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The distribution of intermedinA new biological method of assay and results of tests under normal and experimental conditions.

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THE DISTRIBUTION O F INTERMEDIN : A SEW
BIOLOGICAL METHOD O F ASSAY AND
RE SU L T S O F T E S T S UNDER NORMAL
AND EXPERI,\LENTXL CONDI'L'IONS
L. H. K L E I N R O L Z AND €1. R A H N '
Biological Laboratories, Harvard ZJiiicersity, Canibritlgc, ;).lu.s.scichztsetts
FIVE FIGURES
INTRODUCTION
A hormone from the pituitary gland has for a long time
been known to be coiiceriied in the regulation of tlie dermal
melanophores in lower vei-tebi.ates (cyclostomes : Young, '35 ;
frogs : Smith, '16 ; Atwell, '19 ; Hogben, '24 ; Shen, '39 ;Teague,
Noojin and Geiling, '39 ; elasmobranch fishes : Lundstrom and
Bard, '32; Hogben, '36; Waring, '36; Abramowitz, '39; teleosts : T<leiiiholz, '35 ; Abramowitz, '37 ; Osborn, '38 ; reptiles :
T<leinliolz, '35, '36, '38 a, '38 b ; Parker, '38). The function of
this liormone in those higher vertebrates which show no
pliysiological color cliaiiges has been a matter of speculation,
although recent reports (Jorcs, '35) indicate that it niay have
a metabolic role.
E a r l y investigators were uncertain of the exact origin of
this chromatophorotropic hormone, inany being of the opinion
that it was formed by the posterior lobe, a notion that perhaps
' National Research Council Fellow in the Biological Sciences.
The term, 'inteiinedin, ' although used originally by Zondek to designate the
hormone from the pituitary gland that caused dispersion of led pignieiit in the
ciythropliores of the minnow, Pliosinus laevis, has also been applied to the hormone effecting melanophore dispersion. Whether one or two different hormones
weie involved in this activity has been a disputed point. Bottger ( '37 b ) has
rccently presented evidence which s h o w that the two effects a r e mediated by the
same hoimone. I n this paper iutermedin is employed SJ nonomously with the
melanoplioi e dispersing principle.
157
' 1 ' 1 1 1 ~ 4 \ 4 1 0 \ 1 1 C 4 1 ~ lli('OR1)
\OL
76, 30 2 \ N U
LUPI'I k > l i \ ' l
30
2
158
L. €1. IiLlCISHOLZ A S 1 ) H. ILIHS
i ~ pi a r t was clue to the use of commercial l)i*epai*atiorisof
posterior lobe extract by investigators of iiietaehrosis. It was
suggested (Kleinholz, ' 3 5 ) that the melaiiopliore-dispersill~
principle present in these commercial extracts was pl*obabl:a contamination from the pars intermedia aiicl that care was
~iecessaryin interpreting tlie results obtained by the use of
such extracts in physiological studies of color change. Atwell
aiid Holley ('36) and Atwell ('37) showed convincingly that
in amphibians production of i~iteriiieilinis indel~endentof the
pars neuralis, the intermediate lobe of tlic pituitary glaud
being necessai*y for maiiitaiiiiiig the normal dark coloratioii.
Tissue culture experiments by Anderson aiid Haymaker ( ' 3 5 )
arid by Geilirig and Lewis ('35) have shown that growths of
pars intermedia possess the melanophore hormone.
The pituitary glands of those animals (chickeii : DeLawder,
Tarr and Geiliiig, '34 ; Ralin, '39 ; whales : Valso, '34 ; ('reiling,
'35 ; manatee : Oldham, AIcCleery and Geiliiig, '38 ; armadillo :
Oldham, '38) which lack a n aiiatomically distinct lpars iiitei.media contain appreciable amounts of intermediii in the anterior lobe. Because the anterior lobe of tlie chiclieii liypopli;-sis
shows two cytologically distinct lobes (Rahii, '39) our first
iiiterest was in a possible correlation betweeii the distribution
of intermediii and the localization of specific cell types. A s
the work progressed, we fouiid ourselves speculating also
about the phylogenetic aiid ontogenetic distribution of this
hormone. It was therefore decided to conduct a series of
appropriate studies on this subject. This is the first report,
part of which has already appeared in prcliininarj- form
( Kleinholz and Rahn, '39).
MATERIALS A S D 3IETHOI)S
Although Zoridek and Krolin ( ' 3 2 ) used tlic European iiiiiinow, Phoxinus laevis, a s a test animal in their studies of
intermedin, most investigators in this field h a r e found the
hypophysectomized f r o g a more convenient and reliable animal for this purpose. Earlier work had led us to believe that
tlie so-called Americaii ~hanie1~01i,
hiiolis carolinensis, might
ASSAY F O R IXTERMEDI?;
159
prore to be more sensitive to very low concentrations of intermedin than the test animals used by previous investigators.
W e therefore decided to use the hypopliysectomized lizard as
our indicator for intermedin. The bioiiomics aiid physiology
of color change for Anolis have already been reported (Kleinholz, '33, '36, '38 a, '38 b).
Large male lizards, measuring approximately 3 inclies from
tip of snout to vent, were obtained throughout the year from
the Southern Biological Supply House in S e w Orleans. In
the laboratory the animals m7ere kept in mire-screened cages
and were fed blowflies, milk and orange-juice. Water was supplied by sprinkling the cages daily. Operated animals wcrc'
forcibly fed milk and water.
IIypophysectoniy was easily perf ormecl through the oral
route in a maiiiier previously described (I<leinholz, '38 a ) .
Such operated aiiimaIs live for 8 to 10 weeks and with proper
care against dehydration survive for even longel. periods.3
Pituitary glands from several sources were extracted and
tested for tlieir content of chromatopliorotropic hormone. The
chicken pituitaries wei-e obtained at a poultry farm where thcl
animals were prepared for market. The heads of non-la:%qq,
molting Rhode Island Red liens were obtained at the time of
killing, the entire hypopliysis immediately removed and immersed in pure acetone. The dried anterior lobes mere divided
into three approximately equal portions, a cephalic-, middle-,
ancl caudal-third (fig. 1). Beef pituitary glands were obtained
a t a local slaughter house within a few hours after killing. The
glands were immersed and dried in acetone for 48 hours.
Portions of the pars intermedia, free from the large clumps
of colloid, from six glands were dissected out for extraction.
The anuran glands used were of two different sets of experimental animals. I n one lot twenty adult male Raria pipiens
mere maintainecl in an illuminated white box for 36 hours
(using a 100-watt electric lamp a t a distance of 20 inches from
t We have observed, IloweIer, that iii late s p r i n g and suiiiiiicr ( M a y to A u g u s t )
the surviral of such operated :iniiii:rls i s greatly iedi~ceil, individunlu living only
4-13 ria) s a f t e r li~popli~srrtoiii:.
160
L. H. I i L I E I N H O L Z A N D H. G A H N
tlie floor of tlic container) ; a t the eiid of this period the animals were decapitated aiiti the neuro-intermediate lobes removed and placed i n acetone for 24 hours. In the second lot
twenty-two adult male frogs were kept in total darkness f o r
72 hours ; the animals 117ere tlieii decapitated, the floor of the
skull opened to expose the brain aiid l o allow rapid penetration of the acetone into -\vliichthey were placed. This was done
in tlie weak red light from a pllotographic safe-light consisting
of a 50-watt lamp s o u i w of light and a n Eastman Kodak
Series 2 filter.. According to Rodcwalcl ('35 a ) brief exposure
to such dim i-cd light has no effect 011 the activation of the
iiielwiiophorc lioi~monc. Thirty miiiutes after the clecapitatioii
Fig. 1 Sagittal seetioii of the pituitary gland of tlie cliickeii; anterior end is
t o tlic left. The aiitcrior lobe is showii divided into the three portions used in the
prep:ir:ition of t h e extracts. For the distribution of cell Q p e s in this glaiid, sec
Rahii ( '39).
of the last animal, tlie iicui.o-iiitcrmediatc lobes of tlic pituit a r y glands were removed from the skulls aiid immersed in
acetone for 24 hours. The acetone was then decanted from
the tissues and the glands dried in air at approximately 35°C.
The dried pituitary tissues thus obtained were weighed and
e s t i m t e d by boiling f o r 10 to 15 minutes with N/10 NaOH.
Tlic resulting solutioiis were then neutralized with N,/10 HCl,
using phenolphthalein as a n indicatoiu, and made up with coldblooded Ringer's fluid to yield stock solutions, each containing
0.36 mg. of dried powder in 1.0 cc. The stock solutions were
placed in ampules and capped, then immersed in boiling water
for 20 to 30 minutes, after which they were kept iii a cold-room
at 1°C. No prcscrvatires merc added.
ASSAY FOR I S T E R l I E D I S
161
The stock solutions were diluted to desired concentr a t'ions
with Ringer's fluid and 0.2 cc. samples were injected intraperitoneally (subcutaneous injections of ordinarily inert substances like distilled water can evoke a local darkening due to
the pressure exerted on the melaiiophores by the wheal of
fluid) into each of ten hypophysectomized lizards. The degree
of dispersion of pigment within the dermal niclanophores was
measured by dividing the total color range of this lizard (from
bright green to dark brou-n or black) into five stages according
to the method devised by Hogbeii and Slome ( '31 ), and assigning a numerical value to each stage. The accompanyiiig table 1
shows the stages used.
TAB1,E: 1
COLOR
Green
Green with splotches of yellow on head, flanks, and inid dorsuin
Yellowish-brown
Light brown
Brown
Dark brown t o black
__
~
Darkening in Snolis is due to the release of intermedin into
the circulatory system ; the paling response is presumably
effected by the elimination or inactivation of the melanophoredispersing principle (Iileinholz, '38 a, b). The time relations
f o r adaptations of normal lizards to illuminated black and to
illuminated white backgrounds have been determined by a
number of investigators (Carlton, '03 ; Parker and Starratt,
'04; Iileinholz, '38 a ) . The rate of darkening is rapid, maximum melanophore dispersion taking place in 5 minutes. The
change from the dark to the pale phase is much slower, requiring usually about 20 minutes at ordinary room temperatures. Time-curves for these color adaptations are similar to
the one shown in figure 2.
When the times of darkening a r e plotted for hypophysectomized lizards which have been injected with extract of intermedin, the curves shown i n figures 2 and 3 are obtained. The
rate of darkening decreases a s the injected extract is diluted.
162
I,. H. I i L E l X H O L Z A N D H. RAHN
With very low coiicentrations there is a marked flattening and
displacemeiit of the time-curves to the right, i.e., a lower degree of response spread over a longer time period. Conversely, when more concentrated extracts are injected, the
animals darlieii yapidly and may remain for several days
with their melanophores in a state of extreme dispersion before the effect of tlic hoi.mone wears off.
I
I
2
HOURS
I
4
AFTER
I
I
I
6
INJECTION
Fig. 2 The color response of a n hypophysectomized Anolis after injection with
0.0002 ing. (= 0.2 y) of pituitary powder (chicken 9)extract.
MINUTES
AFTER
INJECTION
Fig. 3 The circles represent the average time required 117 eight hypophysectoniized lizards, each receiring extract containing the equivalent of 0.036 nig.
(= 36 y ) of pituitary powder (chicken A ) , t o attaiii the stages of melauophore
dispersion showii in table 1. The triangles show the a\er:rge time taken by niiie
li!.por)lipsectoiiiized lizards, each iujected with 0.0036 (= 3.6 y ) of the same preparation to reach the iiiilex stages.
rlSS.4Y F O R INTEXIlfEDIN
163
CHICKEN
The results of injecting the three extracts prepared from
the anterior lobe of the chicken hppophysis a r e arranged in
table 2. The curves shown in figure 4 were obtained by plotting
the average color indices of tlie animals comprising a test
group against tlie concentrations in gamma dry weight of
pituitary powder injected as extract.
Fig. 4 Curves showing the quantitative (1istril)ution of the melanophore-dispersing horn1011e in the three regions of the anterior lobe of the chicken lippophpsis
(fig. 1). The portions of the curves bctween stage 1 and stage 2 were used f o r
comparison.
Solutions made from the cephalic-third a r e most active in
their melanophore-dispersing properties ; those of the caudalthird a r e least active; the extracts from tlie middle-third lie
between these two in potency. If we take for comparison a
stage 2 color reaction, which is near the mid-point of the total
mnge of response a i d therefore clearly recognizable, the
cephalic-third is seen to be approximately twenty times more
active than the caudal-third (fig. 4). This confirms a n earlier
observation by DeLawcler, Tarr and Geiling ( '34) who were
164
L. H. 1iLEIXHOLZ A K D H. R A H K
of the opinion that the anterior portion of the pars anterior
of the chicken contained more melanophore hormone than did
the posterior region. These authors found, in addition, that
110 melanophore hormone could be detected in their extracts
of the posterior lobe of the chickcii hypophysis.
ThBI,E 2
Erect of extracts
_ _
_._
~~~
LX'IRACT I N J E C T E D
_ _ ~ Chicken A
Chicken A
Chicken A
Chicken A
Chicken B
Chicken E
Chicken B
Chicken B
Chicken C
Chicken C
Chicken C
Chicken C
Beef
Beef
Beef
Chicken A
Chicken A
Frog (light)
Frog (light)
F r o g (light)
Frog (dark)
~~
-~
.
~~~
LKIB1.41.
t h e clironaatic response
~
~
~.
~
~~~
~~~~
I
3.6
0.36
0.20
0.14
1.4
0.36
0.20
0.14
3.6
i
1.4
1.4
1
,
0.i2
1.4
0.73
0.36
0.36
0.14
0.18'
0.10"
0.07"
0.w
~~~
N U M BER O F
.\XIM.4LS I N J E C T E D
DOSE I N G A 3 1 Y h
(:F D R I P I ) PO\VDER
~
GROUP
5-13
3-13
4-13
4-13
4-13
4-13
4-13
4-13
4-13
4-13
4-13
4-13
16-38
16-38
16-38
16-38
16-38
16-38
16-38
16-38
16-38
OIL
STAGE O F R ES P ON S E A X D
AVERAGE DEVIATION
~
9
10
10
10
9
I
1:
!
I
8
1
I
~
9
8
9
9
10
9
11
10
10
6
8
5
I
I
!
I1
I
~
7
I
If
5
I
~
3.7 & 0.8
3.1 & 0.3
1.9 c 1.4
1.0c 0.4
3.4
0.9
1.8 t 1.1
1.2 c 0.9
0.8 ? 0.6
2.1 c 1.2
1.1 1.2
0.5
0.6
0.6
0.8
2.7 2 1.2
1.6 2 0.9
*
*
*
0.7 t 0.7
2.9 It 1.1
1.0+- 1.0
3.2 2 0.6
1.4 t 0.7
0.7 t 0.8
2.9 t 0.9
~
* These d u e s are half of the actual weight of neuro-intermediate lobe powder
injected. They are given to permit comparison in terms of pars intermedia tissue,
since this portion of the gland is approrimatel:- half the rolumc of the neurointermediate lobe.
Since extracts from the cephalic-third of the pars anterior
a r e the most potent and those from the caudal-third least
active, a regional distribution of cell types might be used to
indicate the site of formation of intermedin. The anterior lobe
of the pituitary gland of the chicken contains four cell types
(Rahn, '39). In the caudal lobe occur typical acidophiles to-
ASSAY FOH IKTEXXIEDIN
165
gether with basophiles and chromopliobes ; the cephalic lobe
possesses basophiles and cliromophobes and a new type of
acidophile, comparatively smaller iii size and with finely distributed granules. One is tempted by the results from the
injection experiments and by the generally accepted cytological observation that degraiiulated secretory cells indicate a
resting phase in the physiological cycle of the cell to eliminate
the chromophobic elements from any major role in the formation of intermedin. Such a n assumption would then more
clearly point to the basophiles a s the type responsible for the
production of this hormone. However, more positive evidence
may be obtained from an assay of embryonic glands where
appearance of the specific cell types occurs a t different times
in the ontogenetic history of the animal.
REEF
Several studies have been made of the content of interinedin
in the pituitary glands of beef. Zondek and Krohn ( ’32) nsiiig
the dispersion of erythrophore pigment in Phoxinus as a n
indicator for the pigmentary hormone, analyzed the component parts of the liypopliyseal body and reported 80,000
Phoxinus units per gram d r y weight of pars intermedia
tissue. Lewis, Lee and Astwood ( ’ 3 7 ) reported 255,000 P.U.
per gram (wet weight) of intermediate lobe tissue. The use of
isolated Yhoxinus skin, while apparently of greater sensitivit:(Bottger, ’ 3 7 ) ,must be more rigidly controlled with regard to
pR-, osmotic, and ionic coiicentrations of the extracts to be
tested. Similar precautioiis must undoubtedly be taken in the
use of isolated frog skin a s a test object.
Hogben (’24) and Oldham ( ’38) have used hypophpsectomized frogs as test animals. Oldham, using extract prepared
from ‘standard’ (U.S.P.) pituitary powder, found that 0.003
mg. produced a perceptible darkening of the test frogs. The
so-called standard powder is, howlever, prepared from posteZondek (’35) defines the Phoiinus unit (P.U.) as ‘‘that quantity of hormone
which produres a deep red coloration of f i o m 4-9 sq.mni. a t the point of attachment of the fin.”
166
L. H. IiLEINHOLZ A N D H. R A H N
rior lobes of tlie glands and may therefore coiitaiii varying
amounts of pars iiitermedia tissue. Iiiasinucli as tlic site of
production of tlie melanophore-dispersing hornioiie is now
generally recognized to be in the pars intermedia of the gland,
use of the U.S.P. preparation a s a standard in the assay of
tlie melanopliore hoi*moneseems to us to be of doubtful value.
If the amount of dry weight of pituitary tissue necessary to
give a perceptible dispersion of the melanophores (slightly
(e),
Fig.5 Plots showing the potency of beef pars intermedia
cliicken A1)ortion of the anterior lobe (O),
aiid iieuro-intermediate lobes fro111 light-adapted
frogs ( 0 ) . I n the latter case, however, since the pars intermedia composes
approximately lialf the volume of the iieuro-intermediate lobe, the figures i n d rating the weight of pituitary tissue iiijeetecl are half of the :ic*tu:il wriglit u w l ,
allowiiig expression in terms of pars intermedia powder.
above the minimal threshold concenti.ation of tlic hormoiic)
is coiisideretl a s a uiiit, then from tlic results of Hogben (’24)
there are 400,000 frog units per gram (dry weight) of palas
intermedia tissue, wliile from Oldhaiii’s data there mould be
350,000 frog units per gram (dry weight) of posterior lobe
1)omder. If tlic same defiiiition is applied to Aiiolis tlieii, using
tlie stage 1response (fig. 5) a s the lower threshold value, there
a r e about 2,000,000 units per gram of pars intciwiedia tissue.
ASSAY F O B INTEItMMEUlK
167 T
The Aiiolis unit for intermedin is therefore defiiied as that
weight of pituitary powder which, when injected in the form
of neutralized NaOH-extract into each of ten hypophysectomized Anolis (each measuring between 5-6 em. from tip of
snout to anus), will elicit a n average stage 1 response. The
above results indicate that liypophysectomized Aiiolis is more
0 as an
sensitive than Phoxiiins or the hypophysectomized frob*
indicator for the chroniatophore hormoiie. Jores ( '33) and
Stelile ( '36) have reported that boiling pituitary powder with
dilute alkali increases the chromatophorotropic effect.j That
the greater sensitivity of Anolis may, in part, be due to the
alkali treatment seeiiis to be ruled out by the experiment of
Bottger ('37 b) who shomed that frogs injected with equal
doses of 'natural' and alkali-treated hormone differed not in
the degree of dai.kening but i n the duration of the response.
FROG
Rodewaltl ('35 a ) published an account of her work on the
effects of light aiid of darkness on the coiiteiit of melanopliore
hormone in the hypophysis of Rana temporaria. Among other
things, she reported that formation of the melanophore hormoiic in the pituitary gland occurs only under the influence of
incident light (reflected light being ineffective in this respect) ;
no melanophore hormone is formed in the hypophysis of animals kept in darkness, the hormone initially present being
soon entirely released; similarly, the glands of eyeless frogs
kept in darkness contain no iiielanopliore-dispersing principle.
In a swoad publication ('35 b) Rodewald indicated that in
addition to cessation of pi'oduction of the melanophore horj The potentiation in nielaaopliorr-dispersing effect that results from treatment
of pituitary powder or extracted iiielanophore hormone with weak alkali has been
shown by Hogben and Gordon ( '30) and by Jores and Lenssen ('33) t o be due,
in part, t o the destruction of inhibiting substances, one of which is presumably
the pressor principle. Jores and Will ( ' 3 4 ) believe t h a t a pro-hormone in the
pituitary gland is activated by the alkali treatment. Stehle ( '36) and Iiottger
( '37 11) disagree with this view and ascribe the effects t o a rhange in the hormone
molecule brought about by the alkali. For a recent discussion of this point see
Stehle's ('38) reiiew of this subject.
168
L. H. I i L E I X R O L Z AK’D H. E h H K
mone in tlie glands of frogs maiiitaiiied in darkness, a substance which rendered the hormone ineffective in the blood
was formed in these animals. This substance she believed wab
bound to the blood corpuscles. J o r e s and Hoeltje (’36) reported that tlie melaiiopho~e-liorriioiie-iiiactivatiii~substance
is not depenciciit upon the preseiice of the hypophysis, siiicc
it was also found in tlie blood of liypopliysectomized frogs.
Rodewald’s results seem to us to be contradicted by several
well-linown aiid generally accepted observations on the physiology of amphibian color change. Hogben and Wintoii ( ’23)
reported that R. temporaria (the same species studied by
Rodewald) when kept in darkness uiidcr otherwise normal
conditions, a r e dark in coloi-. Hogben and Slome ( ’31) find a
similar situation in the case of tlie South African toad, Xeiio1)ns laevis ; blinded individuals a r e intermediately dark in
color after 60 hours in darkness. Since metachrosis in anurans
is regulated by the secretion of the pars intermedia, the cliromatic state of such normal aiid blinded animals in cla i.Imess
must be due to tlie secretion of interniediii,F a conditioii which
immediately conflicts with Rodewald’s report.
A recent paper (Masselin, ’39) which appeared a s this
account was being prepared for publication, not only fails to
confirm Rodewald ’ s experiments but describes quite tlie opposite results. IIassclin’s study of Bufo areiiarum shows that
the pituitary bodies of animals maiiitained in darkness for 14
days o r more show an increase in their content of melanopllore
hormone, while glands from animals kept in an illuminated
environment for a n equal interval show a decrease.
Our results, on the other hand, indicate no significant difference in content of intermedin between the neuro-intermediate lobes of frogs kept in darkness for 72 hours and those of
frogs kept in light 36 hours (table 2). Masselin also found
that if the period in darkness or in light was from 1-7 clays,
‘Slight differences in the responses t o light and to darkness are due to the
direct effect of light on the melanopliores; the quantitative measnrements of these
authors show, however, that the direct effect is a n inadequate esp1an:ttion for the
total difference in the melaiiophore indices.
ASSAY F O E 1K’TElLMEl)IN
169
the intermedin content was the same in both cases a s in normal
toads. The differences between our results and those of Rodewald may perhaps be accounted for by the fact that we extracted the glands of twenty or more animals at one time,
probably eliminating to a considerable degree the individual
variations encountered in the assay of single glands.
We do not believe that the dif€ereiices can be attributed to
the method of extraction. Rodewald (’35 a ) reported that
alkaline extracts of glands from animals under the two experimental conditions did not differ in effect from those prepared
by triturating the hypopliyses in Ringer’s solution. Masselin
found that alkaline, acid, or normal saline extracts of the
pituitaries from animals kept under like coiiditions were equal
in their content of the nielanopliore-dispersiii~hormone.
I n view of the conflicting reports of Rodcwald and of Xasselin, and disagreement of our results with those of Roclewald,
we believe that maintenance in darkness ovei’ short periods
does not lead to any variations in the intermedin content of
the anuran pituitary gland.
SYMMARV
1. Quantitative studies in the distribution of intermedin in
the pituitary glands of several vertebrates have been made,
the liypophysectomized lizard, Aiiolis carolinensis, serving a s
the test animal. An ‘Anolis unit’ is defined.
2. The anterior lobe of the chicken’s hypophysis contains
large amounts of intermedin. The cephalic-third of this lobe
contains twenty times more hormone than the caudal-third.
The same region is also m o i ~potent than equal weights of
pars intermedia tissue from beef.
3. The amouiits of intermedin in the glands of light-adapted
frogs and of frogs kept in darkness do not differ significantly.
LITERATURE CITED
AHRAMOWITZ,
A. A. 1937 The role of the hypophyseal melanophore hormone iii
the chromatic physiology of Fundulus. Biol. Bull., rol. 73, pp. 134-142.
1939 The pituitary control of chrolnatophores in the dogfish. Am.
Naturalist, 801. 73, pp. 208-218.
170
L. H. I i L E I l V I i O L Z hKT) H. CAI-IS
AKDEXSON’,
E., AKD 11’. Ij A Y M A R E R 1935 Elal)or:ition of hornioncs by pituitary
cells growing in ritro. Proc. Soc. Esp. Uiol. and N t ~ l . ,vol. 33, pp.
3 13-3 16.
ATWELL, W. J. 1919 On the nature of the pigmentation ctiaiiges following
hypophysectoniy in the frog larva. Scicnce, rol. 49, pp. 48-50.
1937 Functional transplants of the primordium of the epithelial
hypophysis i n amphibia. Anat. Rec., r o l . 68, pp. 431-447.
ATWELL,W.J., AND E. HOI.LEY 1936 Extirl):ition of the p:irs interiiicdia of the
hypopliysis in the young :tmphibinn with subsequent silvery conditioii
and metamorphosis. J. Exp. Zool., rol. 73, pp. 23-41.
Uber das I’igiiieatllorlnoii. I. Der Test. Ztsrlir. gcs. csper.
€ ~ O T T G E RG
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~~
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SHEN,'1'.
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