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^r. l^iiies Mocazm,







Delivered before the Royal College of Puysicians,
June, 1890.


Prqfe^or of Neuropathology , King's College, Lowion ; Physician to King's College Hospital
and to the National Hospital for the Paralysed and Epileptic.

322 North Craig St.,






Reprinted from the BRITISH Medical Jovbnax.


Lectube I.

Introductory Remarlrs — Effects of Ablation of the Cerebral Hemis-
pheres : (1) in Osseous Fishes — Vision in Brainless Fishes —
Vulpian's Experiments — Steiner's Experiments — (2) in Frogs —
Experiments of Goltz and Steiner — Schrader's Experiments:
(3) in Birds — Flourens' Experiments — Longet's Experiments —
Vision in Brainless Birds — Experiments of McKendrick, etc. —
Munk's Experiments — Schrader's Experiments : (4) in Mam-
mals — Experiments on Rabbits — Longet's Experiments —
Vision inBrain'ess Rabbits — Christiani's Experiments; Munk's
Experiments — Goltz's Experiments on Dog- — General Con-
clusions — Intelligence of the Lower Centres — Spontaneity —
Conditions of Consciousness — Relations of the Hemispheres
to the Lower Centres — The Question of Localisation of
Function — Gall's Opinions — Flourens' Doctrines — Bouillaud's
Experiments and Observations — Dax's Observations; Broca's
Observations — Hughlings Jackson's Views — Brown-S^quard's
Views — Exner's Views on Localisation — Fritsch and Hitzig's
Experiments — The Author's Experiments — Excitability of the
Cerebral Cortex — Mechanical Excitability— Chemical Excit-
ability—Electrical Excitability — Diffusion of Currents — Excit-
obility of the Medullary Fibres — Motor Fibres of the Internal
Ci,psule— Relative Excitability of the Cortex and Medullary
Fibres — Laws of Excitability of the Cortex— Esperiments of

Franck and Pitres pp. 1—24

Lecture II.

Electrical Reactions— Prefrontal Area — Oculo-motor Area — Leg
Area — Arm Area — Area of Face. Tongue, Larynx, etc —Mar-
ginal Convolution — Angular Gyrus and Occipital Lobe —
Superior Temporal Gyrus — Hippocampal Lobule and Gyrus
Hippocampi — Electrical Reactions in Man — Character and
Differentiation of the Electrical Reactions — Signification of
the Electrical Reactions — Sensory and Motor Centres — The
Sensory Centres: (1) The Visual Centres— Electrical Reactions
of the Occipito-Angular Region — The Author's Experiments —
Exp-riments of Luciani and Tamburini, Schafer; Significa-
tion of the Oculomotor Reactions — Experiments of Danillo,
Bechterew, Munk— -The Author's Earlier Experiments on the
Angular Gyrus and Occipital Lobe — Later Experiments of the
Author and Yeo — Munk's Experiments — Experiments of

Horsley and Schafer pp. 27—48

Lecture III.

Visual Centre«< Continued — Experiments of Schafer and Sanger-
Brown— Effects of Lesions of the Occipito-Temporal Region —
Experiments of Sanger-Brown and Gilman-I'hompson — Ex-
periments of Lannegrace — Lesions of the Angular Gyrus —
Experimc^nts of Munk, Schafer, and the Author — Relations of
the Angular Gyrus— Hemiopic Pupillary Reaction— Lanne-
grace's Theory of Cerebral Amblyopia— Pathology of Hemiopia
in Man — Lesions of the Cuneus- Word Blindness— Visual
Centres in Lower Vertebrates — Experiments of Hitzig, Goltz,
Dalton, Munk, Loeb, Luciani, Bechterew, Gilman-Thompson
and Sanger-Brown — Munk's Scheme of the Visual Centres in
Dogs — Loeb's Experiments- Visual Centres in Rabbits— Visual
f'entres in Pigeons — Visual Centres in Owls — General
Conclusions pp. 51— 72



Lecture IV.
The Auditory Centre — Methods of Investigation — Electrical Re-
actions — Earlier Experiments of the Author — Experiments of
the Author and Yeo— Schafer's Experiments— Eecent Experi-
ments of the Author — The Auditory Centre according to Munk
— Luciani and Tamburini's Experiments — Comparative Ana-
tomy of the Auditory Centre— Deafness from Cerebral Disease
in Man ; observations of Shaw, Wernicke and Friedlander —
Word-deafness ; its Pathology — Irritative Lesions of Auditory
Centre — Atrophy of Auditory Centre — Central Relations of
the Auditory Nerve ; observations of Baginsky Flechsig, and
Bechterew. Centre of Tactile and Common Sensation— The
Sensory Paths in the Spinal Cord — Experiments of Ludwig
and Woroschiloff — Ascending Degeneration in Spinal Cord —
The Author's Experiments on the Spinal Cord of Monkeys —
Hemisection of the Spinal Cord — Paths of the Muscular
Sense — Analysis of Recorded Cases — Observations by the
Author .' pp. 75-99

Lectitbe V.
Tactile Centre continued — Course of Sensory Tracts towards Brain
— Position of Sensory Tracts in crus cerebri — Position of
Sensory Tracts in Internal Capsule — Central Relations of
Sensory Tracts — Earlier Experiments of the Author on Hip-
pocampal Region — Experiments of the Author and Yeo —
Experiments of Horsley and Schafer on Hippocampal Region
and Gyrus Fornicatus— Relations of the Falciform Lobe —
France's observations on Descending Degeneration from
Lesions of gyrus fornicatus. Olfactory and Gustatory Centres
— Anatomical Relations of Olfactory Tract — Broca's Views —
Zuckerkandl on the Anatomy of the Olfactory Centre — Com-
parative Anatomy of the Hippocampus ; Sir William Turner's
observations — Relations of the Anterior Commissure — Obser-
tions of Flower, Meynert, Ganser, etc. — Relations of the
Fornix — Electrical Irritation of Hippocampal Lobule — Inves-
tigation of the Sense of Smell in Monkeys — Earlier Experi-
ments of the Author — Experiments of Schafer and Sanger-
Brown — Recent Experiments of the Author — Munk's observa-
tions on Dogs— Luciani's Views — Conclusions from Clinical
Observations pp. 103-126

Lecture VI.
Motor Centres— Rolandic Area — Effects of Destruction of the
Motor Centres in Monkpys — Experiments of the Author,
Horsley and Schafer — Effects of Lesions of the Marginal
Gyrus — The Laryngeal Centres — Bilateral Relations of the
Cerebral Hemispheres — Lesions of the Motor Area in Man —
Characters of Cortical Paralysis — Goltz's Observations on Dogs
— Lesions of the Motor Area and Affections of Sensation —
Observations of the Author, Horsley and Schafer, Goltz and
Bechterew — Analysis of Clinical Cases — Motor Paralysis and the
Muscular Sense — Muscular Sense in Hemianaesthesia — Bastian's
Views — Experiments of Marique, Exner and Paneth — Relations
of the Spinal and Cerebral Motor Centres. The Frontal
Centres— Anatomical Relations of the Frontal Centres— Effects
of Electrical Irritation — Destruction of the Po*t-frontal and
Pre-frontal Centres — I'sychical Effects of Lesions of Frontal
Lobes— Conclusion ... pp. 129 — 152





Ma. President and Gentlemen, — While highly appreciating
the distinguished honour of being appointed Croonian Lecturer of
the College of Physicians, I must confess to having undertaken
the onerous duties of the office with considerable hesitation and
trepidation, for, though the subject which I have chosen is one to
which I have devoted a good deal of attention, and which, in one
of its aspects, namely, The Localisation of Cerebral Disease, I
have already had the honour of discussing before you as Gulston-
ian Lecturer, yet, considering the enormous amount of work that
has been done in this department in recent years, and the nume-
rous problems which still remain unsolved, I have felt that, with
my other duties, time and strength would scarcely permit me to
do justice to my subject. I could not feel satisfied with merely
repeating the views which I have elsewhere, and at various times,
expressed on this subject, and which, to many of you at least, are
sufficiently well known; therefore, it seemed necessary that I
should, for the purpose of these lectures, undertake new investiga-
tions, in order to throw light, if possible, upon some of the points
which are still in dispute. But to compress into practically a
few months of otherwise fully occupied time what might well be
the undivided labour of a long period has proved a difficult task,
and I have fallen far short of what I had hoped to accomplish,
though I trust that some of the results at which I have been able
to arrive may contribute towards a solution of some of the vexed
questions. I purpose in these lectures to sketch the evolution of
the doctrine of cerebral localisation, to indicate the principal data
on which it is baaed, and to discuss, in the light of the most re-


cent investigations, the evidence for and against the existence of
specific centres, and their exact position in the cerebral cortex.

Before considering the facts bearing directly upon the specific
localisation of function in the cerebral cortex, I think it advisable
— nay, even necessary — to consider the effects of ablation of the
cerebral hemispheres in different classes of animals. A due con-
sideration of these phenomena affords, I think, a satisfactory ex-
planation of the chief objections which have been urged against
localisation in general, and, at the same time, also renders un-
necessary certain hypotheses as to the functional substitution of
one part of the cortex by another, which have been — and, in my
opinion, rightly — regarded by the opponents of localisation as
altogether subversive of its fundamental principles.

Recent researches on the effects of the removal of the cerebral
hemispheres, by improved methods, have necessitated some im-
portant modifications of the doctrines which, up to quite a recent
date, have been generally entertained on the subject.

Let us begin with fishes. When in osseous fishes the ganglia
(which correspond morphologically to the cerebral hemispheres of
the vertebrates) are entirely removed, there is little, if anything,
to distinguish them from perfectly normal animals. They main-
tain their natural attitude, and use their tails and fins in swim-
ming with the same vigour and precision as before. It has
generally been said that brainless fishes possess no spontaneity,
but seem as if impelled by some irresistible impulse (occasioned
by the impressions communicated to the surface of their bodies
by the water in which they are sustained) to swim until they are
exhausted by pure neuro-mupcular fatigue. In their course, how-
ever, as was shown by Vulpian, they do not blindly rush against
obstacles, but turn to the right or left, according to circumstances,
as if still possessed of some sense of vision. Vulpian says ^ " In
fact, when the cerebral hemispheres have been removed from a
fish which does not readily succumb tc this kind of operation (a
roach, for example), not only may it be urged to move by bringing
an object before its eyes, but I have proved that it avoids obstacles;
for, by placing a stick to the right or left, a few centimetres from
its eye, I have frequently caused the fish to turn in the opposite

Steiner ^ does not admit the absence of spontaneity in fishes so
operated upon, for he has seen that they occasionally remain at

1 SysUme Nerveux, p. 669.
2 Die Functionen des Centralnervensy stems ; Zweite Abtheilung : Die Fische, 1888.


the bottom, at other times balance themselves at various heights
in the water, and now and then swim about freely, without any
obvious alterations in the conditions by which they are surrounded.
He has also shown, and in this he has been confirmed by Vulpian,^
that they not only see, but are able to find, their food. If worms
be thrown into the water in which they are swimming, they im-
mediately pounce upon them. If a piece of string similar in size
to a worm be thrown in, they are able to detect the difference, and
either disregard it entirely, or drop it after having seized it. Not
only do they seize their food, but they discriminate between
different kinds, selecting some, and rejecting others. They even
to some extent distinguish colours, for when one red and a few
white wafers are thrown into the water, the fish almost invariably
selects the red in preference to the white.

From these facts it would appear that the fish without cerebral
hemispheres can see, distinguish colours to some extent, catch its
prey, discriminate between different kinds of food, direct its
movements with precision, and, in fact, behave to all app'-arance
like a normal animal. The only difference observed by Steiner
was that brainless fishes appeared more impulsive and less cautious
than those which had not been operated upon.

What has been said above applies, however, only to Teleosteous
fishes. Qaite different results appear to follow removal of the
cerebral hemispheres in Elasmobranchs. Thus the dog-fish, ac-
cording to Steiner,* after this operation is entirely deprived of
spontaneity, and is quite unable to find the food (sardines) by
which it is surrounded. The difference between the two orders
of fishes is, however, more apparent than real, for the dog-fish is
guided mainly by its sense of smell, while the activity of the
osseous fish is conditioned more especially by vision; hence, in
the dog-fish, removal of the cerebral hemispheres, which are
almost exclusively related to the sense of smell (Fig. 1.— a),
abolishes all the reactions conditioned by this sense ; while in the
osseous fish, the primary visual centres (optic lobes), being intact,
the ordinary modes of activity, which are conditioned mainly by
the eyes, continue to all appearance unmodified.

Frogs. — According to the researches, more particularly of Goltz'
and Steiner,^ frogs deprived of their cerebral hemispheres behave,

3 Co/nptes Rendus. Tome 102 and 103, 1886.

* Op. cit.
5 Functionen der Nervencentren des Frosches, 1869.
6 Physiologic des Froschhirns. 1885.
B 2


cceteris paribus, essentially like fishes similarly treated; they
maintain their normal attitude, and resist all attempts to over-
throw their equilibrium. If laid on their backs they will turn
over and attempt to regain their ordinary position. If the basis
of support on which they rest is tilted in any direction, they will
clamber up, forwards, or backwards until they gain a position of
stability. Their powers of locomotion are retained, and their
limbs are co-ordinated with precision. If a foot be pinched, or
any irritation applied to the posterior part of the body, they will
hop away ; thrown into the water they will swim, and continue
swimming until they have reached the side of the vessel, up
which, if possible, they will clamber and rest in peace. It would,
in fact, be difficult, so far as their movements and response to

Fig. 1.— Brain of dog-fish (after Steiner). A, cerebral hemisphere ; B, optic
lobe ; c, cerebellum ; o, olfactory lobe.

peripheral stimuli are concerned, to distinguish between a normal
and a brainless frog. If the back be gently stroked the frog will
answer uniformly with a croak, as if of pleasure or enjoyment.
If the animal be put in a vessel containing water, the temperature
of which is gradually raised, it will jump out as soon as the bath
becomes uncomfortably hot. If placed at the bottom of a pail of
water, it will ascend to the surface to breathe. If the vessel be
inverted over a pneumatic trough and filled with water, sustained
by barometric pressure, the frog will ascend to the top as before,
but not finding there the oxygen necessary to satisfy its respira-
tory craving, it will work its way downwards, and ultimately
succeed in making its escape out of the vessel on to the free sur-
face of the trough. Like the fish, the brainless frog undoubtedly
possesses some form of vision ; it does not, when urged to move,
rush blindly against an obstacle, but will leap over it, or turn to


the right or left, or otherwise avoid it. In all these respects a
brainless frog behaves like a normal one, but one noteworthy-
difference has been signalised by most observers— namely, that
the brainless frog, unless disturbed by some form of peripheral
stimulus, will remain for ever quiet on the same spot, until, in
fact, it becomes dried up and converted into a mummy. All
spontaneity— that is, varied activity under apparently the same
external conditions — appears to be annihilated ; its past expe-
rience has been blotted out, and it -views with indifference signs
and threats which would formerly have made it fl^^e. It is also
generally stated that the brainless frog has lost its instincts of
self-preservation, and either feels no hunger or possesses no power
to satisfy its physical necessities, so that it dies in the midst of
plenty. The more recent experiments, however, of Schrader'''
would seem to show that removal of the hemispheres deprives the
frog neither of spontaneity, nor of special instinc'^s, nor of the
ability to feed itself ; for he has observed brainless frogs which
have been kept alive for long periods, apparently "sponta-
neously " jump from the pier of a galvanometer, absolutely free
from all tendency to vibration, alternate between land and water
in the aquarium, crawl under stones, or bury themselves in the
earth at the beginning of winter, and, when cautiously submerged
under water, begin to swim exactly like normal frogs under the
same conditions. These frogs also, after the period of hyberna-
tion, or in the summer, when their wounds were entirely healed,
diligently caught the flies that were buzzing about in the vessels
in which they were kept. It would appear, therefore, if these
observations are correct, that the principal points of distinction
between the brainless and the normal frog — namely, the absence
of spontaneity and the power to feed itself, which are said to
especially characterise the former — are no longer capable of being
upheld, and that the brainless frog behaves precisely like the
brainless fish above described.

Birds. — Let us now proceed to consider the effects of the re-
moval of the cerebral hemispheres in birds, the next higher class
of vertebrates, and more especially in pigeons. These have been
rendered familiar to all by the classical researches of Plourens f
but though the picture he has drawn has been accepted as in the
main correct, there have been, and there still are, some differences
of opinion as to the facts, and more particularly as to their mode

7 Physiologic des Froschgehirns, Pfldger's Archiv fiir Pkysiologie, 1887, Band 41
8 Systeme Nerveux, 1842.


of interpretation. There is, however, no doubt that after this
operation pigeons show no disturbance of station or locomotion.
They maintain their normal attitude, and resist all attempts to
overthrow their balance. Left to themselves they appear, at first
at least, to be plunged in profound sleep. From this condition
they are easily aroused by a gentle push or pinch. When so
urged they march forwards, and should they happen to step over
the edge of the table, on which they are placed, they will flap
their wings and regain their base of support. Thrown into the
air they fly with all due precision and co-ordination. After each
manifestation of activity so induced they subside into their
original state of repose. Occasionally, and apparently without
any external stimulus, they may look up and yawn, shake them-
selves, dress their feathers with their beaks, move a few steps
forwards or backwards, especially after defsecation, and then settle
down quietly, standing sometimes on one leg, sometimes on the
other. They are altogether unable to feed themselves ; but, if fed
artificially, deglutition, digestion, and nutrition go on in a normal
manner, and the animals may be kept alive for an indefinite
period. Flourens was of opinion that the removal of the cerebral
hemispheres annihilated all the senses, and rendered the animals
blind, deaf, and devoid of smell, taste, and tactile sensibility.
These conclusions were, however, disputed by Magendie, Bouillaud,
Cuvier, and, in particular, by Longet^ and Yulpian.^" Longet found
that the animals appeared to see, inasmuch as they would follow
the movements of a flame held in front of their eyes at a sufii-
cient distance to prevent all sensation of heat, and also when
urged to move, occasionally at least, avoided obstacles in their
path. Also they started at loud sounds, such as a pistol shot,
made in their immediate vicinity ; and from their movements and
gestures appeared to feel impressions made upon the nerves of
common sensation. As regards the senses of taste and smell, Le
found it impossible to arrive at any definite conclusions in animals
of this order, and looked upon the statements of Flourens as not
supported by convincing evidence. Longet believed that the re-
moval of the cerebral hemispheres annihilated only perception
proper, as distinct from crude or brute sensation, which had its
centre in the mesencephalic ganglia.

The question as to the sense of sight in brainless pigeons has
been much discussed, that is, whether not mere impressionability

» Anatomie et Physiologie du Systeme Nerveux, 1842.
10 Op. cit.


to light exists, but as to whether the animals see, in the sense of
being able to guide their movements in accordance "with their
retinal impressions. McKendrick^^ was of opinion that removal
of the one cerebral hemisphere caused blindness in the opposite
eye; and Jastrowitz,^^ from his own experiments, arrived at the
same conclusion (on this see further below). The experiments of
Blaschko,^^ under the direction of Munk, led to no very definite
conclusions on this point, though it seemed as if the removal of
the one hemisphere did not cause total blindness in the opposite
eye. But Munk himself^* has made it the subject of a consider-
able number of experiments. He found that in a certain number
of pigeons, from which he had attempted to remove the cerebral
hemispheres, vision was not entirely abolished, and the animals
were able to avoid obstacles placed in their path. Careful investi-
gations, however {post mortem) revealed the fact that in such
cases the hemispheres had not been entirely destroyed, vision
continuing to some extent in the eye opposite the hemisphere, the
extirpation of which had not been absolutely complete. In those
cases, however, in which not a trace of either hemisphere was
allowed to remain, blindness was complete and absolute. These
animals, in their attitude and reaction to peripheral stimuli,
etc., exhibited the symptoms already described. The brightest
light, however, caused no result beyond contraction of the pupil.
The animals, when urged to move, ran against every obstacle
which came in their way. When thrown into the air, they flew
with retracted head and half-raised trunk, outstretched legs, and
dashed against obstacles, or fell bump on the ground and slid a
considerable distance before coming to a standstill.

The phenomena described by Munk certainly indicate total
blindness on the part of his pigeons, and he is of opinion that all
those who have held that ablation of the cerebral hemispheres
does not cause total blindness are in error, owing to the fact of
the extirpation of the hemispheres not having been complete.
Schrader, however,^' describes the phenomena which he observed
in two pigeons, from which, according to the post-mortem exami-
nation of von Recklinghausen, he had entirely removed every por-

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Online LibraryDavid FerrierThe Croonian lectures on cerebral localisation → online text (page 1 of 14)