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Origin of thyroidectomy cells.

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ORIGIN O F THYROIDECTORIY CELLS
HERBERT E L P T M A S
Department of A n a t o m y , College of Physicians and Surgrons,
Columbia Cnivcrsity, New Y o r k
SIX FIGURES
Among the effects of thyroidectomy obscrvcd by the early
investigators were alterations in the anterior pituitary gland.
Accurate cytological description awaited the study of the goat
by Trautmann ( ’16) and the publication of pliotoniicrographs
of the thyroidectomy cells of the rat by Rojima (’17).
Experiments aimed at determining which type of normal
pituitary cell gives rise to the thyroidectomy cell were Iianipercd at first by the lack of adequate cytological methods.
Thyroidectoniy cells were held to be essentially like castration
cells by Severinghaus, Smclser and Clark (’34) and by
Severinghaus ( ’38) while the difference between thcrn were
emphasized by Zeckwei-, Davison, Kcllcr and 1,ivingoocl ( ’3t5),
Zeckwer ( ’37) and Reese, Koneff and Wainniaii ( ’43).
The modern aspects of the prolnleni of thyroidectomy cell
origin date from the introduction of histochemical methods
into pituitary cytology. Following a lead supplied by Goniori
( ’50)’ aldehyde fuchsin was found by Halnii ( ’50) to be a
specific stain for the beta cells of Romeis (’40). The term
“tliyrotroph” was suggested a s a descriptive name for this
cell by Purves and Griesbach ( ’?la, b) on the basis of evidence
that it is the source of thyroid stimnlatiiig hormone.
Ability to distinguish thyrotrophs from gonaclotrophs and
thyroidectomy cells from castration cells did not provide an
unequivocal solution to the problem of tliyroidectoniy cell
Aided by a grant f r o m the National Science Foulidation.
* W i t h the technical assistance of Mary Clarke Miksic and Cynthia Anii Galazzi.
119
120
tlE1:BEI;'I' ELFTMAS
origiii. Altliougli Halmi ( '52), Farquhar and Xiiieliart ( '54)
and Yurves and Griesbach ( '56) have summarized evidence
ivliich points strongly to the development of thyroidectoiny
cells from thyrotrophs, this origiii ~ v a sdenied by Scharf and
B'orstei. ( '54). The iiiconipleteiicss of tlic present evideiicc
stenis from the fact that thyrotrophs lose tlieir diagnostic
i*eccptivity for aldeliycle fuelisin some time before the thy]-oiclectomy cells develop their characteristic periodic acitlScliiff ( P A S ) positive granules. The current ~sperinieiits
are concerned with one method of hridging this gap.
JIATERIALS A S D METHODS
The r a t s used in these esperinients were of the Long-Evaiis
strain and belonged to a larger group utilized for liistocliemical
research on the pituitary. Thyroidcctomy was performed 011
3 2 animals and their pituitaries collcctecl for histological
study at periods ranging u p to 49 clays after operation. The
effect of a goitrOg.cn, propyltliioui-acil, on the pituitary was
investigated in 97 rats by niisiiig it with their food to tlie
extent of 0.1% aiitl collecting the tissues at intervals up to
286 days.
Sumerous fixatives were used for the pituitary gland i i i the
early stages of this investigation but only 4 methods of
preparation proved to be sufficiently satisfactory for inclusion in this report. (1) F o r general pituitary cytology,
chrome-alum fixation (Elftrnan, '57a) was followed by either
PAS or aldeliyde fuchsin with oraiige G for the acidophils.
( 2 ) F o r finer cellular detail, osmichrorne fixation (Elftman,
'57c) was superior for those features susceptible of visualization by PAS or by aiiilin acid fuchsin. ( 3 ) F o r specific drnionstration of the Golgi apparatus tlie direct silver method (Elftman, '52), followed by chrome-alnni fixation, was most useful ;
after gold toning, the P A S procedure was applied with
oraiige G following. (4) F o r phospholipids, including tliosch
of the Golgi apparatus, controlled chromation usiiig dichromate-sublimate (Elftman, '54, '5713) was followed by P A S processing before Sudan black staining.
OBSEKVATIO
Identification of thjrotrophs.
The appeai*aiice of a gi*oup
of thyrotrophs stained with aldehyde fuclisin is illustrated iri
figure 1. The details of the surrounding cells have intentionally been rendered inconspicuous by photographic filtering in
order not to detract from the observation of the thyrotroplis.
Tlie angular appearance of many of the thyrotrophs offers an
easy way of making a preliminary identification but has liniitations as a diagnostic characteristic since rnaiiy thyrotrophs
do not exhibit this shape. The processes extending out from
the main body of the cell may be c w x longer than the oiic
n.liich is evident in the lower central poi-tioii of figure 1. Sucli
extensions of thc protoplasm, n~licnthey can be folloiwil to
their termination, typically end on a sinusoicl, apparently making it unnecessary for the main p a r t of the cell to occupy estensive sinusoidal frontage. It is evitlcnt that thc sliapc clisplayed hy thyrotrophs in sections of tissue will depend on the
orientation of the section and angularity is not a sufficient
criterion for identification.
The staining of thyrotrophs by aldehyde fuchsin fm-nislics
a specific means of recognition in the rat, provided the stain
is applied after proper fixation and the cell is in a iiornlwl
functional state. The affinity of cellular constituents for aldehyde fuchsin is markedly affected Iny oxidation, either during
fixation or later, but chrome-alum fixation preserves its spccificity for the thyrotrophs among the anterior pituitary
cells. Unfortunately for the investigation of thyroidectonip
cell origin, the thyrotrophs lose their capacity to bind alclchyde fuchsin before they acquire the cytoplasmic staining
characteristics of thyroidectomy cells.
A more durable characteristic of the thyrotroph may be
seen by close scrutiny of the large cell located near the left
border of figure 1. The nucleus can be recognized as an unstained sphere and lying in close contact ahove it is the dark
mass of the Golgi apparatus.
ItIen,fification of tlzyroidecto+i??ycells. Thyroiclcctonip cells
from a pituitary gland prepared 49 days after the operation
arc shown in figure 2. Tlie cell most casily stutlietl Iiws its uiistaincitl nucleus in the center of the figure. Tlie cptoplasrii
contailis ~iunie~’ous
tliyroidectorny granules which stain lwilliaiitly with the PAS technique. Tlic Golgi apparatus occupic\
a11 :ii~’aoiilp slightly smaller than that taken up hp th(1 iinclens.
It is closely applied to the nucleus. The thyroidectoiup cclls
a r e relatively large, their size being indicated by the distrihntion of the granulcs. Their general shape is u~itle~iial)ly
i.eiiiini\ccnt of the thyrotroplis although they give the inipression of being more trapezoidal than angular. The ccintral
thyroitlectorny cell of fig~11.e2 slio\vs another thprotropli charactcristic, that of rcacliiiig out to a sinusoitl hy I ~ I C H I I S of ii
protoplasmic process.
0 lg i Ct]lpCI T ( l f 21 S O f t 11 jJ 1’0t T 0 1)?1 (1 T?(1 f h .I)T 0 I 1 C C f 0 7 11 jJ C 0 11h .
Tlic similarity ill the Golgi apparatus apparent in tliprotro1)li
and tliproidectomy cells fixed f o r general study suggwts the
application of moi-c specific Golgi niethods. This lias 1)een tloiie
in figures 3 and 4 by using the direct silver nietliod ant1 iii
figuws 5 and 6 l)y m e a n s of phospholipid p ~ s e r r ttioii
i
coiiti*ollecl cliromation.
Wit11 both of these techniques it is possible to apply tlic
P A S reaction, in the case of the silver method after tlic silv(1i’
has heen replaccd by gold, and so to identify the tliyroitlec. .
torriy granulcs in figures 4 and 6. Tlie positive PAS staiiiing
of the thprotroph cytoplasm in figure 3 is not specific, siiice
gonadotrophs also stain, hut it is a n aid in locating the ctllls.
TVhen the two thyrotrophs which occupy tlie center of figure 3 a r e compared with the two thyroidectomy cells similarly
placctl in figure 4, tlie similarity in the Golgi apparatlis 1 ~ cotncs apparent. I t s chai.actei*istics include a geiieral density
which contrasts with the vesicular appcaranccl of the gonadotroph Golgi ; a size almost, but not quite, equal to that of t h c
nucleus; and a close association with the nucleus, \vhicli
liave referred to as capping.
Inspection of figures 5 and G show the similarities of the
Golgi apparatus in tlie two types of cells when it is visnalized
by Suclaii black staining of its phospholipid component. Tho
~
(
1
0L:IGIS
O F T HYROIDE CTO MY CELLS
12::
aiigular tliyrotropli in the upper left portioii of figure 5 a i d
the less angular one in the center both show a dense Golgi
mass in close contact with the unstained nucleus. Tlie tliyroidectoriiy cells of figure 6 are co~ispicuousbecause of their
tliyroitlcctorriy granules, red in tlie original, but they a r e only
partially obscuring the Golgi apparatus.
UTSCUSSIOS
T h r striltiiig similarity between tlie Golgi apparatus in the
tliyrotropli and in tlie tliyroidectomy cell provides a connecting
link between them whicli considerahly streiigtlicns the coilception of tliyrotrophs becoming tliyroidcctorny cells by internal modification. The functional significance of the close
relation of the Golgi apparatus to the nucleus in these cells,
contrasting as i t does with the situation in goiiaclotrophs, invites further study.
One intercisting feature of the type of Golgi apparatus illustrated here f o r the thyrotrophs and tliyroidectomy cells is
that it bears a closer resemblance to the type which Scveringhaus ('38) found to be characteristic of acidophils than t o
his basophil type. This may be responsible for some of the
confusion which has attended this problem ; with the improved
techniques for the Golgi apparatus now available it is to hc
hoped that further study of this organelle will again coiitribute to the clarificatioii of pituitary cytology.
Tlie detailed differences between the Golgi apparatus of
the various types of pituitary cells cannot be discussed in
this paper but attention has already been called to one typical
difference between gonadotrophs and the cells with which we
are concerned. This difference lies in the more vesicular appearance of the gonadotroph Golgi body after either direct
silver or controlled chromation compared with the denser
construction in thyrotrophs and thyroidectomy cells. It is
consequently interesting that Farquhar ( '54), working at the
electron microscopic level, found that vacuoles a r c moro coilspicuous in the Golgi apparatus of the castration hasophil
and the lamellar constituent in the thyroidectorny cell.
\\’hethcr the differeiiccs a r e natural or produced by fixation,
they indicate some fundaniciital difference between these two
varictics of Golgi apparatus.
h u JI 11A 11 Y
Tlic origin of thyroidectorriy cells from thyrotrophs has
pl.ovc siiiccl tlic diagnostic characteristics of
the tliymtrophs arc usually lost before those of the tllyroidcctorriy cclls appear. Study of the Golgi apparatus by the direct
silver a r i d the coiiti*ollctl chi*omation methods has sho\vn it to
f o r m a. dense aggregate capping the nucleus in both thyrotroph
and tliyroidectomy cclls. Since this Golgi structure differs
fi-om that charactci*istic of goiiadotrophs it may ho regartled
as a coiiriecting liiik bct\vceii tliyrotrophs and thyroidectomy
cells.
wrmhTcm C r T m
~ ) ( y q l(lifficult to
ELFTMl x , H. 19.52 A d i r w t silver method f o r the Golgi apparatus. Staiii Tech..
2 7 : 47-52.
1954 Controlled chronixtion. J. Histocheni. & Cytochern., 2: 1-8.
_ _ _ ~ 1957a
~
-4 chrome-alum fixative f o r the pituitary. Stain Tech., 3‘2:
23-28.
1957b Pliospholipitl fixation by dicliroiiiate-sublimate. Ibiil., 32:
29-31.
19.57~ Osmichrorne fiuation. Quart. ,J. Micr. Sci., 98: 15-18.
FARQUHAR,
M. G . , A R D J. 3’. RINEIIART 19.54 Cytologic alterations in the ailterior pituitary gland following thyroideetomy : an electron microscopic study. Endocrinology, 5 5 : 857-876.
GOMORI, Q. 1950 Aldehyde fuchsin: a new stain f o r elastic tissue. h i . J.
Clin. Path., 20: 665-666.
H I L N I ,S . 8. 1950 Two types of basophilx i n the anterior pituitary of the r a t
:rnd their respective eytophpsiological significance. Endocrinology, 47 :
289-299.
____
1952 Two types of basophils in the r a t pituitary: “thyrotrophs”
and “gonadotroplis.” Tbid., 5 ’ 1 : 140-142.
KO.JIXA,M. 1917 Tlic relations of the pituitary body with the thyroid and
parathyroid and certain other endocrine organs in the rat. Quart. J.
Exp. Physiol., I 1 : 319-338.
P U R V S S , H. n., AND w. E. (:RIESRACH
1951a The site of thyrotrophin and
gonadotrophin production i n the rat pituitary studied by MeManus
Hotchkiss staining f o r glycoprotein. Endocrinology, 49 : 244-264.
195lb The significance of the Gomori staining of the bxsophils of
the r a t pituitary. Ibid., 49: 6.5-662.
_ _ _ 1956 Changes in the basopliil cells of the r a t pituitary after thyroidectomy. J. Endocrin., 13: 363-375.
OItIGIN O F T HYBOIUE CTO MY CELLS
1%
REESE,J. D., A. A. KONEFFAND P. WAINMAN1943 Cytological differences be
tween castration and thyroidectomy basophils in the r a t hypophysis.
I n “Essays i n Biology,” Llniv. of Calif. Press, 471-485.
ROMEIS,B. 1940 Die Beta Zelle. In von Mollendorff’s Hand. d. mikr. Anat. d.
Menschen, Berlin, Springer, 6 ( p a r t 3 ) : 99-111.
SCIIARF, J. H., AND W. FORSTER 1954 Das Zellbild cler Rattenhypopliyse. &it.
f . Zellforsch., 40: 117-138.
SEVERINGHAUS,
A. E. 1938 The cytology of the pituitary gland. Assoc. Res.
Nerv. Mental Dis., 1’7: 69-117.
REVERINGIIAUS, A. E., G. K. YXELSER A N D TI. M. CLARK 1934 Anterior pituitary
changes in the adult m d e rat following thyroidectoniy. Proc. So(.. Exp.
Biol. & Med., 31: 1187-1129.
‘I’RAUTMANN, A.
1916 EIypophysc und Thyroidektomie. Frank. %it. f . Path.,
18: 173-304.
1937 Morphological changes in the pituitaries of rnts icsultiiig
Z:F.CK\VEk, I. T.
from combined thyroidectoniy :uid gonailectomy. Ani. .J. L’ath., 13:
985-991.
Z W K W E R , I. T.,1,. w. n A V I S O N , T. B. I(ELLER A N D c. S. I J V I N G O O D
1935 Structural cliniiges i n the pituitaries of tIiyroitlc.ctoiiiize(1 r;lts. Am. .J. M w l .
Sci., 1 9 0 : 145-157.
PLATE 1
ESPLANATIOX OF FIGURES
All figures are photoiiiicrograplls ( x 1000) of anterior pituitaries of LongEvaiis r a t s takeii o n Eastnian Conrmercial Ortho fill11 through a Wratten 60 (green)
filter except for figure 1, for which n Wratten 13 (yellow) filter was used.
The (lark cells are thyrotroplis of a 28 day norrrlal male, fixed i n c l ~ r o n ~ - a l ~ n ~
fixative and stained with aldehyde fuchsin and orange G. Although the tliyrotrophs of the r a t are frequently angular and niay have processes exteiicliiig
long distances, they niay also have a rounded shape indistinguishable f r o m
that of other basophils. Kear the left border of the figure is n large tllyrot,roph in which a (kirk Golgi apparatits forms a e:ip over tlie lightly st:iined
nucleus.
Several thyroidertoniy cells may be seen i n the ccwter of this figure f r o m :I
female thyroidectomized 49 days before autopsy, chron~e-alumfixation an(1
PAR-orange G staining. The thyroidect,omy granules are PAS-positive :11ic1
servr to identify the thyroidectomy cells. 'rho dark (iolgi :i1J1xlr:ltus forllls :L
characteristic cap over the light nucleus.
A group of thyrotroplis, two t,ypir:il ones lying :ibove :inti I)elo\v t,lw ccIi1tc.r
of tlie figure, from a normal male pituitary prepared b y the tlircct silvcsr
tec,hnique f o r the Golgi apparatus, followed by chrome-alum fixative. Sections
3 p thick were gold toned nnd stained with P A S :tilt1 o r a l ~ g eG. Tht. (;olgi
:Lpp:ir:itus forins :i characteristic cap over tlie unstainetl nucleus.
Tliyroidc~ctonip cells of :i feiiiale 25 days a f t e r the operation, prep:ird b y
the techniques described for figure 3. A t the center of the figure is a tppic:il
cell with sparse thyroidectoniy granules :ind with the Golgi apparatus c:rpl)inp
the Illicleus.
Norn~aI~ n a l epituitary wit.11 1)reserv:ition of tlic pliospliolipitls by c.oiitrollt.tl
clironuition using dicliromate-subliniate. I n the center :ind further t i p and
slightly to the left are t w o tllyrotroplis showing the typic:il Golgi ap1):ir;itiis
stained with Su d an black.
Tliyroidectoniy cells froin :I frin:ile litter ni:itc of t.lic. snbject of figurc 2 fistvl
f o r pliospIiolipi(1s :IS dewri1)ed tinder figure 5. 'JYir use of the P A S tec.hniquc~,
with or:iiige G , before the zipplicntion of Sudnii 1)l:ick results in retl t,liyroidectoniy graniiles coiitrasting with t l i c 1)l:ick Golpi a1qiar:itus c:ipping tlie
nucleus.
127
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