely a continuous stream, over more or
fewer parallel paths and produces rhythmical performances. Thus,
whatever makes the rhythm is downstairs; it need not be in the cerebral
cortex or in the basal ganglia. These rhythmical movements originate
downstream. By destruction upstream, you may release the same thing.
DR. LINDSLEY: Dr. Lashley's discussion of the temporal schema im-
posed upon the motor system, was of considerable interest to me. I
don't know whether the spontaneous rhythms of the cortex have any
relationship to this temporal schema, but there are a good many sug-
gestive things that we can pick up in a rather superficial look at cortical
activity. We can get, through the electroencephalogram, things which
have been rather impressive to me. No one has been able to demon-
strate conclusively that any of the cortical rhythms of spontaneous
character has a direct relationship to the resting tremors of the normal,
or, for that matter, the neurologically abnormal person, despite the
fact that their frequencies may be similar in range. However, in the
brain waves of new-born infants and even of fetal infants, it is possible
to demonstrate rhythms, particularly in the motor area of the cortex,
and they seem to be at least grossly correlated with some of the serial
and sequential orders of motor activity. The correlation must remain a
very rough one at the present state, because we have rather inade-
quate techniques for investigation.
144 The Problem of Serial Order in Behavior
The frequency that one finds in the fetus or in the new-born infant
over the motor areas of the brain is much slower than that in the adult
stage. Whether this has any relation to the similar lack of sequential
organization which one finds in these early stages of development, I do
not know.
Another thing which has been impressive to me is the nature of the
frequencies that one records from the central nervous system—the
multiple or submultiple character of the rhythms one encounters, not
only as a developmental feature of the maturing brain, but in the ma-
tured individual when the normal spontaneous rhythm breaks up and
multiplies itself or divides in half. Just what relationship these changes
in rhythm bear to actual behavior, again I do not know.
I have been particularly interested in trying to devise experiments
to help get a better correlation between these superficially observed
electrical phenomena of the cortex and the behavioral activities of the
motor system. I find this very difficult to deal with in terms of se-
quential activities which are continually flowing along. It is very hard
to devise a system which will do more than suggest possible correlates.
On the other hand, there are phenomena we encounter both in the
normal and the abnormal individual, which seem to bear some rela-
tionship to this serial order. I'd like to have Dr. Lashley comment if he
will on whether the breaks in a sequence may not perhaps give us clues
to some of the relationships. By this I refer to various types of blocking
that one encounters, the blocking in the tapping of a motor rhythm, the
blocking in the speech pattern of normal individuals and stutterers, the
blocking in color naming, and a variety of others. Some of them (for
example, in free associations) represent a longer time scale of block-
ing, but will do essentially the same thing. I have tried in some in-
stances to correlate certain phenomena of the alpha rhythm with these
patterns, such phenomena as the blocking of the alpha rhythm, or the
out of phase character of the usually synchronized activity of the two
cerebral hemispheres. I found, in some instances, very suggestive cor-
relations, but I must confess that the proof that alpha blocking and the
out of phaseness in the two hemispheres are related to blocking in
behavior has not yet satisfied me.
DR. LASHLEY: I have been rather embarrassed by some of the flattering
remarks made today. I think that here we must also consider back-
ground effects. Boring has pointed out that no progress is ever made
until the time is ripe for that progress, and that the individual who
makes the contribution is a very minor matter. If one doesn't, another
will very shortly. What I have said today is really scarcely more than a
compilation and summary of the thoughts of the people who have
 Discussion 145

studied the aphasia problem. I have summarized again a series of

problems which have confronted the students of aphasia for many
years, to which Monakow and many others have contributed, Pick
especially, and I have found their views helpful in considering such
problems.
I have also been very much impressed with the similarity of the
problems with which the behaviorist is confronted, and the problems
with which the student of embryology must deal. The problem of form
seems to me to be too similar in the two cases to be altogther due to
different kinds of mechanisms.
There have been perhaps too many problems and questions raised
here for me to attempt to answer them in detail. With regard to Pro-
fessor Halstead's remark about the removal of my frontal lobes: In
fantasy, I have thought perhaps that my most important contribution
when I reach retirement age would be to have my frontal lobes re-
moved and see what I could do without them. I have less confidence
than Dr. Halstead that it would preclude the production of something
of interest. We have little experimental evidence of intellectual defect
from uncomplicated removal of the prefrontal lobes.
The conception of cerebral organization that I have sketched here
is the result of repeated failures to confirm simpler hypotheses. At-
tempts to trace the course of conditioned reflex arcs or to discover
the location of the traces of specific habits have been repeatedly un-
successful. They have revealed areas specialized for different functions,
but within those areas the parts have proved to be equivalent or equi-
potential for a great number of functions. Analysis of effective stimuli
and of adaptive responses has shown that these are not dependent
upon particular neurons but upon the spatial and temporal relations
of excitation within shifting fields of nervous activity. Attempts to
develop theories of neural integration in terms of chemical gradients
or electrical potentials are difficult or impossible to relate to the es-
tablished facts of axon conduction, however well they may seem to
account for the facts of behavior.
I agree thoroughly with Dr. McCulloch that the transmission of
excitation by the individual neuron is the basic principle of nervous
organization. However, the nervous activity underlying any bit of be-
havior must involve so many neurons that the action of any one cell
can have little influence upon the whole. I have come to feel that we
must conceive of nervous activity in terms of the interplay of impulses
in a network of millions of active cells.
In perception, in action, and in the preservation of memory traces
there are systems or anatomic structures whose parts seem to be equi-
146 The Problem of Serial Order in Behavior
potential. The equipotential systems of which we have more definite
knowledge all work by reduplication of elements or parts, of molecular
or gross mechanical structure. The problem of how a reduplication of
functional units might arise in the nervous system has seemed to me a
basic one in neurology. The analogy with wave form and interference
patterns is an attempt to meet that problem. In wave interference,
patterns are reduplicated in different parts of the field, and I think
it likely that nerve impulses, spreading at definite rates through a
nerve net, may also produce reduplicated interference patterns. These
would make possible the production of structural changes constituting
memory traces at numerous points throughout the system. Such a con-
cept is highly speculative, difficult to apply to concrete instances of
behavior, but is as far as I have been able to go toward a solution of
the problem of equipotentiality.
I mentioned briefly the problem of the dynamic level of nervous
activity and the probable role of non-specific facilitation in the pro-
duction of the shift from internal to overt speech or in determining
the availability of memories. There are certainly other variables which
affect the level of activity: changes in the chemical environment and
the metabolic activity of the neurons. I believe that many of the
symptoms following cerebral trauma or disease are due to such wide-
spread chemical changes, resulting perhaps from vascular damage,
rather than to actual destruction of nervous tissue. Aside from sensory
defects, clinical symptoms more often suggest changes in dynamic
level than the loss of specific integrations.
Dr. Kohler has raised the question of the stationary state of the
world around us. I have tried to deal with this in terms of the constant
space coordinates maintained by the postural system. The conceptual
construct of the data obtained by scanning is closely integrated with
this space system. I confess that I can form no clear picture of the
actual physical mechanism.
Dr. McCulloch has given a beautiful illustration of the spatial
determination of rhythmic activity by spread of excitation from a focal
point of stimulation. I wish that such a mechanism were applicable
to the problems of the syntax of action with which I have been dealing
but I see no way in which such a simple mechanism can be used to
account for the more complicated behavior. As for the mathematical
representation of time as a fourth dimension: This may help to con-
ceptualize motion, but the actual functions of the brain are carried out
by a three-dimensional system. How is time or temporal order repre-
sented within the three dimensions? Mathematical symbolization of
time as a fourth dimension does not suggest a clue.
 Functional Differences between
the Occipital and Temporal Lobes"
WITH SPECIAL REFERENCE TO THE
INTERRELATIONS OF BEHAVIOR AND
EXTRACEREBRAL MECHANISMS

HEINRICH KLUVER
The Division of the Biological Sciences,
University of Chicago

I shall first briefly consider certain behavior alterations following

bilateral removal of the occipital lobes in subhuman primates, more
particularly, the changes observed in rhesus monkeys.

FUNCTIONS OF THE OCCIPITAL LOBES

I shall not describe the techniques I have developed for testing such
animals (34-37, 40). Nor shall I bother you with the details of the
experimental analysis. I shall simply point out that such animals in
which the geniculostriate system has been eliminated have been
studied for a long time before and after the operations. In some in-
stances, thirty to forty-five trials per day have been given for more than
2000 days after the lobectomy. In studying the behavior of the bilateral
occipital monkey I have been chiefly concerned with analyzing the
responses of the dark-adapted animal to luminous stimuli differing in
brightness, area, shape, color, and/or distance from the eye as well as
the responses to intermittent light stimuli differing in light-dark ratios
and flash frequencies per second. The results of this laborious analysis
may be briefly summarized by saying that all or practically all dif-
ferential reactions of the bilateral occipital monkey to visual stimuli
can be understood by assuming the effectiveness of differences in the
density of luminous flux at the eye or, expressed otherwise, differences
in the quantity of light entering the eye (39, 41, 43). The eye of such
* Aided by the Dr. Wallace C. and Clara A. Abbott Memorial Fund of the
University of Chicago and by the Commonwealth Fund.
147