247
THE EFFECTS OF HYPOPHYSECTOMY AND
ANTERIOR LOBE ADMINISTRATION
ON THE SKIN AND THYROID
OF TRITON CRISTATUS
BY A. ELIZABETH ADAMS.
(From the Department of Animal Genetics, University of Edinburgh1,
and the Department of Zoology, Mount Holyoke College, South Hadley,
Massachusetts, U.S.A.)
(Received 22nd November, 1932.)
(With Two Plates.)
THE skins of many vertebrates react to variations in the amount of hormones of the
endocrine glands, especially of the thyroid and pituitary. Moulting in Triturus
viridescens is inhibited after both thyroidectomy and hypophysectomy or both
operations simultaneously performed. The cornified layers continue to form and
the accumulation of these makes the animal appear grimy (Adams and Richards,
1929; Adams, Richards and Kuder, 1930; Adams, Kuder and Richards, 1932). The
series of experiments here reported demonstrates that a similar effect occurs after
hypophysectomy in immature or mature Triton cristatus (Adams, 1931a). The
moult in Triturus viridescens operated in the above ways can be induced by various
methods. Supplying thyroid hormone by thyroid grafts or thyroxin injections, or
administering iodine to thyroidectomised or hypophysectomised animals or to ones
lacking both glands, or stimulating the thyroid glands of hypophysectomised animals
either by giving pars anterior grafts or injections or by pinching them in situ, will
cause the shedding of the cornified layers (Adams et al, 1930, 1932; Adams, 19316,
1932). Similarly in hypophysectomised Triton cristatus, the moult can be induced by
administration of pars anterior substance or by injection of certain extracts of mammalian anterior lobe.
MATERIAL AND METHODS.
Immature Triton cristatus (10 males and 10 females, average length 7-7 cm.) and
mature specimens (8 males; average length 12-6 cm.) were hypophysectomised by
the buccal route. Fourteen of the immature and six of the mature tritons were
totally hypophysectomised, i.e. the pars anterior, pars intermedia, and pars nervosa
1
I wish to express my appreciation to Prof. F. A. E. Crew for the hospitality of the laboratory
the
Animal Breeding Research Department, the University of Edinburgh, during the autumn of
in
1930 when part of these experiments was carried out. I am indebted also to Dr B. P. Wiesner and his
co-workers for supplying me with the extracts used in the experiments.
248
A. ELIZABETH ADAMS
were removed but the pars tuberalis was left in place. The pars anterior alone was
extirpated from the remaining six immature and two mature animals. Four of the
immature and two of the mature set died within 4 days after the operation. They,
with unoperated, normals served as controls. The remaining sixteen immature
animals lived from 24 to 45 days. Seven of these were given injections of extracts
of bovine anterior lobe ("119," "192," " 196 C)1 or of pregnant human urine
(" 198 E ") 2 or of salt solution at various times after the operation (Table I). Of the
six mature males living more than 4 days after the operation, one (THcJ 7) died on
the thirty-second day after hypophysectomy and the remaining five were given
various treatments (Table I).
For fixation, Bouin's, Zenker's, and Helly's fluids and 10 per cent, formalin were
used. Pieces of skin, the brains (examined to check the completeness of the operation), and the thyroid glands were sectioned. Sections of skin and brains were
7-5 or 10 fx in thickness, those of the thyroids, 5 or 7-5 p. Delafield's haematoxylin
and eosin were used for staining.
OBSERVATIONS.
The first reaction observed externally within 24 hours after total hypophysectomy
was a paling of the orange-black colour of the animal to an orange-brown. After
the removal of the pars anterior alone, no such colour change occurred. Later,
usually within 10-14 days, all the operated animals became grimy. This griminess
increased until the entire animal presented a blackened appearance, with the pattern
considerably or wholly masked (compare PI. I, fig. 1 of a normal animal, with PI. I,
fig. 2 of TH(J 7 hypophysectomised for 32 days). Sections of the skin revealed contracted chromatophores in totally hypophysectomised animals, thus explaining their
orange-brown coloration, while in normal specimens or ones lacking pars anterior
only, the chromatophores were well expanded. Sections also proved the griminess
to be due to the accumulation of cornified epidermal layers which had continued to
form but had not been shed. Normally one, or rarely two, layers of cornified cells
cover the semi-cornified and uncornified layers of the epidermis (PI. I, figs. 3 and 4),
but in the hypophysectomised individuals, three or four (PI. I, figs. 5 and 6) and
occasionally five were found.
1
For the preparation of " 119," see Wiesner and Crew, 1930, Proc. Roy. Soc. Edinburgh, 50, 79.
In a personal communication to the author, Dr Wiesner describes the preparation of " 192" and
" 196 C " as follows:
"Extract 192 is made in the following way: Anterior lobe is finely minced and boiled for a few
minutes with 0 5 per cent, aqueous solution of acetic acid. Filter through paper. The filtrate is
concentrated on a water-bath and adjusted to approximate neutrality with sodium bicarbonate.
10 c.c. corresponds to 2-0 gm. of fresh anterior lobe (beef pituitary).
"Extract 196 C is prepared in the following way: Extract 192 is deproteinised by addition of a
small volume of 20 per cent, solution of sulphosalicylic acid. The proteins are rejected (centnfuged)
while the supernatant liquid alone is used. Alcohol is added to the supernatant liquid until the precipitation of the alcohol-precipitable fraction is complete. The precipitate is well shaken with distilled
water and the mixture is then centrifuged. The fraction which is insoluble in water and is obtained
by centrifugation is taken up in 0 5 per cent, aqueous solution of acetic acid. This yields the ultimate
extract which is adjusted with bicarbonate to approximate neutrality. 1 o c.c. corresponds to 20 gm.
of fresh anterior lobe."
1
For the preparation of " 198 E," see Wiesner and Marshall, 1931, Quart. Journ. Exp. Physiol. 21,
H7.
Effect of Hypophysectomy on the Skin of Triton
249
Twelve animals received treatment after hypophysectomy in attempts to induce
the moult (Table I). Of these, only one (TH $ 5, PI. I, fig. 7) of the two receiving
"119" and the three inoculated with anuran pars anterior moulted. In the latter
three, inhibition of or induction of the moult occurred as inoculations were alternately discontinued and resumed and the condition of the skin at death depended on
whether treatment was or was not being given (compare PI. I, figs. 8 and 9 of
TH <$ 8, being inoculated at time of death, and PI. I, figs. 10 and 11 of TH $ 6, in
which inoculations ceased 92 days before death). In three cases among the twelve
(JH 3 4 with " 192"; JH (J 2, JH cJ 6 with " 198 E") the time of survival after
treatment (8, 16, 30 hours) was too short to expect a moulting reaction. All anterior
lobe extracts ("119," "192," "196C") caused expansion of contracted melanophores of totally hypophysectomised animals, indicating their contamination with
pars intermedia hormone (cf. Spauf, 1927).
The thyroids of a number of the operated animals were examined to see whether
any changes could be detected. In all specimens dying within 4 days after hypophysectomy, they appeared very similar to those of normal individuals (PI. II, fig. 12),
in that the follicles varied in size, the colloid was bordered by or contained many
vacuoles, the follicular epithelium was low or high cuboidal, with rounded or oval,
uncrowded nuclei, that often extended the height of the cell. The thyroids of
individuals hypophysectomised for some time (24-44 days) appeared inactive
(PI. II, fig. 13). Their epithelium consisted of very flattened cells with oval nuclei
lying parallel to the surface of the follicle and often appearing to touch each other.
The colloid was abundant, and in the majority of the follicles, chromophobe vacuoles
bordering the colloid were scarce. The thyroids of animals that had received treatment of various kinds gave dissimilar pictures. The two that had been injected with
extracts of pregnant urine (" 198 E") and had died within the first day after treatment did not differ noticeably from those of hypophysectomised animals. Of the
two (JH 3 10, TH <$ 5) that received aqueous alkaline extracts of bovine anterior
lobe ("119 "), the thyroid of TH <$ 5 alone was usable1. It showed marked stimulation (PI. II, fig. 14). Vacuoles were abundant along the border of the colloid, the
follicles were small and looked as if considerable colloid had been released. The
colloid also stained faintly. Except for follicular size, the condition resembled that
found in the thyroids of the normal animal (cf. PI. II, fig. 12) and, except for degree
of activity, it suggested that in two normal animals (T cr $ 2, T cr $ 4) that had
received injections of " 119" in almost the same amount (4-5 and 4-65 c.c.) over
periods of 12 and 14 days respectively (9 and 11 injections) (cf. PI. II, fig. 15). Those
receiving aqueous acid extracts of anterior lobe ("192," "196C") showed very
slight evidence of stimulation in that the follicular epithelium was cuboidal rather
than flattened and the nuclei were rounded and did not appear crowded. Chromophobe vacuoles were beginning to appear at the border of the colloid (PI. II, fig. 16).
There were, however, few if any collapsed follicles that would indicate a sudden and
extensive release of the colloid. It is interesting that of the seven animals receiving
1
Crossly the thyroid of JH S 10 appeared to show slight stimulation but its cells had undergone
some post-mortem disintegration and its interpretation was therefore doubtful.
jBB-xiii
17
A. ELIZABETH ADAMS
250
Table I. Treatment of Hypophysectomised Triton cristatus.
Treatment*
Amount of
injection (cc.)
or number of
inoculations
(pars anterior
or muscle)
Duration of
treatment
(days after
hypophysectomy)
3°
119
4"5 c c .
30-38 (no injection
33rd day)
JH <J 10
42
119
1-3 cc.
42-45
JH<J4
36
0-7 % NaCl
175 c.c.
192
0-25 c.c.
36-42
45
JH$8
35
196 C
275 c.c.
35-43
192
Animal
Day after
hypophysectomy
when
treatment
began
TH3S
JH$i
36
196 C
015 c.c.
125 c.c.
JH?6
40
196 C
1-5 c.c.
TH<?4
32
Muscle (R.
vuigaris)
E94
192
7 M.
4 0 c.c.
1-5 c.c.
0-25 c.c.
JHc?2
35
JHQ"6
39
198 E (1 : 10
dilution)
0-25 c.c.
T H cJ i
33
Pars anterior
(R. vulgarit)
3 P-A-
198 E
2 P.A.
7 P.A.
2 P.A.
32
Pars anterior
17 P.A.
(R. vuigaris)
TH.J8
30
Pars anterior
(R. pipiens)
25 P.A.
Pars anterior
(R. vuigaris)
18 P.A.
Pars anterior
(R. pipiens)
22 P.A.
43
36-40
40-43
32-40 (no inoculation 35th,
39th days)
Results
Moulting 35th day after
2 5 c.c. injected; thin
moult followed heavy
one; death 39th day, 19
hours after last injection.
Death 46th day, 18 hours
after last injection.
Death 45th day, 8 hours
after last injection.
Death 44th day, 17 hours
after last injection.
Death 41st day, 16 hours
after last injection.
Nearly dead 43 rd day, 5
hours after last injection.
Nearly dead 49th day, 6
hours after last injection.
38-45
47-49
Nearly dead 36th day, 16
hours after last injection.
Death 40th day, 30 hours
39
after last injection.
Moult beginning 37th day
33-35
Thin moults 47th, 52nd,
55th days; nearly dead
38-39
56th day, 4 hours after
46-52
last inoculation.
55-56
32, 34-36, 38, Moult 47th day (15th day
after first inoculation);
43-45, 47-49,
thin moult 47th, 52nd,
53-56t
56th day.
Re-darkened.
Gonadectomi8ed
91st day.
94,97,101,105,
108, n o , 112, Moult beginning 96th day
115, 117, 1 1 9 , (3rd day after first ino122, 124, 1 2 6 , culation) ; thin
moult
133, 136, 140, 101st, 105th, 108th, 115th
129th,
143, 157, 161, 122nd, 126th,
157th,
199th
164, 168, 171, 143rd,
178, 185, 192
dayj.
Re-darkened by 213th day;
death 284th day; heavy
cornification.
30-32, 35, 36, Moult beginning 34th day
(4th day after first inocu45-56 §
lation); thin moult 38th,
49th, 52nd day.
Re-darkened by 79th day.
94, 97, 101,
Moult beginning 97th day
105-106,109,
(3 days after inoculation);
112, 115, 117,
thin moult 101st, 105th,
119, 122, 124,
109th, 112th, 122nd,
126, 133, 136,
124th,
143rd,
157th
140, 143, 157,
day ||.
161, 164, 168, Death 174th day.
35
171
Effect of Hypophysectomy on the Skin of Triton
251
either "119," " 192," or " 196 C" (Table I) only TH ^ 5 moulted and its thyroid
alone appeared unusually active.
Of the three individuals that received anuran pars anterior (Table I), TH <$ 1
and TH <J 8 were being inoculated at the time of death. In the former, the follicles
of the thyroid glands (PI. II, fig. 17) were widely separated by vascular spaces, and
although many of them contained colloid, they were smaller than those of an hypophysectomised animal and the colloid was bordered by numerous chromophobe
vacuoles. Many of the smaller follicles contained vacuoles throughout the colloid. The
gland seemed like a transition stage between that of an hypophysectomised and of a
normal animal (cf. PI. II,figs.12 and 13). The thyroids of TH $ 8 showed hyperplasia
of the follicular epithelium with a marked increase in the height of its cells (PI. II,
fig. 18). Further, the colloid was almost wholly absent from the gland and stained very
faintly. The thyroids of TH $ 6 did not present the completely typical condition after
hypophysectomy, although 92 days had elapsed since the last inoculation with pars
anterior (cf. PI. II, fig. 19 with PI. II, fig. 13). The gland contained considerable
colloid, but its follicular epithelium was not as flattened as after hypophysectomy
without treatment. But neither was there evidence of colloid release. Interpreted in
the light of the appearance of the thyroid in TH <J 8, it seems likely that this gland had
been similarly stimulated during the second period of inoculations (25 pars anterior
in 98 days) and that even the lapse of 92 days without treatment had not sufficed to
re-establish completely the condition found after hypophysectomy without treatment.
DISCUSSION.
The cutaneous reaction in both immature and mature Triton cristatus after
either total or partial (pars anterior removal) hypophysectomy resembles that in
mature Triturus viridescens after similar operations (Adams et at. 1930, 1932). In
both forms, an inhibition of moulting occurs, the cornified layers of epidermis continue to form, and the animals gradually darken as these layers increase in number.
A similar reaction occurs in some other amphibians (toads) after pars anterior
removal, either with or without the other parts of the pituitary (Giusti and Houssay,
1921,1922,1923,1924; Puente, 1927), although in frogs the cuticular change does not
take place (Houssay, 1917; Houssay and Ungar, 1924). The skin and its derivatives
in other vertebrates also show the effect of too little anterior lobe secretion1.
It is also evident that the thyroid glands of the hypophysectomised Triton
cristatus present a picture of lowered activity, similar to that following this operation
in Triturus viridescens (Adams et al. 1932) and in other forms1. The glands contain
1
See discussion in Adams et al. 1932, Joiirn. Exper. Zool. 63, 1.
Notes to Table I.
• In this column, 119 = aqueous alkaline and 192 and 196 C = aqueous acid extracts of bovine
anterior lobes; 198 E = extract of human pregnant urine; E 94 — placental extract. For preparation
of E 94, see Wiesner, 1930, Edinburgh Med. Journ. February.
t On 53rd and 56th days 2 pars anterior were inoculated.
X Moults not recorded in interval between 157th and 199th days.
§ On 56th day 2 pars anterior were inoculated.
|| Moults not recorded after 157th day.
17-3
252
A. ELIZABETH ADAMS
considerable amounts of colloid with relatively few chromophobe vacuoles, the
follicular epithelium is flattened, its nuclei are oval in shape and lie parallel to the
surface of the follicle. Interpreted in the light of the experiments on Triturus, it
seems that the inhibition of moulting is really an indirect rather than a direct effect
of the removal of the pituitary, that is, it is due to the inactivation of the thyroid.
Although few of the hypophysectomised Triton cristatus were subjected to treatment likely to induce the moult after hypophysectomy, they bear out the above
interpretation. Only an aqueous alkaline extract of bovine anterior lobes (" 119")
and inoculations with anuran pars anterior gave positive results (Table I), and after
these treatments alone was there evidence of thyroid activation (see p. 249). In one
case (TH $8), the glands had been so stimulated by continued inoculations that
they resembled the condition in human exophthalmic goitre, or in spontaneous
hyperplasias in other mammals (Marine, 1907), or that in a dog made acromegalic by
anterior lobe injections (Putnam, Teel and Benedict, 1929; Teel and Cushing, 1930),
or that in an hypophysectomised puppy receiving daily transplants of anterior lobe
material (Reichert, 1928). Similarly in Triturus viridescens (Adams et al. 1930, 1932)
moulting only occurred in hypophysectomised animals when their thyroid glands
were induced to release their hormone.
The question naturally arises as to why only the pars anterior inoculations and
the aqueous alkaline extract of bovine anterior lobes caused the release of sufficient
thyroid hormone to induce the moulting reaction. In the former cases, the fresh
material administered probably contained all of the hormones produced by the pars
anterior, and it was to be expected that they would stimulate the thyroid gland.
Activation by the use of similar material is in line with the work of many investigators on normal amphibians and mammals (Swingle, 1921; Allen, 1929; Ingram,
1929, 1930; Grant, 1930, 1931; Uhlenhuth and Schwartzbach, 1926, 1927, 1928a
and b; Aron, 1929, 1930a, b, c, d, 1931; Loeb and Bassett, 1929, 1930; Loeb,
Bassett and Friedman, 1930; Loeb, 1932a and b; Watrin and Florentin, 1929)
and on hypophysectomised ones (Allen, 1929; Smith and Smith, 1922, 1923;
Adams et al. 1930, 1932; Smith, 1927, 1930; Reichert, 1928). It was hoped, however, that the injections of extracts might throw some light on the question of
whether a specific thyroid-stimulating hormone exists in the anterior lobe (Crew
and Wiesner, 1930) and indicate whether it was contained in both alkaline and acid
extracts of anterior lobes. Moulting and the histological evidence of thyroid activation after the injection of " 119 " (the alkaline extract) show the presence of a thyroidstimulating substance in this extract. Its existence in similar extracts has also been
demonstrated by Loeb and Bassett (1929,1930), Loeb, Bassett and Friedman (1930),
Loeb (1932a and b) and Schockaert (1931, 1932) in guinea-pigs and ducks, by
Adams (1932) using phyone of Van Dyke and Wallen-Lawrence (1930) on normal
and hypophysectomised Triturus viridescens, by Teel and Cushing (1930) in dogs.
The experiments with acid extracts, however, are wholly negative, a fact which may
be due to inadequate dosage or inadequate survival periods after treatment (Table I).
It should be mentioned, however, that the thyroid-stimulating potency of acid
anterior lobe extracts is evidenced in the work of Loeb and Bassett (1929, 1930),
Effect of Hypophysectomy on the Skin of Triton
253
Loeb, Bassett and Friedman (1930), Loeb (1932a and b) in guinea-pigs and in that
of Spaul (1930) on tadpoles and axolotls. Further, from the work of Loeb and his
collaborators (1929-32) in which thyroid activation in the guinea-pig was secured
with equal facility with either acid or alkaline extracts and at the same time a gonad
response (which, however, did vary with the type of extract) occurred, it seems
doubtful whether the method of extraction of anterior lobe substance can separate
a specific thyroid-stimulator or even whether a specific thyrotropic hormone (Crew
and Wiesner, 1930) exists.
As for the other extracts employed, it is not surprising that that from human
placenta did not activate the thyroid, although 4 c.c. were injected over a period of
8 days. A similar impotency was found by Crew and Wiesner (1930) and by Klein
(1929) and Aron and Klein (1930) in tests on axolotls and guinea-pigs. The
effectiveness of the urine of pregnancy (" 198 E ") cannot be judged since survival
was too brief to reveal any change (Table I). It should be noted, however, that
Aron and Klein (1930) claim thyroid-stimulating power for both normal and
pregnant urine when tested on guinea-pigs, while Loeb (1932a and b)findspregnant
urine incapable of causing hypertrophy in the guinea-pig thyroid.
There is one further point worth brief mention, although it cannot be settled by
the data available at present. If the pituitary stimulates the thyroid to release its
hormone, can the hormone be renewed in the absence of the anterior lobe? It was
thought that TH <$ 6 might harbour a clue to the answer to this question, but the
condition of its thyroids suggests recovery after excessive stimulation rather than that
which might have been expected as a result of the release of their colloid content
followed by no further stimulation. The presence, however, of a considerable
amount of colloid in the follicles (PI. II, fig. 19) suggests that secretory activity may
possibly occur even when the anterior lobe is absent. Whether the pituitary controls the secretory activity of the thyroid as well as the release of the thyroid hormone
merits further investigation.
SUMMARY.
1. An inhibition of moulting occurs after hypophysectomy (total or pars anterior
removal only) in mature and immature Triton cristatus. Cornification of the outer
epidermal layers continues and the animal gradually becomes grimy as these layers
accumulate. The thyroid glands of such animals become inactive.
2. Moulting was induced in hypophysectomised animals by means of inoculations
of fresh anuran pars anterior lobes and in one case by injections of an aqueous alkaline extract of bovine anterior lobes. A study of the thyroid glands of these animals
supports the conclusion that the moulting is brought on by the stimulation of the
thyroids to release a considerable fraction of their colloid content. It thus appears
that the inhibition of moulting is really an indirect result of pars anterior removal.
Hypophysectomy results in an inactivation of the thyroid gland and the resultant
lack of circulating thyroid hormone is responsible for the failure to moult.
254
A. ELIZABETH ADAMS
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JOURNAL OF EXPERIMENTAL BIOLOGY, X, 3
PLATE I.
. 11.
ADAMS—'I'HK KKFI-XTS OF HYPOPIIYSECTOMY AND ANTERIOR LOBE
ADMINISTRATION ON THE SKIN AND THYROID OF TRITON CRISTATUS (pp. 247-255).
JOURNAL OF EXPERIMENTAL BIOLOGY, X, 3.
18
PLATE II.
19
ADAMS—THE EFFECTS OF HYPOPIIYSECTOMY AND ANTERIOR LOBE
ADMINISTRATION ON THE SKIN AND THYROID OF TRITON CRISTATUS(pv- 247—255)-
Effect of Hypophysectomy on the Skin of Triton
255
VAN DYKE, H. B. and WALLEN-LAWRENCE, Z. (1930). Journ. Pharmacol, and Exp. Tlierap. 40, 4:3.
WATHIN, J. and FLORENTIN, P. (1929). Compt. Rend. Soc. Biol. 101, 1200.
WIESNER, B. P. (1930). Edinburgh Med. Journ. 37, 73.
WIESNER, B. P. and CREW, F. A. E. (1930). Proc. Roy. Soc. Edinburgh, 50, 79.
WIESNER, B. P. and MARSHALL, P. G. (1931). Quart. Journ. Exp. Physiol. 21, 147.
EXPLANATION OF PLATES.
All figures except figs. 3 and 5 are of mature male Triton cristatus; figs. 3 and 5 are of immature
females of the same species. Whole mounts of skin, x 4; sections of skin, A 215 ; sections of thyroids,
.
100.
PLATE I.
Fig. 1. Whole mount of skin of normal animal, Triton cristatus 3 1. Note distinctness of pattern.
Fig. 2. Whole mount of skin of TH 3 7, 32 days after hypophysectomy. Note masking of pattern by
accumulation of cornified epidermal layers.
Figs. 3 and 4. Sections of skins of normal immature female (JH 2 9) and normal mature male
(Triton cristatus $ 1) respectively. Note single layer of cornified epidermis.
Figs.sand6. Sectionsofskinof immature female (JH '? 2) and mature male (TH ^ 7), 24 and 32 days,
respectively, after hypophyaectomy. Note increased number of cornified epidermal layers.
Fig. 7. Section of skin of hypophysectomised animal (TH <J 5) showing lack of cornified layers after
moult had been induced by an aqueous alkaline solution of bovine anterior lobes (119) 35 days after
hypophysectomy. See Table I.
Figs. 8 and 9. Whole mount and section, respectively, of skin of hypophysectomised triton (TH c? 8)
showing clear pattern (fig. 8) and lack of cornified layers of epidermis (fig. 9) at time of death.
Successive moults had been induced by inoculations of anuran pars anterior. See Table I.
Figs, ioand 11. Whole mount and section, respectively, of skinof hypophysectomised Triton cristatus
(TH ^6) showing masked pattern (fig. 10) and several cornified layers of epidermis (fig. 11) at
time of death, 284 days after hypophysectomy. These cornified layers represent the third accumulation after hypophysectomy, moults having been induced by anuran pars anterior twice since the
operation for hypophysectomy. See Table I.
PLATE II.
Fig. 12. Thyroid of normal animal, Triton cristatus 3 1.
Fig. 13. Thyroid of TH 3 7, 32 days after hypophysectomy. Note flattened follicular epithelium.
Compare with fig. 12.
Fig. 14. Thyroid of T H 3 5, which had been induced to moult its accumulated cornified layers
35 days after hypophysectomy by injections of an aqueous alkaline extract of bovine anterior lobes
(119). Thyroid has released much of its colloid and its follicular epithelium is cuboidal. Compare
with figs. 12 and 13. See Table I.
Fig. 15. Thyroid of a normal animal (Triton cristatus 3 2) which had received 45 c.c. of an aqueous
alkaline extract of bovine anterior lobes (119) in nine injections over 14 days. Note its very active
condition. Compare with normal thyroid (fig. 12) and with thyroid of hypophysectomised animal
activated by the same extract (fig. 14).
Fig. 16. Thyroid of hypophysectomised animal (TH ^4) after injection of r j c.c. of an aqueous acid
solution of bovine anterior lobe (192). Thyroid shows some evidence of activation in heightening of
follicular epithelium, but colloid release has been very slight. This animal did not moult
following treatment. See Table I.
Fig. 17. Thyroid of hypophysectomised animal (TH 3 1) after inoculations of anuran pars anterior.
Collapse of follicles and marked vascularity indicate colloid release. See Table I.
Fig. 18. Thyroid of hypophysectomised animal (TH 3 8) showing marked hyperplasia following
long-continued inoculations of anuran pars anterior. See Table I.
Fig. 19. Thyroid of hypophysectomised animal (TH 3 6) showing only partial effects of hypophysectomy. Animal had been induced to moult at two different times by inoculations of anuran
pars anterior after hypophysectomy and then had re-darkened for the third time. See figs. 10
and 11. Probably the thyroid had been so stimulated by these inoculations (compare fig. 18) that
even after their cessation it had not wholly reverted to the inactive condition characteristic of
hypophysectomy.