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even in the case of the arm after the brachial plexus has been severed
(Fig. 91). In the frog it is produced also with the brain and spinal cord
completely destroyed, when salt solution containing suprarenal extract is
allowed to flow through the arteries. Under these circumstances the
flow of fluid, which, without the suprarenal extract, may have been com-
paratively rapid, becomes almost completely stopped, and this can only
be due to the direct action of the extractive substance upon the muscular
tissue of the smaller arteries.

The enormous rise of blood pressure. which is got after the vagi have
been cut, is shown in the tracings (Figs. 89 and 90) ; the pressure may rise
to four or five times its original height. Hardly any other agent will
produce such an enormous increase of pressure, except direct stimulation
of the vasomotor centre. It is not the case, however, that the elevation
of blood pressure, and the contraction of the arteries, is due to the
stimulation of the vasomotor centre by the drug, as was supposed by
Cybulski and Szymonowicz, for, as we have seen, the action is essentially
a peripheral one. As shown by Oliver,^ it will occur if the extract be
directly applied to the vessels of the mesentery, either during life or
in the " surviving " condition.

The effect of intravenous injectiou upon the blood pressure passes off in the
course of a few minutes. After a dose, no matter whether small or large, has
been injected into a vein, and has produced the results which we have
recorded, the blood vessels slowly resume their ordinary calibre, the augmenta-
tion and increased frequency of the heart's heats become gradually lessened,
and the blood pressure recovers its normal condition. Whilst the pressure is
raised under the action of suprarenal extract, there is apparently no possibility
of inhibiting the arterial contraction ; even the strongest stimulation of the
depressor nerve, which under ordinary circumstances produces through the
vasomotor centre a marked dilatation of the arterioles, is without result during
the activity of this extract. The question naturally arises, How is it that the
effect so soon disappears % In what manner is the active principle eliminated %
It is not eliminated by the kidneys, for the effect passes off just as quickly
even although the renal arteries are clamped. It is not eliminated by the
suprarenals themselves, for the same fact holds good for the suprarenals. It
passes off almost equally quickly if the aorta and vena cava are tied in the
upper part of the abdomen, so that there is no circulation of blood whatever in
the abdominal organs. It is not oxidised or otherwise destroyed by the blood,
for it retains its full potency even after it has been twenty-four hours in
contact with that fluid. The most probable explanation of the disappearance
of the effect seems to be that the active principle becomes packed away, and
eventually rendered innocuous in certain organs. That the muscles take most
part in this storage is probable, from the fact that the physiological effects upon
the skeletal muscles are manifested for a long time after the effects upon the
heart and arteries have disappeared,

1 In man the effect of taking suprarenal extract by the mouth is to produce a general
diminution in calibre of the arteries as measured by the arterionieter (Oliver, "Croonian
Lectures," Lancet, London, 1896).

2 " Proc. Pliysiol. Soc," Joi/ni, Physiol., Cambridge and London, March 1897.



956 INFLUENCE OF DUCTLESS GLANDS ON METABOLISM.

Source of the active material. — The physiologically active material
is yielded entirely by the medulla of the capsules : no appreciable amount




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can be obtained from the cortex. This result, which was arrived at by
Cybulski and Ijy ourselves from an investigation of the mammalian



so UR CE OF THE A CTIVE MA TERIAL OF SUPRARENAL. 957

organ, has been confirmed in an interesting manner by the observations
of Swale Vincent upon the glands of fishes.^ Elasmobranchs possess
two sets of organs, which appear from their structure to represent the
suprarenal capsules of other vertebrates ; the one of these, the inter-renal
hocly of Balfour, lies between the posterior part of the kidneys in the
middle line ; the other, the imired bodies of Balfour, forms a series lying
on either side, segmentally arranged, on the branches of the dorsal aorta.
Teleosts possess only one kind of gland representing the suprarenal ;
this in its structure is similar to the inter-renal of Elasmobranchs. As
Vincent has shown, the minute structure of the paired bodies of
Elasmobranchs resembles that of the medulla of the suprarenal of other
vertebrates, while the inter-renal body is similar to the cortex of the
ordinary vertebrate suprarenal.^ The physiological test shows this in a
striking manner, for injection of an extract of the j)aired bodies of
Elasmobranchs produces in a marked degree the phenomena which are
characteristic of the medulla of the mammalian suprarenal, while extracts
of the inter-renals of Elasmobranchs and of the corresponding organs
of Teleosts have no such effect.^

Dose. — One of the most interesting and important facts regarding
the material which is yielded by the suprarenals, is the minuteness of
the dose which is necessary to produce the results. As little as 0-0055
grms. (5i mgrms.) of dried suprarenal is sufficient to obtain a maximal
effect upon the heart and arteries in a dog weighing 10 kilos. For each
kdogramme of body weight, therefore, the necessary quantity to produce
a maximal effect is 0'00055 grms., or little more than half a mgrm.^
The active principle is, however, contained only in the medulla of the
gland, not in the cortex, and the medulla in all probability does not form
more than one-fourth of the capsule by weight. Of the dried medulla
certainly not less than nine-tenths is composed of proteid and other
material which is not dialysable, and which otherwise does not conform
to the chemical properties which are associated with the active substance
of the gland. So that, if we take these facts into consideration, we find
that,in order to produce a maximal effect, a dose of not more than fourteen-
millionths of a grm. of the active material per kilo, of body-weight
is all that is necessary. Now it is certainly true to say that one-
fourteenth of this dose will produce some effect, although not perhaps a
very large one. We thus arrive at the astounding conclusion, that the
active principle of the suprarenal capsules, administered in the pro-
portion of not more than one-millionth part of a grm. per kilo, of body
weight, which would be equivalent to xu^o-o gi^nis. (less than gi-y of a
grain) for an adult man, is still sufficient to produce distinct physio-
logical results upon the heart and arteries.^

^ Anat. Anz., Jena, 1897, S. 47 ; " Proc. Physiol. Soc," Journ. Physiol., Cambridge and
London, March 1897, and Proc. Roy. Soc. London, 1897, vol. Ixi. p. 64: and vol. Ixii. p. 176.

- These homologies were long since inferred by Leydig from a study of their structure
(" Fische u. Reptilien," Berlin, 1853), and later by Balfour from a study of their develop-
ment (" Comparative Embryology," 1881, vol. ii. p. 549).

^ It is, however, difficult to avoid contamination with the paired bodies in extracting
the inter-renal. Vincent has also found, in an experiment which is not yet published, that
an eel will survive, for some weeks at all events, removal of the glands which appear to be
the only representative of the mammalian suprarenals, but contain no medullary tissue.

* The proportion of suprarenal capsule to body weight is given by Langlois as from
5Ttnr to TTTTii- in the dog.

^ The chemical nature of this active principle is still obscure, since, in spite of much work
on this subject, it has never been isolated. The history of this has been already dealt
with by Halliburton, along with the chemistry of the suprarenals, on pp. 90-92.



958 INFLUENCE OF DUCTLESS GLANDS ON METABOLISM.



Conclusions. — It may be considered probable that the suprarenal
capsules are continually secreting into the blood an active material, which,
although present in that fluid only in minute quantities, may yet be suf-
ficient to produce very distinct effects upon the metabolic processes of
muscular tissue, and especially the muscular tissues of the vascular

system. It has, in fact, been stated by
Cybulski, and this statement has been
confirmed by Langlois and by Biedl,^ that
the blood of the suprarenal vein contains
a sufficient amount of the active prin-
ciple of suprarenal extract to produce a
marked rise of blood pressure when intra-
venously injected. I have, in spite of
careful experiments, not been able myself
to confirm this statement. Nor is it
easy to understand how it can be true,
■4. since such blood is constantly flowing





Fig. 92. — A. Ergograph tracing of a person suffering from Addison's disease.
B. Tracing from the same person after six weeks' treatment with' supi'arenal
extract. — Langlois. A, natural size ; B, reduced to one-half.



into the vena cava in larger quantity than these observers injected.
But whether we are able to show it experimentally or not, there is
very little doubt of the fact that the materials formed pass somehow
or other into the blood ; and when we compare the results of supra-
renal injection with the effects obtained from the removal and from
disease of these organs, we can come to no other conclusion than
that we have before us a notable instance of internal secretion ; and
that the effect of such secretion passed into the blood is beneficial
to the muscular contraction and tone of the cardiac and vascular
walls, and even of the skeletal muscles, appears very evident from
the results both of removal of the organs and of injection of their
extracts.^

^ Arch. f. d. ges. Physiol., Bonn, 1897, Bd. Ixvii.

■-' This conclusion, which is the one arrived at by Oliver and myself, is ditlerent from that
of Abelous and Langlois, which they formulate thus: — " Les cajisules surrenales ont pour
fonetion de neutraliser on de detruire des substances toxiques ^labor(^es au cours des
echanges chimiques et specialement au cours du travail des muscles." This statement,
which may be taken to set forth the auto-intoxication theorj'' {supra, p. 949), was niHde in
1891, and therefore before the effects of injection of the extract of the medulla were known.



I NFL UENCE OF THE SPLEEN ON ME TAB OLLSM. 959

In advanced cases of Addison's disease, with complete degeneration of the
medulla of the suprarenals, an extract of these organs is devoid of all physio-
logical activity.^ Such patients show, as already stated, extreme muscular
weakness, and very rapidly become fatigued, and their capability of raising a
weight, as estimated by Mosso's ergograph, is extraordinarily small. In one
such case, which was treated by exhibition of fresh capsules of the calf, the
amelioration of this condition was found by Langlois to be very manifest.
The above is well illustrated by the accompanying tracings — Fig. 92, A, in a
patient with Addison's disease ; Fig. 92, B, in the same patient after six weeks'
treatment with suprarenal capsules of the calf.- The tracings are not strictly
comparable, for the second one was taken with half the weight, but on the
other hand it has been reduced to one-half.

Influence of the Spleen on Metabolism.

The constant occurrence and relatively large size of this organ in
vertebrates, the very large supply of blood which it receives, and its
intimate anatomical relationships with the digestive organs, would seem
to render it probable that it must have important functions to perform
in connection with the nutrition of the body.

Effects of ramoval. — This supposition is not, however, borne out by
the result of experiment, for it has been abundantly proved that
the spleen can be completely removed in animals and in man without
their exhibiting any abnormal symptoms whatever.^

Whether the functions of the organ can be taken up under these
circumstances by other organs, such as the lymphatic glands, is a point
which has not yet been determined. In a dog from which I had removed
the spleen several months previously, and which was examined for me
with regard to this point by Swale Vincent, there appeared to be a
larger number of haemal lymphatic glands than in normal dogs,* but it
would require a long series of observations to establish this point
conclusively. Certainly such a function as the formation of lymph
corpuscles may well be carried on by the abundant lymphoid tissue
which is present in other organs of the body; but the spleen has
undoubtedly, in addition to this, a certain influence upon the haemoglobin
of some, at least, of the blood corpuscles which are passing through its
tissue, for we find haemoglobin in various stages of transformation into
other kinds of pigments within the cells of the organ, and also find a
relatively considerable amount of iron in loose organic combination. It
has therefore been supposed that the cells of the spleen pulp may
produce disintegration of effete red blood corpuscles, and that their
pigment may pass to the liver, either as free haemoglobin or as formed
bile pigment. Neither free hfemoglobin^ nor bile pigment can, how-
ever, be detected in the blood of the splenic vein.

On the other hand, the function of producing new red blood
corpuscles has been ascribed to the spleen, on the grounds that

^ Oliver and Scliafer, lot. cit.

^ For other references and observations on the connection between the suprarenals and
Addison's disease, see H. D. Rolleston, Goulstonian Lectures, Brit. Med. Journ., London,
1895, vol. i. ; and Langlois, " Maladie d' Addison," " Dictionnaire de physiologic de Ch.
Richet," Paris, 1895.

^ There is considerable literature on this subject. It has been collected by Pernet, and
is given by him in the British Medical Journal for November 26, 1896.

* For an account of these organs, see Swale Vincent and Harrison, Journ. Anat. and
Physiol., London, 1897, vol. xxi. p. 182.

^ Schiifer, "Proc, Physiol. Soc," May 1890, in Journ. Physiol., Cambridge and
London, vol. xi,



96o INFLUENCE OF DUCTLESS GLANDS ON METABOLISM.

in some animals, e.g. rabbits, after extensive blood-letting, nucleated
erythroblasts, sucb as those which are found in bone-marrow, occur
both in the spleen pulp and in the blood of the splenic vein ; and that
if a spleenless animal is bled, the recovery of the usual percentage of
red corpuscles is less rapid than in a normal animal.^

Whatever may be the nature of its functions in relation to the blood,
it is certain that the organ is in no way essential to the normal
nutrition of the body. It is, on the other hand, not at all improbable
that the main function of the spleen is to serve a mechanical pur-
pose, answering as a reservoir at certain periods of digestion for the
blood which has to pass through the portal system ; and the fact that,
as was first shown by Eoy, the spleen normally exhibits regular rhythmic
contractions and dilatations, seems to point to its exercising an influ-
ence in assisting the flow of blood through the portal vein, and thus
through the liver.^

1 Laiidenbach, Arcli. de physiol. norm, et path., Paris, 1897, p]). 200, 385, and 893.

- The functional connfiction of the spleen with the vascular system is dealt with in the
article on " Circulation " in the next volume. Extracts and decoctions of spleen appear to
have no specific effect, either when injected subcutaneously or intravenously. Nor is any-
thing known as to any specific functions possessed by the thynnis body or by the carotid
and coccygeal glands. Extracts and decoctions of the Ihynuis appear to have no specific
effect when injected intravenously (Oliver and Schafer. Jouroi. Physiol., Cambridge and
London. 189.5, vol. xviii.) or subcutaneously (Vincent, iUd., 1897, vol. xxii.). It has
been stated that removal of the thymus in frogs is followed by a fatal result (Abelous and
Billard, Arch, de pthysiol. norm, et j'cith., Paris, 1896, p. 898), but the statement requires
corroboration.



INDICES.



VOL. I.— 6 1



1



II^DEX OF SUBJECTS.



PAGE

Abrin, 55

Absorption of acid albumin, . . 437

,, ,, alburaose, . . 437, 439

,, alimentary, channels of", . 432

seat of, . 432, 541
,, of alkali albumin, . . 437

,, ,, aromatic decomposition

products, . . .469

,, bile salts, , . 391, 563

,, by blood vessels, 302, 303, 306,

309, 900
,, of carbohydrates, . 431, 434

,, ,, dissolved foodstuffs, . 433

,, by epithelial cells, 435, 436, 451,

449, 454, 457, 486, 488, 659, 685
„ of fats, 369, 392, 443, 449, 451,

459, 462

,, ,, fatty acids, . 450, 454, 457

intestinal, 284, 302, 432, 433, 436,

442, 557

,, of isotonic fluids, . . 304



by lacteals,
of lymph, .
„ oxygen by

canal, .
,, peptone,
,, proteids, 431,
by rectum,
relation,

by renal tubules,
,, skin,
of soaps,
by stomach,
of tyrosine,
water.



457, 462, 610
. 302, 306
alimentary

. 730
. 437, 439
436, 437, 440, 900
. 436
. 215
. 650
. 685, 688
451, 456, 457
. 432, 541
. 469
. 433



Acetic acid, 5, 19, 31, 34, 75, 355, 464, 471,

615, 673
Acetone, . ... 616, 881, 928

Acetylene hfemoglobin, . . . 242
Acetyl-lactic acid, .... 106

Achromic point, 322

Achroo-dextrin, . . . .16, 395

Acid-albumin, . . 50, 167, 404, 408

,, ,, absorption of, . 436, 437

,, ,, carbohydrate from, . 64

,, ,, vegetable, . . .51

Acids of bile. See Bile acids.

,, of body, ..... 1
Acrite, ...... 6

Acrolein, 18, 133

Acromegaly, ..... 946

Acrose, . . . . . . 5, 6

Acrylic acid, . . . . 18, 31, 32







PAGE


Activity co-efRcient,




261,


268


Adamkiewicz's reaction.






47


Addison's disease,




948,


959


Adenine, . 60, 65, 66,


67," 8?


, 93,


596


Adenyl, ....






67


Adenylic acid.






66


Adipocere, ....




20,


933


Adipose tissue, fat of, .






17


,, ,, pigment of, .






20


Admaxillary glands, .




476,


479


Adsorption,






275


Aerotonometer, .






775


Aethalium septicu7n,






80


Affinity, mechanical.






275


Air, alveolar.






774


,, changes in, during respiration.


754,


756


Alanine, ....






31


Albino rabbit.




37,


173


Albrecht's Glaskorper,






222


Albumin, ....






49


,, acid. See Acid albumin.






,, alkali, .


29, 50


, 96,


436


, , , , reducing substance from,


64


, , of aqueous humour.






122


,, ash-free.




.


25


,, assimilation of.






878


,, of blood plasma, .




161,


163


,, ,, cells.




81


, 82



,, chyle, . . . .183
coagulation temperature of, . 43
egg. See Egg albumin.
formula of, .... 26
of intestinal juice, . . 557

,, kidney, .... 92
,, lens, . . . .124

,, liver, .... 86

,, lymph, . . . .182
,, milk. See Lactalbumin.
molecular weight of, . . 26
of muscle, . . .96, 97, 98
,, nervous tissues, . . 118

,, peas, osazone from, . . 64
precipitation of, mechanical, 43
,, ), by salts, . 42

rotatory power of, . . 46

of sebum, .... 674
See Serum albumin.



,, ,, serum.

,, thyroid, .
,, in urine,

, , vegetable.
Albuminates,

,, vegetable.

Albuminoid degeneration.



. 89
437, 604
51, 54
28, 40, 50
. 51
. 74



964



INDEX OF SUBJECTS.



Albuminoids,

,, digestion of,

Albumino-mucous glands,
Albuminous cells,
glands,



PAGE

1, 2, 69, 114
. 429

. 478
. 477, 478
477, 503



,, residue of lifemoglobin, 243, 244

Albuminuria, ..... 604

,, alimentarj^, . . . 437

Albumoid, 124

Albumone, ...... 41

Albumose, 50, 401, 403, 404, 405, 416, 899

437, 439

. 466

64

. 410

. 167

353, 355



412,



absorption of,

,, in bacterial digestion

,, carbohydrate from,

,, classification of, .

,, from fibrin,

,, of gastric juice, .

,, influence of, on coagulation, 146,

147, 177, 181

,, „ ,, epitlielium on, 440

,, of intestinal juice,

,, molecular weight of, .

,, nutritive value of,

, , primary,

,, secondary, .

,, separation of,

, , of urine.

Albumosuria,

Alcapton,

Alcaptonuria,

Alcohol, .....
,, action of, on proteids,

cerotyl, ....
ethyl, ....
,, hexatomic,
,, influence of, on body tempera

ture, ....
,, nutritive value of, .
,, ortho-nitrobenzyl, .
Aldehyde, aspartic, . . . '

,, benzoic.
Aide palmitic acid,
Aldoses, .....
Aleuron grains, ....



557

27

878

416

. 412

411, 412

. 604

. 604

630

630

7

41

20

470

4

820



607,
607,



Alimentary albuminuria,

, , glycosuria,

Alkali-albumin, .



5

38, 39

34

133

4
51, 52



437

436, 609, 881, 945
29, 50, 96, 436



,, absorption of,

,, carboliydrate from,

,, vegetable, .

Alkaline tide, . . . . .

Alkaloids, action of, on body tempera-
ture, .



437
64
51

579

821
664



animal,



milk secretion,
pancreatic secre-
tion, , 548, 550
renal secretion, 648
salivary secretion, 512
skin secretion, 673, 679
34, 58, 673



Alloxan,



in bacterial products, . 59, 465
. _ . . _ . . .597

,, reaction of proteids, . . 48

,, ring, 597

Alloxantin, ...... 592

Alloxuric bases, . . 66, 67, 88, 98, 597

,, nitrogen, .... 597
Aluminium, ..... 78

Alveolar air, ..... 774

,, surface of lungs, . , . 754



I'AGE

Alveoli, Toammary, . . . 662, 666
,, pancreatic, .... 546

,, salivary, .... 477, 507

Anianitine, ...... 60

Amido-acetic acid, . . 31,373,378
Amido-acids, action of liver on, . . 906
,, from bile acids, . . 378

,, ,, proteid decomposi-

tion, . 29, 30, 31, 32, 403
,, in digestion, 419, 420, 421,



nutritive value of.
Amido-butyric acid,
Amido-caproic acid,
Amido-ethy] sulphuric acid,
Amidogen, .
Amido-glucose,
Aniido-isethionic acid,
Amido-isobutylactic acid,
Amido-oxethylsulphonic acid,
Amido-pyrotartaric acid,
Amido-suecinamic acid,
Amido-succlnic acid,
Amido-valerianic acid
Amido-valeric acid,
Amidulin,
Amigdulin, .
Amines,



880

. 31

28, 31, 421

. 373

. 395

9

. 379

. 28

379

32, 421, 426

. 425

29, 32, 421, 425

72, 421

. 31, 32, 82

14, 395

. 395

. 34



Ammonia from decomposition of albumi-
noids, 71, 72,
73
,, ,, ,, ,, cerebro-

sides, 119
,, ,, ,, ■ ,, proteids, 28,

30, 31, 32, 34
29. 76, 427, 472
.473
. 76
. 346
76, 585, 905



,, in digestion,

,, ,, putrefaction,

,, ,, saliva,

,, ,, urine,

Ammonio-magnesic phosphate, 78, 473, 632

Ammonium carbamate, 582, 583, 673, 906,

907
, , carbonate,

,, chloride,

,, lactate,

, , purpurate,

,, salts in blood, .

,, ,, ,, body, .

,, urate,

Amphibia, respiration of,

,, skin secretions of,

Amphicreatinine,



78, 682

78, 907

905, 906, 909

592

905, 907, 919

. 78

. 78

. 703, 723

. 673

61, 101

. 417

104, 405, 416, 418

. 576

14, 395

. 133

. 74

70, 73



Ampho-albumose,
Arapho-peptone, .
Amphoteric reaction,
Amylodextrin,
Amyloid, animal,

,, degeneration,

, , substance,

Amylolytic ferments. See also Ferments. 393

Amylopsin, . . 326, 328, 336, 338, 393

,, action of, on amyloses, 326, 394

,, influence of reaction on, . 339

,, ,, ,, temperature on, 339

Amyloses, action of enzymes on, . . 326

Anabolism, 869, 892

Aniline, ...... 34

Animal alkaloids, . . 34, 58, 673

,, amyloid, ..... 133

,, dextran, . . . . .16



INDEX OF SUBJECTS.



965



Animal gum, 14, 16, 62, 65,



,, heat,

,, proteids, .

,, proteid poisons,

,, quinoidine,

,, sinistrin,
Annelids, haemoglobin of,
Anthrax toxin,
Antialbnmate,



PAGE

126, 133, 158,
613, 665
785
49
55
59
64
187
58
402, 404, 406, 408
Antialbumid, . 403, 404, 405, 406, 408
Antialbumose, . . . 405, 407, 418
Antilytic secretion, .... 522

Antiparalytic' secretion, . . . 522

Antipeptone, . 103, 405, 416, 418, 420

,, molecular weight of, . 27

Antitoxins, ...... 55



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