PROCEEDINGS

OF THE

PHYSIOLOGICAL SOCIETY,
October 17, 1931.
Nerve endings in Pacinian bodies. By DONAL SHEEHAN (introduced
by Prof. J. S. B. STOPFORD). (Department of Anatomy, University
of Manchester.)
Preparations were shown, stained by the intra-vitam methylene
blue technique, demonstrating the different types of nerve-endings in
the Pacinian bodies in the mesentery of the cat. Fibres could be seen
passing straight through one Pacinian body to terminate in two adjacent
corpuscles.
After sub-diaphragmatic section of both vagi, the normal staining
of the nerve-endings was obtained three weeks later. Bilateral section
of the splanchnics, in a series of animals, gave a negative methylene
blue result for the Pacinian bodies. Control experiments were carried
out in every case. From experimental work it would thus appear that
the main afferent nerve fibre of the Pacinian corpuscle reaches its cell
station in the posterior root ganglion via the splanchnic nerves.
Work is still in progress on a fine plexus of un-myelenated fibres
with nerve ganglion cells and occasional free nerve-endings, found
throughout the mesentery.
Provisional results show that this plexus is in connection with the network of un-myelenated fibres around the blood vessels. After bilateral
section of the splanchnics, and also after division of both vagi below
the diaphragm, the fibres of this fine plexus could still be traced, though
much reduced in number. A further series of experiments has been
carried out to exclude the possible source of afferent innervation to
the mesentery from the upper lumbar sympathetic ganglia.
There appears to be some definite histological evidence underlying
the assumed somatic innervation of the mesentery, but these results
await confirmation from present experimental work.

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PROCEEDINGS OF THE PHYSIOLOGICAL
The nervous origin ofthe Wever and Bray effect. By E. D. ADRIAN,
D. W. BRONK and GILBERT PHILLIPS1.
One of us [Adrian, 1931] recently criticized Wever and Bray's interpretation of the potential changes which they found in the auditory nerve
when sounds fall on the ear. These changes follow the content of the
sound waves so closely that, after amplification, they can be made to
reproduce speech as well as notes of high and low pitch. We v er and B r a y
give reasons for supposing that the effect is due to true action potentials
in the auditory nerve, but on repeating their experiments it was found that
the effect could always be detected in the neighbourhood of the cochlea
and that a lead from the nerve was unnecessary. Since the application of
novocaine etc. to the nerve did not affect the electric changes appearing in
it, it was concluded that these were generated within the cochlea and
were not due to impulses in the intracranial nerve fibres.
Further experiments have now been made and the objections raised
to Wever and Bray's explanation turn out to be not very serious.
It seems probable that the changes in the cochlea are due to nervous
structures of some kind: the situation of the nerve fibres in it may
favour the development of large potentials, and the intracranial part of
the nerve may often appear to be acting as an inert conductor because it
is unfavourably placed for electric recording and much more liable to
damage than the intracochlear part.
The experiments which favour a nervous origin are as follows:
Survival period. When the bulla of the guinea-pig is opened and a
moist thread electrode placed on the cochlear projection we find that
sounds are still reproduced for an hour or more after the death of the
animal or the failure of the blood supply to the inner ear. When the
circulation fails there is a sudden fall in intensity, most pronounced for
sharp isolated sounds like the clicking of a pair of forceps. These usually
become quite inaudible, but sustained sounds are less affected: speech is
still quite intelligible, and after the initial reduction its intensity remains
at about the same level for 1-14 hours. Sounds then become faint and
distorted and the effect fades rapidly. In the cat with a lead from the
foramen rotundum the survival period is about half an hour. This
result can be produced by cutting the vessels to the inner ear without
interfering with those to the middle ear. The prolonged survival at fairly
constant intensity suggests that the initial diminution is due to a
1 Fellow of the Rockefeller Foundation.

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SOCIETY, OCTOBER 17, 1931.

8P

mechanical cause, e.g. the collapse of vessels, and not to the failure of the
oxygen supply. The persistence of the effect in the absence of blood supply
does not rule out a nervous origin, for peripheral sense organs and nerves
may remain excitable for long periods after removal from the body.
The final disappearance makes it likely that the electric changes are due
to the activity of living cells and not to movements of fluid in the inner
ear.
Cocaine. The injection of 10-20 p.c. cocaine solution into the foramen
rotundum or through the wall of the guinea-pig's cochlea causes a rapid
failure of the effect. The injection of saline usually leaves it unimpaired if no great damage is done. Although the variable amount of
damage confuses the issue the effect of cocaine seems unmistakable.
Cooling the cochlea. It was reported previously that ice placed on the
petrous bone in the cat did not weaken the effect. In the guinea-pig,
however, the cochlea can be reached more easily from the bulla and
cooling it with ice causes a definite weakening. If the circulation is
intact there is complete recovery when the ice is removed, but if it is
applied during the survival period there is no recovery and the effect
soon fades completely.
These results all indicate that the Wever and Bray effect is due to
living cells and ceases with their death, and the action of cocaine makes it
likely that nerve endings, nerve cells or nerve fibres are responsible. The
large size of the potential changes in the cochlea suggests that the nerve
fibres are not the only structures giving rise to them, but it is true that
the synchronous activity of many fibres in a nerve can give large potentials, and it is possible that the arrangement of the bony channels
in the inner ear may give particularly favourable conditions for recording
electric effects. It is also worth remarking that preparations of the touch
receptors and their nerves can be made to give potential fluctuations
reproducing the frequency of the stimulus over a limited range, and it is
even possible to reproduce a sinusoidal wave form if enough receptors
are in action.
REFERENCES.
Adrian, E. D. (1931). J. Pkysiol. 71, 28 P.
Wever, E. G. and Bray, C. W. (1930). J. Exp. Psychol. 13, 373.

a2
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PROCEEDINGS OF THE PHYSIOLOGICAL

4P PP
The mechanics of Pecten muscle. By A. D. RITCHIE.
The isolated slow adductor muscle of Pecten opercularis stimulated
under isometric conditions takes several seconds to relax. If the muscle
is rapidly stretched and released again to the same length during relaxation, the rate of fall of tension is greatly increased. With a series
of stretches and releases the base line of tension is reached in a second or
two. The stretching and releasing does not appear to injure or permanently
alter the muscle in any way. Release followed by stretch has no effect.
The phenomenon is believed to be due to a momentary lowering of
viscosity by the mechanical shaking up of the tissue and to be comparable
to the lowering of viscosity of a "'thixotropic" gel [Freundlich, 1927],
which liquefies on shaking and sets again on standing.
If it is assumed that a similar thixotropic effect is produced in an
excited muscle fibre immediately after the excited state passes off, i.e.
while the fibre is relaxing, certain anomalies observed in Pecten muscle
can be explained. (1) Relaxation from a large contraction is quicker
than one from a small contraction, so that the relaxation curves are not
superposable [Bozler, 1930]. The greater the number of fibres that
have been excited, or the greater the contraction of any individual
fibre, if contractions are summated, the greater the thixotropic effect.
(2) The state of "contracture" found in the muscle immediately after
isolation is abolished by a strong stimulation [Bayliss, Boyland, and
Ritchie, 1930]; and also by stretch and release. (3) The intact animal's
reflex relaxation may be much more rapid than the relaxation obtained
in the same muscle after isolation [Bozler, 1930]. This is probably
because the contractions of the isolated muscle produced by ordinary
methods of stimulation are always submaximal and therefore produce
less thixotropic effect than will follow a maximal reflex contraction.
The thixotropic effect has been observed in the adductor muscle of
the oyster, and less conspicuously in the retractor of the foot of Mytilus
and in crab's claw muscle. In tortoise skeletal muscle its presence is
doubtful. The body wall muscle of Holothuria does not appear to show a
thixotropic effect, but the interpretation of the results obtained is doubtful, as a series of rapid stretches and releases often acts as a stimulus.
REFERENCES.
Bayliss, L. E., Boyland, E. and Ritchie, A. D. (1930). Proc. Roy. Soc. B, 106, 363.
Bozler, E. (1930). Z. vergl. Phy8iol. 12, 579.
Freundlich, H. (1927). Protop7.ma, 2, 278.

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SOCIETY, OCTOBER 17, 1931.

5P

Fibrillarytwitchings in skeletal muscle during the post-ischbmic

period. By CHARLES REID; (Prince of Wales Medical College, Patna,

India.)
Ischemnia of the hand and forearm was produced by application of
an Esmarch bandage round the hand and forearm. The armlet of a
blood-pressure recording apparatus was placed on the upper arm, the
pressure on which was then raised and maintained at about 20-30 mm.
Hg above the systolic level. On removal of the bandage the forearm
was ready for use as an ischaemic limb.
In the course of the second or third minute after the re-admission
of blood following ischaemias of from 15 to 35 minutes' duration, involuntary fibrillary contractions constantly appear in the muscles of the
hand and forearm, the small muscles being affected particularly, and
disappear usually in the course of the sixth or seventh minute after the
return of blood to the limb. Their intensity and duration vary to some
extent with the duration of the ischwemia. Evidently they are of peripheral origin, involving an excitatory process which constitutes a definite
phase in the behaviour of the muscles some minutes after the re-entry
of blood and are no doubt of the same nature as those observed in the
tongue, etc. of the dog, following experimental ischaomic conditions by
Guthrie, Pike and Stewart [1908].
So far the writer has not demonstrated similar fibrillary twitchings
in the limb muscles of the dog, cat or rabbit subsequent to ischaemic
conditions induced similarly as for man. In order to show, however, a
possible chemical basis for the origin-in part at least-of the above
abnormal excitatory process, the total calcium [D a v en p or t, etc., 1929]
of the post-ischamic muscle in animals was compared with that of the
normal. It was easily shown that the Ca contents of corresponding
muscles from the hind limbs or fore limbs of animals anaesthetized with
intra-peritoneal amytal were similar. Normal muscles were then compared as regards their Ca content with corresponding muscles after the
circulation had been re-established for about 5 minutes.
Animal
Rabbit (13)

Ca content (mg. p.c.)

Normal muscle
21

Post-ischamic muscle
16-7
18-2

22-2
Dog (7)
The Ca content of the blood [Tisdall, 1923] obtained from the deep
vein of a post-ischeemic limb (dog) was always slightly higher (less
than 1 mg.) than that of blood from the deep vein of a normal limb.

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6P6PIPROCEEDINGS OF THE PHYSIOLOGICAL

Ft A'ther evidence, suggesting that the onset and maintenance of the

post-i 3chaemic fibrillary process are associated with the setting in of the
periout of most active interchange between the muscles and the blood,
is afforded by serial estimations of the lactic acid content of samples of
blood [Cotonio, etc., 1927] proceeding from a limb in a period extending
for half an hour after the re-establishment of the circulation. The blood
lactic acid in a number of observations on different dogs quickly increased in the course of a few minutes from 8-10 mg. p.c. to 12-15 mg.
p.c. and then subsided slowly to about 9-11 mg. in 30 minutes.
The mechanism of these fibrillary twitchings is probably essentially
similar to those described by Langley [1915] for the dog's sartorius
placed in pure NaCl0 6 p.c. and then transferred to Ringer. The twitchings
set up cease in three to seven minutes and are considerably greater if
the Ca content of the Ringer is low.
REFERENCES.
Cotonio, M., Friedemann, T. E. and Shaffer, P. A. (1927). J. Biol. Chem. 73, 335.
Davenport, H. A., Dixon, H. H. and Ranson, S. W. (1929). J. Biol. Chem. 83, 737.
Guthrie, C. C., Pike, F. H. and Stewart, G. N. (1908). Amer. J. Physiol. 19, 17.
Langley, J. N. (1915). J. Phy8iol. 49, 29P.
Tisdall, F. F. (1923). J. Biol. Chem. 56, 439.

Cellobiose in relation to the mammary gland. By L. B. WINTER.

(Physiological Laboratory, Manchester.)
It is usually assumed that in the formation of lactose by the mammary
gland galactose is formed from glucose brought by the blood and then
linked with glucose to form a fl-galactoside, but all attempts to show
the presence of an enzyme which will synthesize lactose from glucose
and galactose in vitro have failed. Free galactose may not be formed
in the gland, and there are other possibilities of which the following is
an example. Two molecules of glucose may be joined to form a ,B-glucoside, and then the H and OH attached to C 4 of the non-reducing
component may be transposed. Such a change occurring in the glucose
/3-glucoside cellobiose will give rise to lactose directly, for these disaccharides have the same structure [Haworth, 1929]. In the present
work mammary glands of rabbits killed 10 days after parturition were
extracted and the extracts were incubated with cellobiose at 37. The
sugar was subsequently recovered from the mixture by the lead-barium
method [Winter, 1930]. Cellobiose octa-acetate was prepared by the
method of Haworth and Hirst [1921], and converted into the free
sugar by Zemplen's method [1926]. I take this opportunity of thanking
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7P
SOCIETY, OCTOBER 17, 1931.
Dr E. L. Hirst for showing me the preparation, as experience is necessary
to obtain satisfactory yields. Three methods of extracting the mammary
tissue were used:
(1) Dehydration with acetone, and extraction of the dry powder
with saline.
(2) Extraction with 50 p.c. glycerol.
(3) Direct saline extraction at low temperature (5C C.).
Each flask on incubation contained 10 c.c. of a 3 p.c. solution of
cellobiose, 8 c.c. 0*9 p.c. NaCl or 0*4 p.c. Na2HPO4, and 2 c.c. of " enzyme)"
solution. When incubation was for more than 3 hours, toluol was added.
The following are representative figures.
Method of
extraction
Acetone
Control
Glycerol
Control

Saline

[a] 5461 of the

recovered sugar

Time of
incubation

(hr.)

Initial

48
27-1
48
28-3
48
44.3
48
27-0
48
307
48
25-4
3
34*0
Control
3
25*0
t Contained sodium phosphate.

Final

39.4

40-0

63.0*

39.9
400t

41 Ot

409
41-0

The presence of a small amount of lactose in the recovered sugar

would show itself by raising the initial and final specific rotatory powers
above those of cellobiose, and by masking the mutarotation (since
lactose crystallizes as the a and cellobiose as the , compound at room
temperature). In no experiment were these three requirements satisfied,
and in only one (marked with an asterisk) was the final [a] raised; this
may well be due to splitting of some cellobiose to glucose by bacterial
action. Experiments in which cellobiose was injected into rabbits gave
no evidence that the sugar was utilized when the-animal was lactatinag.
Sex
Male
Female
Female (lactating)

Weight
(kg.)
27
3-1
3-0

Cellobiose
injected
(g.)
10
10
10

Extra
"sugar') in
urine (g.)
0 45
074
050

The sugar was excreted unchanged by the lactating rabbit; from

the above urine 0-19 g. was recovered with M.P. 222, and [a] 5461 = + 22*4.
REFERENCES.
Haworth, W. N. (1929). The Conatitution of Sugars. Arnold, London.
Haworth, W. N. and Hirst, E. L. (1921). J. Chem. Soc. 119, 193.
Winter, L. B. (1930). Biochem. J. 24, 851.
Zemplen, G. (1926). Ber. deuts. chem. Gme. 59, 1254.

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8P8PPROCEEDINGS OF THE PHYSIOLOGICAL

The precise anatomy of the Hensen's cells in the cochlea of
the guinea-pig and its physiological significance. By

C. S. HALLPIKE. (Ferens Institute of Otology, Middlesex Hospital.)

(Preliminary note.)
These cells situated to the outer side of the external group of hair
cells and associated Deiter's cells form part of the so-called supporting
elements found in Corti's organ.
The fact that these cells contain fat globules has been known for
many years. Retzius [1881] represents them as being first described

Fig. 1.

Fig. 2.

by Hensen. Details as to arrangement, however, are here, as elsewhere,

lacking. The arrangement found in young adult guinea-pigs is shown
in the appended camera lucida drawings of osmicated celloidin preparations (Figs. 1 and 2).
It may be described as follows: The fat globules are absent in the
basal coils, appear first half way round the second coil and show a
finely graded increase towards the apex where their bulk is considerable.
Anatomical differences in various parts of the cochlea have played an
important part in support of the Resonance Theory, in particular the

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9P
SOCIETY, OCTOBER 17, 1931.
observations of Hensen [1863] regarding differentiation in length
of the fibres of the basilar membrane, and of Gray [1900] regarding
differentiation in tension by the spiral ligament. Both appear to
indicate the presence of tuning mechanisms by which the apex of the
cochlea is adapted to the lower part of the scale. It is suggested that
the present observation indicates yet other possibilities, and that in the
guinea-pig at any rate the arrangement of the fat globules described
represents a graduated loading mechanism which works towards the
same end, i.e. tuning of the apical part of the cochlea to low tones.
The arrangement of fat globules described does not appear to be
present at birth although differentiation as regards fibre length in the
basilar membrane and tension in the spiral ligament are already accomplished. Thus ontogenetically the loading mechanism described
appears to be the final achievement in the development of tuning
mechanisms in the cochlea of the guinea-pig if the suggestion made
regarding the function of the fat globules is acceptable.
REFERENCES.
Gray, A. (1900). J. Anat. Lond. 34, 324.
Hensen, V. (1863). Z. Wl88. Zool. 13, 492.
Retzius, G. (1881). Das Geh6rorgan der Wirbeltiaere, n, 296.

Effect ofactivated charcoal on frog heart perfasate. By E. DAVIS

and S. J. FOLLEY. (Department of Physiology, University of

Manchester.)
A cannula was inserted into the posterior vena cava of the frog, the
heart was isolated, and perfused with Ringer solution through the
cannula. The same fluid was continuously re-circulated through the
heart. Hearts treated in this way would beat up to 36 hours. When
the contractions were nearly extinguished, activated wood charcoal
(" Klarit") was added to the Ringer, and circulated through the heart.
The beat was much accelerated and augmented. The contractions
resembled those of the fresh heart, and the heart continued beating for
2 to 21 hours. At the end of the experiments, the charcoal-Ringer solution
was collected, and the charcoal centrifuged off. The fluid remaining
stimulated the perfused frog heart and particularly weak hearts. The
charcoal was eluted with distilled water, Na2HPO4, and NaH2PO4, but

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10P PROCEEDINGS OF THE PHYS. SOC., OCTOBER 17, 1931.

the solutions obtained had but little effect upon the perfused heart.
Slight inhibitory effects were sometimes obtained.
After acetylcholine had been shaken with " Klarit" for three hours,
the inhibitory action of the drug on the heart was slightly reduced.
A frog heart, perfused with "Klarit" suspended in Ringer solution,
responded but feebly to acetylcholine.

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