Experimental Researches applied to Physiology and Pathology.
By E. Brown-Sequard, M. D., of Paris.
(Continued.)
XXVIII.—INFLUENCE OF RED BLOOD ON MUSCLES AND NERVES
DEPRIVED OF THEIR VITAL PROPERTIES.
James Phillips Kay* has found that blood, injected into limbs
of dead animals, just after irritability has disappeared, is capable
of regenerating this vital property. I have gone much farther, and
have discovered that blood is able to regenerate the vital proper-
ties of nerves and muscles, even in limbs which have lost their
irritability and have been rigid for several hours. I have ob-
tained this result from the following experiments :
’‘Treatise on Asphyxia. London, 1834
1st. On the body of a rabbit, in which cadaveric rigidity had
already existed for 10, 20, and in one case, 33 minutes, I
divided the aorta and the vena cava in the abdomen, immediately
above the bifurcation of these vessels. By means of small tubes,
a communication was established between their peripheric ex-
tremity and the central extremity of the corresponding vessels
divided in a living rabbit. The blood of this living animal
circulated immediately in the posterior limbs of the dead one.
After about six, eight or ten minutes, rigidity disappeared,
and, a few minutes afterwards, movements took place when
I excited the muscles or the muscular nerves.
2d. I have obtained a like result from an experiment more
easily made than the preceding, and which I have performed
more frequently. I divided transversely the body of a living
guinea-pig, or rabbit, into two halves, on a level with the lower
border of the kidneys, leaving no communication between the
two halves, except by the aorta and the vena cava. I then tied
the aorta immediately below the origin of the renal arteries.
The muscular irritability gradually diminished, and in a very
variable length of time* it gave way to cadaveric rigidity. I
waited until rigidity had been fully developed in all the muscles,
and then the ligature was relaxed and the circulation re-es-
tablished. Rigidity disappeared slowly, and the muscles and
the motor nerves resumed their vital properties.
* Sometimes 30, 20, or even only 10 minutes in weak animals, and from
1 to 8 or 9 hours in strong animals.
3d. In order to ascertain if voluntary movements and sensi-
bility could be restored to limbs that had been in a state of ca-
daveric rigidity, I tied the aorta immediately behind the origin
of the renal arteries, in several rabbits. Shortly afterwards,
sensibility and the voluntary movements disappeared in the
posterior limbs. I waited until muscular irritability had given
way to what is called cadaveric rigidity ; and when that peculiar
rigidity had existed for at least twenty minutes, I relaxed th
ligature. Then circulation took place, and, in consequence of it,
sensibility and voluntary movements re-appeared.
From this experiment it results, that not only local life, but all
the properties and actions of full life, can be restored in limbs
that have been in the state called rigor mortis, cadaveric or
post-mortem rigidity.
4th. On a man, 20 years old, who was guillotined on the
18th of June, 1851, in Paris, I made an experiment similar
to some of the preceding. The decapitation took place at
8 o’clock A. M. Ten hours afterwards, i. e. ten minutes after
6 o’clock P. M., the muscles of the hand, upon which I intended
to experiment, exhibited some slight manifestations of irritability.
At 7 and at 7 ’ o’clock P. M. I ascertained that they had lost
their irritability. Shortly after they were in a state of cadaveric
rigidity.
I began the injection of blood 10 minutes after 9 o’clock P. M.
As I wished to inject fresh human blood, and as I could not
obtain any from the hospitals at such an hour, I was obliged to
make use of my own. My friends, Drs. F. Bonnefin and Des-
lauriers, drew from one of the veins of my left arm half a pound
of blood, which was immediately beaten and completely defibri-
nated and filtered through a cloth.
As, in opposition to the general opinion, I had found that it is
not necessary, in transfusion, to make use of blood at a tempera-
ture not far from that of warm-blood animals. I left the blood
employed in this experiment freely exposed to the atmosphere
during all the time of the operation. The temperature of the
air was 19° centigr. (66°-2 Fahr.) I regret not having taken
the temperature of the blood when I began to inject it, but it
was probably about the same as that of the atmosphere.
The injection was made into the radial artery, a little above
the wrist. The whole quantity of the blood was injected in about
8 or 10 minutes. The arm operated on had been separated from
the body, and the blood injected came out from all the divided
arteries and veins.
Having saved nearly all the blood which flowed from these
vessels, I injected it anew. The last injection was made 45
minutes after 9 o’clock P. M. Ten minutes afterwards I found
that cadaveric rigidity had ceased in the hand, and that two
muscles only, out of the nineteen existing in that part, had not
resumed their irritability. Three muscles had become so very
irritable that a slight mechanical excitation was followed by a
contraction in the whole length of their fibres.
At half past one o’clock A. M.,—seventeen hours and a half
after decapitation and four hours after the injection of blood,—
there was still a slight irritability in the muscles of the hand.
In this experiment I found that half a pound of defibri-
nated human blood was sufficient to give irritability, for seve-
ral hours, to seventeen of the muscles of a hand.*
• For a full account of the circumstances of this experiment, see my paper
in the Gaz. Medic.de Paris, t. vi.—1851, p. 421.
5th. An experiment on another guillotined man gave me
more interesting results. The decapitation had taken place
at 8 o’clock A. M. on the 12th of July, 1851. At 51 o’clock
P. M., cadaveric rigidity existed in almost all the muscles of
the arms and fore-arms. I separated them from the body, and
at 61 o’clock I ascertained that cadaveric rigidity was increased,
and that only a few muscles were still slightly irritable. At 8
o’clock P. M. (12 hours after the decapitation) the muscles of the
two arms were completely deprived of irritability, and in full rigi-
dity, and the muscles of the forearms contracted only locally under
the influence of a mechanical irritation, and not at all when ex-
cited by a powerful magneto-electric current. Two other exami-
nations made, one at 91 and the other at 10 o’clock, gave the
same results.
At 10| o’clock two or three bundles of fibres of one of the
muscles of the fore-arm were the only parts where a mechanical
excitation produced a slight local contraction. All the other
muscles were perfectly stiff and deprived of irritability.
Twenty-five minutes after 10 o’clock there was no appearance
of irritability remaining in any muscle.
I then began the preparations for the injection of blood, with
the assistance of Drs. Martin-Magron, F. Bonnefin, Crouzet, and
Mr. Moyse.
We drew about a pound of blood from the carotid of a strong
dog. The blood was beaten and defibrinated before coagulation
could take place in it, and 10 minutes after 11 o’clock the injec-
tion was begun. It w'as made in the brachial artery of the left
arm, in the middle of its length, where the arm had been ampu-
tated. As soon as the blood had been thrown in the artery,
some reddish spots appeared in different parts of the skin of the
fore-arm, of the hand, and more particularly of the wrist. These
spots became larger and larger, and the skin had the appearance
it has in rubeola. Soon after, the whole surface of the skin was
of a violet reddish hue. In a few minutes this color disappeared,
and was replaced by the natural hue of the skin during life. The
skin became elastic and soft, as in a living man, and we saw the
bulbs of its hair becoming erected and presenting the appearance
called cutis anserina. By increasing and diminishing alternately
the impulsion given to the blood, we succeeded in producing the
beatings of the pulse in the radial artery. The veins were dis-
tinct and full as during life.
A short time after, the fingers, which had been extremely stiff,
relaxed, and rigidity disappeared also in the other parts of the
limb.
Forty-five minutes after 11 o’clock P. M., irritability had re-
turned in all the muscles of the limb operated on. The degree
of irritability, more particularly in the muscles of the arm, (tri-
ceps, biceps and others) was very considerable, and much greater
than I had seen it at the time the corpse was first examined
(about five o’clock P. M.) Irritability was still present in almost
all the muscles of this limb at 4 o’clock A. M., (20 hours after
the decapitation,) when I was obliged, from extreme fatigue, to
abandon further investigation.
The blood injected was at 23° centig. (73°.4 Fahr.) when I
began the operation, and the atmosphere was at 191° centig.
(66°.66 Fahr.)
In this experiment, about one pound of defibrinated dog’s
blood gave irritability for more than five hours to all the
muscles of a limb, from the middle of the arm to the hand.
6th. Every one knows the singular fact, that Vibriones and
other Infusoria, when desiccated, will live when they are put into
water. It is also perfectly known that seeds, after many centu-
ries, may grow when put in the earth. I have found something
of the same kind in higher organisms; it is that muscles, in a
certain condition, after having been separated from the body
for many days, may recover their irritability.
Dr. Coze, of Strasbourg, has found that chloroform injected
into the main artery of a limb produces instantly the strongest
rigidity, and that if blood is allowed to circulate again in the
limb, life appears again in it. I have gone farther, and found
that if a limb, in which an injection of chloroform has been made,
is separated from the body, it is able, under the influence of an
injection of blood, to recover its muscular irritability 2, 3, 4, 5
knd (in one case) 10 days after the rigidity was produced.
I think Mr. Edouard Robin is right in admitting that chloroform
prevents the chemical changes that take place in organic bodies
after death, and, if it is so, we can understand why an injection
of blood made so long after the limb has been separated from the
body, may reproduce irritability. One day is not more than one
hour, if, during it, there is no alteration produced in the muscles.
It appears, nevertheless, that chloroform does not entirely
prevent the alterations of muscles, because, in my experiments, I
have found that the longer the limbs had been separated from
the body, the greater was the quantity of blood necessary to re-
produce irritability.
7th. I lately made an experiment, with the view of ascertain-
ing how long a limb, separated from the body of an animal, may
be kept alive by means of injected blood. I succeeded in
retaining local life in one of the limbs of a rabbit more than
41 hours. The animal was a very vigorous, full grown one. I
killed it by hemorrhage, and, two hours afterwards, rigidity had
begun in most of the muscles of the two posterior limbs, and only
a few bundles of muscular fibres had still a slight irritability. A
first injection of defibrinated blood was then pushed in the femo-
ral artery of the right posterior limb. Fifteen minutes after the
beginning of the injection, local life (i. e. irritability) was re-
stored in the limb receiving blood, and cadaveric rigidity had
disappeared.
The manner of testing this irritability was the same as that of
Glisson, Gorter, and all the experimenters of the two last centu-
ries,—I mean by mechanical excitation. I did not use galvanism,
as it exhausts muscular irritability too much, as Autenrieth,
Pfaff and many other observers have shown long ago. Being
aware of this fact, I have always, in my preceding experiments,
made use of galvanism for a very short time only.
Three hours after the death of the rabbit, irritability still
existed in the right limb (the injected one,) while the left was
perfectly rigid and had not the slightest irritability. Half an
hour later, rigidity had begun again in the right limb ; blood
was injected anew, rigidity disappeared, and local life returned.
From this moment until 11 o’clock, P. M., (death had occurred
at 6 o’clock, A. M. of the same day,) blood was injected many
times. Rigidity did not return, and the vital property of the
muscles was maintained. Of course the left limb, during that
time, remained rigid, and had not the slightest irritability.
From 11 o’clock, P. M. until 6 o’clock, A. M. the succeeding
day, an abundant injection of blood was made every twenty or
twenty-five minutes. The irritability was not powerful, but it
existed in all the muscles of the limb. There was no rigidity
at all.
The injections were then made more frequently—once in each
quarter of an hour—until three o’clock, P. M., at which time I
was obliged to stop them for an hour and a half.
At half past four I found the limb rigid, and only a few
bundles of muscular fibres still irritable. A very abundant
injection was then practised, and rigidity soon disappeared,
giving way to irritability. From this time to 11 P. M., a great
many injections were made, and irritability was maintained. I
was then obliged to give up the experiment. At that moment
irritability was strong in all the muscles of the injected limb,
except some parts of their pelvic extremities that had not
received a sufficient quantity of blood.
The next morning that limb was in full and energetic rigidity.
The other limb had already lost its rigidity, and had an evident
smell of putrefaction. The third day after the death of the
animal, rigidity was strong in the injected limb, while the other
was in an advanced state of putrefaction.
If we compare these two limbs, we find, 1, That the injected
one had a strong irritability at the end of forty-one hours after
the death of the animal; 2, That its rigidity gave way to putre-
faction only at the eightieth hour; 3, That it was in complete
putrefaction only at the ninety-fourth hour. The other limb
■was in full rigidity at the fifth hour after the death of the
animal; its rigidity gave way to putrefaction at the forty-
eighth hour; and it was in complete putrefaction at the seven-
tieth hour.
From all the experiments above related, it appears that life
may be reproduced or maintained in muscles and nerves by
mere injections of blood. I have found, also, that life may be
rep oduced by the same means in the spinal cord and in the
brain. I will publish these facts in another article.
It is nearly indifferent in these experiments whether we use
venous or arterial blood ; but it is absolutely necessary to employ
red blood, i. e. oxygenated blood.
I have tried, sometimes, arterial blood, rendered black by
the substitution of nitrogen or hydrogen for a great part of its
oxygen, and I have found that such blood was unable to repro-
duce the vital properties of nerves and muscles.
Oxygen is necessary, either because it prevents the blood-glo-
bules from being altered, or because it acts directly on muscles, as
Gustavus Liebig has found it does on their external surface,
when exposed to air. I believe it is necessary for both these rea-
sons.
I cannot say how long after the beginning of cadaveric
rigidity in a muscle, oxygenated blood can reproduce local life.
In the second of the two decapitated men, on whom I ex-
perimented, rigidity had existed at least five hours before the
injection was begun. I believe that the stronger the animal is,
the more easy it is to reproduce local life in rigid limbs, by
injection of blood. In limbs of weak rabbits, I have found it
impossible, two hours after the beginning of cadaveric rigidity,
to reproduce local life. In a very strong dog I have reproduced
muscular irritability four hours after rigidity had been fully
developed.
Ten, twelve, or fourteen hours after rigidity had taken place,
in human limbs, I have tried in vain to re-establish local life.
I have ascertained that pure serum of blood, or milk, or albu-
men of eggs, are unable to produce any apparent change in
rigid limbs.
The following conclusions are to be drawn from the facts
related in this article :
1st. Red blood, i. e. richly oxygenated blood (arterial or
venous) is able to revive irritability in muscles, four or five hours
after these organs have lost this property.
2d. Red blood is able to revive the vital properties of
nerves and nervous centres, when these properties have not been
lost for more than about an hour.
3d. Muscular irritability can be maintained for more than 41
hours, by mere injections of blood, in limbs separated from the
body of a rabbit.
4th. Muscular irritability may be re-established in limbs ren-
dered rigid by chloroform for many days, even ten days.
XXIX____CASES OF LOSS OF SENSIBILITY ON ONE SIDE OF THE BODY.
AND LOSS OF VOLUNTARY MOVEMENTS ON THE OTHER SIDE.
It has been objected to me that if the transmission of sensitive
impressions, in the spinal cord, takes place, as I have tried to
prove in a former part of this sketch (Art. XIX,) so that those
coming from the left side of the body, are mostly conveyed to
the sensorium along the right side of the spinal cord,—et vice
versa—physicians should have some times found in man the
same thing that I have discovered in animals.
Many reasons have prevented physicians from making such a
discovery: In the first place, an injury or a pathological altera-
tion, limited to a lateral half of the spinal cord, is very rare.
Besides, the idea that there is no crossing of fibres in the spinal
cord, has been an obstacle to a thorough examination of many
pathological cases, and it has been so in a case observed by Boyer.
There are but few cases on record in which there was a loss
or a diminution of sensibility on one side, and of voluntary
movements on the other. I will give here a short account of
some cases of that kind, which are very interesting.
The first one I will relate has been observed by Boyer:
A drummer, of the National Guard of Paris, received a wound
in the back of his neck. A sword had been thrown at him, and
had penetrated the superior part of the right lateral half of the
neck. An incomplete paralysis of movement took place in the
right side of the body, and, some time after, it was accidentally
discovered that sensibility was lost in many parts of the left
side of the body. After twenty days the wound was cured, and
the man went out of the hospital, still paralysed.
From what we know of that case, it appears that the sword had
incompletely divided the right lateral half of the spinal cord.
The paralysis of motion on the right side of the body was
certainly produced by the division of a part of the anterior
column, and, as the instrument had penetrated the right side of
the back of the neck, it must have divided the parts between the
anterior column of the spinal marrow and the external surface of
the right side of the neck. These parts, besides the muscles and
bones, are the lateral and posterior columns and the gray matter
of the right half of the spinal cord. So that in this case nearly
the same injury and also the same morbid phenomena had ex-
isted as in the animals on which I have divided a lateral half of
the spinal cord.
The following case is still more interesting. It has been re-
corded by Dr. R. Dundas, Surgeon of the Hospital of Bahia.
A mason fell on his back from an height of 20 feet. After
having recovered his consciousness, he discovered that all the
left side of his body, from the shoulder to the foot, was paralyzed
as to motion, without the slightest alteration of sensibility, and
that the right side in which the movements were free, was com-
pletely deprived of sensibility.
Three important facts, precisely like those I have discovered
in animals after the transversal section of a lateral half of the
spinal cord, existed in this case :
1st. A morbid exaltation of sensibility in the side where move-
ment was lost.
2d. A diminution of temperature in the side where the para-
lysis of sensibility existed.
3d. An increase in temperature in the side where the paralysis
of movement existed.*
* In a former part of this sketch (Art. xxii.) I have related facts proving
that animal heat may be increased after injuries to the spinal cord. I have
learned since, that Prof. D. Gilbert has observed a case of fracture of the
spine, in which the temperature of the paralyzed parts was increased. Prof.
Dunglison has also stated that the paralyzed side in hemiplegic patients
may have an elevation of temperature.
When Dr. Dundas published this curious case, the patient was
living and improving ; so we do not know what was the altera-
tion existing in the spinal cord.
II. Ley, in a letter to Sir Charles Bell, relates the following
case:t
■[The nervous system of the human body. By Ch. Bell. 3d ed. London,
1844, p. 245.
Mrs. W., after a profuse hemorrhage, became paralytic. Upon
one side of the body there was a loss of sensibility, without, how-
ever, any corresponding diminution of power in the muscles of
volition. The breast, too, upon that side, partook of the insen-
sibility, although the secretion of milk was as copious as in the
other. She could see the child sucking and swallowing, but she
had no consciousness, from feeling, that the child was so occu-
pied.
Upon the opposite side of the body there was defective power
of motion, without, however, any diminution of sensibility. The
arm was incapable of supporting the child ; the hand was power-
less in its grasp ; and the leg was moved with difficulty, and with
the ordinary rotatory movement of a paralytic patient; but the
power of sensation was so far from being impaired that she con-
stantly complained of an uncomfortable sense of heat, a painful
tingling, and more than the usual degree of uneasiness from
pressure, or other modes of slight mechanical violence.
She again proved pregnant. Iler delivery was easy : but after
about ten days she complained of numbness on both sides.
Her articulation was indistinct; she became more and more in-
sensible, and sank, completely comatose.
No positive disorganization of the brain could be detected.
The ventricles, however, contained more than usual serum ; and
there were found thickening and increased vascularity of the
membranes, with moderately firm adhesion in some parts ; in
others, an apparently gelatinous, transparent and colorless de-
posit interposed between them.
Unfortunately, no examination of the spinal cord was made.
In this case there was very likely, as in my experiments, an
increase in the temperature of the side paralyzed of motion. The
writer merely says that the patient was constantly complaining
of an uncomfortable sense of heat. There was, as in my animals,
an evident increase in sensibility on that side.
M. Monod* has related the case of a man who, after having
felt a sudden pain in his back, became paralyzed in the motion
of the right inferior limb. Sensibility was entire on this side,
but on the left side, where the movements were entire, sensibility
was entirely lost from the breast to the foot. There was at first
no fever. The patient died 34 days after the beginning of this
affection.
* Bulletin de la Societe Anatorniqiie, No. xviii. p. 349.
The brain and its membranes were normal. A hemorrhage
had taken place, and blood was found in the right side of the
central gray matter, in the neighborhood of the anterior column
in the dorsal and lumbar regions.
This case is assuredly a very remarkable one, and in accord-
ance with my experiments.
The conclusion to be drawn from these four cases is, that in
man as well as in animals, there appears to be a crossing of the
sensitive nerve-fibres in the spinal cord.
XXX.—ON THE DIFFERENT DEGREES OF EXCITABILITY OF THE
DIFFERENT PARTS OF THE SENSITIVE NERVE-FIBRES.
It is a well-known fact, that an excitation of the skin or of a
mucous membrane, produces a greater pain or a greater reflex
action than that of the nerve trunk, from which these parts re-
ceive their nerve-fibres. For instance, a slight excitation of the
laryngeal mucous membrane produces coughing, while an excita-
tion of the vagus nerve very rarely produces the same effect.
Therefore, there is a notable difference between the peripheric
extremity of a nerve-tube and its part contained in a nerve-
trunk.
The existence of a peculiar organ in the skin (the corpuscles
of touch of Wagner) has not much (if it has anything) to do with
the different degrees of excitability of nerve-tubes in the skin
and in the trunks of nerves. The corpuscles of touch do not
exist in the mucous membranes, and if they exist in the skin of
frogs, turtles, etc., it is in a very small number ; and, neverthe-
less, the degree of excitability of nerve-fibres in these parts is
much superior to that of the fibres of the nerve-trunks.
Some very striking differences exist in the degree of excita-
bility of centripetal nerve-fibres in the five following different
parts of their length.
1st. The part contained in the skin.
2d. The part of a nerve extending from the skin to the spinal
cord.
3d. The posterior roots of the spinal nerves.
4th. The part of the posterior roots attached to the spinal
cord.
5th. The part of the cutaneous nerve-fibres contained in the
gray matter of the spinal cord.
The fibres existing in the gray matter of the spinal cord ap-
pear to be inexcitable, at least by our ordinary means of excita-
tion. Of the four other parts, the less excitable is the nerve be-
tween the ganglion and the skin. The excitability of the pos-
terior roots is less than that of the skin and that of their part
attached to the spinal cord. Of these two last parts the skin is
less excitable than the other.
I measured the excitability by the degree of pain or of reflex
action. The differences are much more easily found for the re-
flex action than for the pain.
Is it because they have been connected with the cells of the
central gray matter of the spinal cord, that the centripetal fibres,
contained in that gray matter, are not excitable? If it is so,
there is a difference between these cells and those of the gan-
glions on the posterior roots, because the connection of these fibres
with the cells of these ganglions does not prevent their being
excitable.*
* Recently, Dr. Cl. Bernard has communicated to the Societe de Biologie,
of Paris, a fact which would be very important if it were exact. He says
that some of the fibres of the posterior roots of the spinal nerve, in frogs,
do not pass through the ganglion,—that they are the sensitive fibres, and
that the ganglionic fibres are merely for reflex action. I have made, alone
or with my friend Dr. Henry Lolliot, the minutest examination of the spinal
ganglions, and I am satisfied that all the fibres of the posterior roots pass
in the ganglion, and that Dr. Bernard has been mistaken.
From the facts above related I conclude that the same nerve-
fibre, in different parts of its length, may have very different de-
grees of excitability.
(To be continued.)