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Spermatogenesis in a sex-reversed female and in normal males of the domestic fowl gallus domesticus.

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SPERMATOGENESIS I N A SEX-EEVERSED FENALE
AXD I N NORMAL MALES OF' TIIE DOBIESTIC
FOWL, GALLUS DOMESTICUS
RICHARD AVERY MILLER
State University of Iowa
TIIIRTB-FOTJR TEXT B'IGURES AND THREE PLATES (ELEVEN FIGURES)
IXTRODUCTION
Genetical studies indicate that in birds the female is heterogametic, the male homogametic with respect to the sex determining factor. Cytological investigittbns which were undertaken to verify this conclusion by chromosome studies show
a notable lack of agreement in the observations of the different
workers. Table 1summarizes the literature.
Most of the work has been on the Gallinaceae, particularljthe domestic fowl. Several papers report on the pigeon, duck
and parakeet. U r g e r ('36) has recently investigated two
species of passerine birds.
The work on chromosomes may be divided into three periods.
Pr i o r to 1923 the number of chromosomes reported for any
species is small. From 1923 to 1930 the number described
for the fowl falls within a range of 29 to 45, with the average
about 35 to 37. Since 1930 the increase in the number of
small chromosomes observed has raised the total count. Obviously these changes a r e due to improvement in technique as
well as i n accuracy of observation.
A11 investigators a r e agreed that males a r e homogametic,
the female heterogametic. Guyer ('16) reported a n unpaired
element in the chromosome complex of both male and female
Aided by grants from the National Research Council, Committee for Research
in Problems of Sex; grants administered by Prof. Emil Witsehi.
155
g
c1
-______
Saguchi, '30
Kemp, '30
Goldsmith, '28
_-
Akkeringa, '27
Shiwago, '24
Hance, '24
Crew. '23
-
I-
.-
--
-
-
--
_ ~ -_
-
-_
Rhode Island Red,
Leghorn
Ban kiva
Barnenelder
Ankona
Fl Seideii 9 X Bank.
F,Bank. X Seideii
-
-
Oal!us domestieus
RACE OR SPECIES
-~
I
-
-1
_______...
Cutler, '18
__
Boring, '23
(Stevens)
Guyer, '16
Boring, Pearl, '14
LBcaillon, '10
Guyer, '09
Sonnenbrodt, '08
Loyez, '06
AUl'HOR
.
_
~
_
-__
Tissue culture
Tissue eulture
Embryos
__
--
-
.
36-38
29-39, pro. 42
meta. ca. 30
_ _ ~ _ _ _
36
I
I
_
30-34 soma.
~~
32 or more
_
32 spg.
18-20 SI'g.
~
12 I
12 somitic
1
fusion)
14.5 (iiieom.
18-22
a - l o
in pairs)
pi-
No rcduetioii
9
8 bival.
S-16 prs.
N
OHROIOSOMI NUNBER
_ - ~
18 fipg-
I
~
15-19 spg.
prob. 17
__
_
Embryonic gonad
adult testis,
tissue culture
~
Adult testis
Testis
Adult testis,
embryos
Testis
-
__
Gonads, newly
hatched to laying
_. __ Adult testis
--
MATEZIAb
I
TABLE 1
.-
zz
zz
~
~-
Homo:ametic
~-
I-
I
__
_ -
~~
I
~
--
-
-
Largcst chromosome
.
Largest chromosome
~
_
_
Largest chromosome
Second largest
~
Largest chromosome
_____
Unable to confirm
Guyer
~-
Largest chromosome
heterogametic in 6
-
SEX CHROMOSOME
-
_
-1
CL
Q
-
-
_---
-
__
Italian
Plymouth Rock
-
-~
___--
Melcagris dom.
I
--
~-
-
~
__
Sokolow, Tiniakow,
M. gallopavo
Trofimov, '34
___Sokolon, Tiniakow,
M. gallopavo
Trofimov, '36
- _
Cutler, 71s
Pheasant X fowl
Phasiaiius
Phasiaiius
Trofimov,
Tiniakow, '33
colcliicus
- - - _____Sokolow, Tiniakow,
Ph. colchicus
Trofimov, '36
__
-~
Unger, '36
Ph. torqnatus - Sokolow, Tiniakow,
Nyct hemerus
Trofimov. '36
argentatus
Werner, '31
-____-
____I___
-
~
-
--
-~~
-
- -
Embryoiiic
-__ - testis
Embryos
Embryos
- -
Embryonic gonad
-
__.--
_
Sterile male
Embryos
-- -
-
_ _ _ - Embryonic testis,
Emb. membranes
-
Testis : young 8
ovariotomized 0
Testis : adult 8
_Amnion
- -
Embr yoi,
Embryonic testis
Embryo, testis
Embryo, soma
White Leghorn,
Minorka, R. I. Red
Wyandotte,
Emlsryos, soma,
Leghorn X Orpingtor Embryonic testis
Wetlnga X Seiden
White Leghorn
Embryos
Embryos, adult
testis smears
Amnion,
Embryonic gonad
I
_
_
_
I__
2alliis domesticus
Sokolow, Tiniakow,
Several races
Trofimov,-'36- -Miller (present
Buff Orpington
investigation)
Brown Leghorn
Shiwago, '29
Mrleagris
Unger, '36
__
Sokolow and
Trofimov, '33
___
Popoff, '33
White, '30
Suzuki, '30
-
_-
18 -20
20-22
46 ?
49r3
-
-
-
-
- --
76 8
_____
77 0
46 soma.
46 8 soma.
51-60
32-71 most
ca. 50
~-
30-45 meta.
60-70 pro.
74 SPg
7- 3 ~oog.
66 -c 2
1
10-11
38
37
i8
I
I
137 (36
+ Z)
--
1
I
I----
1
ZZ
zz___
iZ
%Z
zz
ZZ
zz
__-___
-
1
I
ZO
Z
I
I
'
I
I
~
-
--
___--__
4th chromosome
4th chromosome
4th chromosome
______
-
-
.
___ -chromosome
Largest and secoud
largest cliromosomcs
4th chrornosomc
-Largest
_.
5th chromosome
--____._
5th chromosome
__ - - _ _ - - -
_____-Idargcst chromosome
Largestchromosome
___I_-
Smallest V-chromosome
- - --__-
Z
Z
Z
Z
I
-I
-1
ZWw
zw
ZO
---
_--
Z
Z
I__
:1
________
Guyer, '09
____
I
-
_
I
_
_
_
Aythya ferina
Cairina moschata
Lampronesa spoiiaa
hfareca peiielopc
Turtur risorius
Columba livia
domestica
C. domatiea
~Columba hybrid
____
C . livia dom.
___-Alias boschas
-
Jentsch, '35
Crew, Laniy, '35
Unger, '36
Unger, '36
Turdus merula
-- -
1
undulatus
M. rindulatus
1 Melopsittacus
-
1 Linota eannabina
- _
_ _-_
-'
-1
I
-
-
~
- -
-
-
16 el.
16 spg.
____16 spg.
62 spg., som.
61 9 (emb.)
60-85
ca. 16
Amnion, embryos,
Embryonic gonad ~
~
Amnion, embryos, 64-85, most
Embryonic gonad
68-75
-~
50-60,
Embryos
ca. 52
Amnion, embryos, 42
Testis
- - -
*
2N
8
38
~-
_
- - ----
lstcyte
1
i
I
2ndcyte
_
4 (occas. 8)
_____
N
______
-_
CHROMOSOME NUMBER
ea. 1G spg.
36-58
_
I
_
Adult testis,
Embryos
- - -_
Eggs
Tehtis
_
Adult testis
-
Embryo,
gonads
Embryonic
- _
Embryonic testis,
membranes
___--
Embryo,
Embryonic gonads
Adult testis
Embryos
Embryonic testis
-
_I_______-
Harper, '04
Smith, '12
Oguma, '27
Guyer, '02
Guyer, '02
$ gchoneberg, '13
CL
-
~
Pavo cristatus
______
Tetrao tetrix
RACX OR SPECIE3
- - - __-__
Numida meleagris
domestica
Sokolow, Tiniakow, Numida meleagris
Trofimov, '36
-Iiidiau Runner Duck
Werner, '27
Trofimov, '36
Sokolow, Tiniakow;
Trofimov. '36
Tiniakow, '34
AUTHOR
I
Z
~
-~
_ _Nale
_ _ 1l Female-
___
SEX
DETERMINATION
__
--
-
Largest chromosome,
-
Largest and second
largest chromosomes
- -
4th or 5th chromosome
I______~
5th chromosome
5th chromosome
-_
5th chromosome
SEX CHROMOSOME
SPEEMATOGEKESIS I N THE DOMESTIC FOWL
159
fowl. IIe stated that in males the unpaired element passes
undivided to one pole in the primary spermatocyte division.
he thonglit,
Later the spermatids wliirh lacked the Z eleme~it,~
degenerated. Therefore the male becomes homogametic in
accordance with the requirements of genetic experiments.
Later work has not confirmed Guyer 's observations.
Werner ('27, '31) reported that in ducks and turkeys the
€emale has one more chromosome, the largest element, than
the male. According to her scheme two classes of eggs
mature, one carrying the Z element, the other a W w complex ;
J V is the largest element, w a chromosome similar to the 2
element.
I n the domestic fowl, Hance ( 'as),Shiwago ( 'as),Akkeringa
( '27), Goldsmith ( '28) , White ( '32), and Popoff ( ' 3 3 ) assert
that the largest chromosome, a bent element with unequal
arms, is unpaired. Eowever, according to Sokolow and
Trofimov ('33) and Unger ('36) it is the fifth chromosome
which is unpaired in the female. This chromosome is a Vshaped element with arms of equal length; it is perhaps the
same chromosome that Suzuki ('30) considered as the sex
chromosome. Sokolow, Tiniakow and Trofimov ( '36) have
summarized their investigations of several of the Galliiiaceae.
I n size and morphology the sex chromosome of these species
resembles the fifth element of the fowl complex. Unger ('36)
reports that the sex chromosome of fowl, ring necked pheasant,
the thrush, Turdus mcrula, and the linnet, Linota cannahina,
are very similar.
I n parakeets Koller (Crew and Lamp, '35) believes that the
largest chromosome, and Jentsch ('3.5) that the second
largest chromosome is unpaired in females. I n pigeons the
largest element is the sex chromosome (Ognma, '27).
L4 majority of the workers assume that the Z elemcnt is
not paired with a 3V chromosome, or leave the question nnsettled. However, Shiwago ('24), Skkeringa ('27), and
Jentsch ('35) believe that one of the smaller elenieiits has
* I n the light of recent investigations it 110 longer seems desirable to use the
symbol XY t o represent heterognrnetic females. Accordingly in our work and in
citing other iiivestigatioiis we have used the symbol ZW.
160
RICHARD AVERY MILLER
the significance of a ITT chromosome. Because of the variability i n the number of small cliromosomes, only by the
observation of the maturation divisions could the identity of
the UTchromosome be established if present. Since the
maturation divisions d o not OCCLIT until the egg is fully formed
and ovulated, technical difficulties have prevented observation
of the chromosomes during meiosis. Harper ('04) reported
sixteen chromosomes in the oogonia and eight in the oocytes
of the pigeon. The sex chromosomes were not identified.
An interesting possibility of observing maturation in the
female is suggested in the work on ovariotomy in the fowl.
Following sinistral ovariotomy, the vestigial right gonad
regularly hypertrophies into a testis-like structure, At the
present time among 208 ovariotomized fowl examined histologically, twenty-four have shown fertile seminiferous tubules
in the right gonad, with germ cells undergoing spermatogenesis. Of these Benoit ('23, '32) found four among thirtythree animals (Zawadowslsy ( '26) two in ten ; Domm ( '29, '30)
nine in 126, and Padoa ('34) nine in thirty-nine ovariotomized
fowl. This paper reports one more case.
None of the previous investigators have considered the
chromosome complexes of these exceptional feniales. Since
such material contains female germ cells undergoing spermatogenesis, it can well be expected to reveal the identity of the
sex chromosomes.
I n birds spermatogenesis has apparently not been studied
in its entirety since the early papers of Guyer ('16) and
Schoneherg ( '13). Hance ( '26) remarks on the difficulty of
finding mitotic figures in adult testes of the domestic fowl,
and Foley ( '28) had little success with the English sparrow.
I n the present study the observations on maturation in tlie
female are supplemented by a study of spermatogenesis in
the normal male.
This problem was undertaken at the suggestion of Prof.
Emil Witschi. The writer wishes to express his sincere appreciation for helpful advice and criticism given during the
course of the investigation.
S P E R M A T O G E N E S I S I N THE D O N E S T I C FOWL
161
XATERIBL AND METHODS
To obtain testes showing mitotic figures we have followed
the technique suggested by Riley’s work on diurnal spermatogenetic cycles ( ’ 3 7 ) . He showed that increased daily rations
of light o r the injection of gonadotropic hormones stimulates
testes of sexually inactive sparrows. When the testes of
such birds were preserved between midnight and dawn he
found mitotic figures in abundance.
A Buff Orpington female was sinistrally ovariotoiiiized 4
days after hatching. Previous t o killing at the age of 10
months and 25 days, the bird was injected for 14 days with
serum of a pregnant mare. One cubic centimeter was given
daily during the first week, arid 2 cc. daily the second week.
The bird was killed a t midnight. The lumbosacral region including the gonads, accessory ducts, and the stump of the
cloaca mas prcserved in Bouin’s fixative. A detailed description of this bird is given in a later section. A series of Brown
Leghorn females ovariotomized at birth is not included in
this paper.
A Brown Leghorn rooster, 11 months old, was killed at
2 AX. after a 2-meek period of injection; 1 cc. (10 R.U.) of
pregnant mare serum was given daily. The germ cells in
this material are in all stages in maturation.
Each of three Buff Orpington males 33 days old a t autopsy
had been injected daily for 4 days with 0.2 cc. of pregnant
mare serum (0.1 cc. equals 10 R.U.), with 20 mp. of a ppridiiie
extract of bull anterior pituitary (8 mg. equals 1 R.U.), o r
with 20 mg. of dried turkey anterior pituitary (25 mg. equals
1 R.U.). Sectioned testes show spermatogonial divisions and
synizesis stages. The bird injected with turkey pituitary
shows fewer divisions.
A fourth male was injected with 20 mg. of dried turkey
anterior pituitary daily for 8 days and was killed at 4 A.M.
on the following moriiing at the age of 37 days. Testes show
enlarged seminiferous tubules but only spermatogonial divisions.
162
RICHAED AVEEY MILLER
A fifth male was injected with 20 mg. of the pyridine extract of bull anterior pituitary f o r 8 days. After an interval
of 3 days, 0.2 cc. of pregnant mare serum was given f o r 10
days and the bird was killed a t 2 A.M. The age at autopsy
was 50 days. Testes were not greatly stimulated. Sections
show gonial divisions and syiiizesis stages.
The sixth male was injected with 0.2 cc. of pregnant mare
serum for 8 days. After a n interval of 3 days, six daily
injections of 20 mg. of the bull extract were made, and following a second interval of 3 days, 20 mg. of turkey pituitary was
given for 3 days and the bird was killed at 2 A.M. The age at
autopsy was 52 days. Testes show all stages including a few
spermatozoa.
Injections were made approximately 6 hours previous to
the hour a t which the bird was to be killed a t the end of the
experimcnt. Dcring the period of injection the length of the
day was increased to 16 hours with 4 hours of artificial
illumination.
Tissues were fixed in Bouiii’s, Allen’s B 15, Champy’s, and
acetic sublimate. Of these Bouin’s regularly gave the best
results and most of tlie observations liave been made from
this material.
After fixation the tissues were dehydrated i n a graded
series of 3570, 50% and 70% diosan in water, followed by two
changes of pure clioxan. Melted paraffin was added over a
period of 2 to 4 hours, the tissue was then transferred to melted
paraffin, changed three times, and embedded. Equally good
preparations resulted from dehydration following tlie aniline
oil technique suggested by Painter ( ’22). Sectioiis were cut
at lop.
Tissues were stained with IIallory ’s, with Delafield’s hematoxylin counterstained with Congo red, with basic fuchsin by
the Feulgen method counterstained with fast green and with
Heidenhain’s iron hematoxylin. With the last stain the seetions were destained in iron alum, o r preferably in a saturated
solution of picric acid as suggested by Tuaii ( ’30).
S P E R MAT OGE NE S IS I N TLLE D O MESTIC F O W L
163
Drawings were made with a Spencer camera lucida and a
Spencer microscope provided with 1.5 apochromatic objective,
aperture 1.3, and a 20 X ocular.
DESCEIPTION O F THE OVARIOTOM IZED FEMALE
At the time of autopsy the ovariotomized female had long
sickle tail feathers. Neck and saddle hackle feathers were
mixed, some of male and some of female type. The comb
was 9.8 em. long and 5.4 em. high; wattles were 5.1 em. long.
Both were bright red. The bird was not observed to crow.
Examination after autopsy showed at the site of the right
gonad a large white, somewhat lobulated, testis-like structure
(fig. 1). After preservation it measured 2.5 X 1.7 em. Study
of the sectioned material shows a testis composed of seminiferous tubules of variable diameter, and cords of germ cells,
or single cells separated by connective tissue resembling the
tunica albuginea of the normal testis.
Covering the gonad is a thin layer of flat cells taking the
place of the ovarial cortex. I n localized regions it appears
thickened, consisting of cuboidal cells several layers deep.
Such patches are connected with the medulla by cords of
connective tissue breaking through the tunica albuginea (fig.
35).
Fertile sections of seminiferous tubules contain all stages
of germ cell maturstion including spermatozoa (fig. 36).
For the most part spermatogenesis is entirely similar to that
observed in our normal niale material, but in some tubules,
development evidently takes an atypical course (fig. 3 7 ) .
There are giant cells which in division show large numbers
of smaIl chromosomes apparently formed by fragmentation.
W e hare also observed spermatids with pycnotic nuclei.
Sterile tubules (fig. 38) are filled with a fibrous cytoplasm
in which n o distinct cell boundaries can be seen. The nuclei
are located about the periphery of the tubules. The sterile
tubules are scattered throughout, the gonad but are more
numerous in the medial half of the gland.
164
RICHARD AVERY MILLER
Ruiining from the region of the right gonad and opening
into the cloaca is a promiiient wolffiaii duct (fig. 1). The
anterior two-thirds is not coiled but is quite irregular because
of annular constrictions. The posterior part is distiiwtly convoluted until just before it joins the cloaca laterally. At the
Fig. 1 Ovariotornixed female, showing right coinpensatory gonad. LO, left
oviduct; RO, riglit oviduct; WD, wolffian duet ; RG, right coinp~iiwttorytestis-like
goiisd ; I-, ureter. Camera lucida drawing.
anterior end the duct enlarges and coils about itself in a.
structure which resembles tlie epididyinis of the male. Cells
lining the tubnles are columnar and ciliated. A ~-ell-de.i.eloped
rete-like formation is visible but there is no open connection
with seminiferous tubules. No sperm were fourid in the wolffian duct.
SPERMATOGENESIS I N THE DOiLlESTIC FOWL
165
On the right side is an oviduct 2.4 cm. long (fig. 1). It is
thin-walled structure with a distinct lumen but without
ostial or cloaca1 opening. The left oviduct is 7.5 em. long
and 3 mm. in diameter through the middle coiivoluted portioii
(fig. 1).
On the left side a t the site of the ovary there is a large
smooth body (fig. 1) which 011 sectioiiing proved to be fat.
No ovarial tissue appears in serial sections, but a small nodule
of lymphoid tissue is attached to the epoophoron. The epoophoron is iiot as exterisive as on the right side, although much
of it may have been removed in the operation. The left
Tvolffiaii duct is continuous to the cloaca. I t s development is
riot as extensive as that of the right wolffiaii duct.
A
CYTOLOGICAL STCDY OF SPERMATOGENESIS
The histology of the riornial avian testis is sufficiently
familiar that i t need not be described here. Figure 39
illustrates the general appearance of a testis in active mitosis.
The course of maturatioii may be followed from the periphery
of a seminiferous tubule, where the spermatogoiiia are found,
t o the lumen. About the lunieii the nearly mature spermatozoa
are arranged in groups attached to Sertoli cells. Spermatogenesis does not proceed in a regular wave, but groups of
adjacent cells mature synchronously ; primary spermatocyte
divisions may occur adjacent to resting spermatogoiiia, o r
spermatids beside primary spermatocytes. Several clifferelit
stages occur simultaneously within a tubnle but no one tubule
ever shows the complete course of maturation. Some stages
a r e apparently of short duration and arc found only after
considerable searching.
The course of spermatogenesis has been followed in tlie adult
rooster, in tlie youiig stimulated males and in tlie ovariotomized female. Spermatogonial divisioiis were difficult t o study
in the small crowded cells of the adult testis. I n the young
males, howover, the gonin are large and the clironiosomes
widely spread. The spermatogonia of the ovariotomized
female were almost as large as those in the ponng stimulated
males.
166
KICHARD AVERY MILLER
The syerinatogonin
There are no significant differences in appearance or behavior of spermatogonia in normal males and in the ovariotomized female. Resting spermatogonia typically show a slightly
oval, lightly staining nucleus lying eccentrically in the cell
(fig. 2). A fine network is visible upon which appear from
place to place small condensations of chromatin. Fine granules
of chromatin border the nuclear membrane. There are one
to several, but usually two, chromatin-nucleoli. Although
these take the usual nuclear staiiis, they do not appear in
2
3
4
Fig. 2 Resting sperrnatogonia.
Fig. 3 Spermstogonia shortly before division, showing two central bodies lying
in the idiozome.
Fig. 4 Spermatogonial division, side view.
Camera lueida drawings. X 2760.
Feulgeii preparations. Between them there is an achromatic
nucleolus, larger than the chromatin-nucleoli.
Shortly before spermatogonia divide, there appears a cap
of dense cytoplasm, the idiozome of Neves, usually on the
side of the nucleus toward the periphery of the seminiferous
tubule (fig. 3). Lying in the iodiozome is the central body.
Spermatogonial divisions proceed much as do ordinary
somatic mitoses. The central body divides into two bodies
which migrate to opposite poles of the nucleus. The metaphase spindle is short and broad (fig. 4). I n the telophasic
reconstruction the central bodies can no longer be seen.
SPERMATOGENESIS IN THE DOMESTIC FOWL
167
Spersnatopaial chromosorn Ps
Spermatogonial metaphase plates (figs. 5 to 8) typically
show a number of small chromosomes grouped in the center
of the field and surrounded by larger chromosomes at the
periphery. Arranged in order as to size, there are six pairs
of large chromosomes, a number of medium sized rod-like
elements, which are of various lengths, some so short as to
be indistinguishable from the small chromosomes lying in
the center of the plate.
The large elements show a distinctive morphology. The
largest of these is a bent chromosome with submedian spindle
fiber attachment. The short arm is a little more than half
the length of the long ~ I I Y L . In some figures the first chromosome tapers from heavy ends toward the point of spindle
fiber attachment (fig. S), but often the chromosome has the
same diameter throughout its length (figs. 5, 8). The apex
is oriented toward the center of the field in metaphase and
toward the poles in anaphase.
The chromosomes of the second pair in order of size are
likewise J-shaped, with submedian spindle fiber attachment.
In this pair the short arm is less than half the length of the
long arm. Orientation is the same as in the first chromosome.
The chromosomes of the third pair are straight, tapering
from a broad end to a feebly pronounced knob-like enlargement at the other end. At anaphase the small end is directed
toward the pole. Spindle fiber attachment is probably terminal. The third chromosome is slightly shorter than the long
arm of the first chromosome.
The fourth chromosomes are J-shaped. Spindle fiber attachment is slightly subterminal but so near the end that in
many cases the chromosomes appear as rods. I n length they
are less than the long arm of the second chromosome, or
about half the length of the first chromosome.
The chromosome of the fifth order is the only V-shaped
element in 1,he entire complex. It is slightly shorter than the
chromosomes of the preceding pair, but its morphology is
entirely distinctive. Spindle fiber attachment is median. I n
THE b N A T D Y I C A L RECORD, VOL. 711, SO. 2 ANT) S Z ' P P T J X E P T N O 2
168
RICHARD AVERY MILLER
male complexes there are two of these elements ; in spermatogonia of the ovariotomized female we have never found more
than one chromosome of this size and shape. This clement
then, is the Z chromosome.
The sixth chroinosonies are rods which taper slightly. The
smaller end i s directed toward the center of the metaphase
plate. They are shorter than the short arm of the first chromosome but longer than that of the second chromosome.
The seventh pair are rod chromosomes of uniform diameter.
They are half as long as the sixth pair. This pair of chromosomes is the first of the rod-like elements of medium size. It
and the others of this group differ from the preceding chromosomes in that they show no distinctive morphology.
The material does not permit any conclusive statement as
to the number of chromosomes. Spermatogoiiial figures from
both sexes show a broad variation, fifty-one to sixty in the
best preparations. The variation, as reported by other investigators, is always in the number of small elements.
Primary spermatocytes
Resting spermatocytes are found against the wall of the
tubule or close to it. Their general appearance is similar to
that of spermatogonia. There are fewer granules along the
nuclear membrane, and the network on which appear the
chromatin condensations is finer. The idiozome which surrounds the central body at this time is a thick cap on one side
of the nucleus. ,4s maturation proceeds it spreads along the
wall of the nucleus to occupy half its circumference. Chromatic
and achromatic nucleoli arc unchanged in appearance. While
Figs. 5 a n 6 Polar views of sperinatogoiiial divisions i n the testis-like gonad
of the ovariotomized female.
Figs. 7 and 8 Polar views of sperinatogonial divisions in the testis of a young
Buff Orpington male.
Figs. 9 and 10 Polar vicws of primary spcrmatocyte dirisioiis in the testis-like
gonad of the ovariotornized fcmale.
Figs. 11and 12 Polar views of primary spermatocj te divisions in the testis
of the Brown Leghorn rooster.
Camera lucida drawings. X 1900.
169
SPERMATOGENESIS IN THE DOMESTIC POWL
7
9
Figures 5 t o 13
170
R I C H A R D AVERY M I L L E R
still in position on the ~7allof ihe seminiferous tubule, the
spermatocytes grow to thcir definitive size (fig. 13). Now begin the characteristic changes in the nucleus. Chromatin
nucleoli disappear and the iiucleus stains more heavily. The
nuclear network increases in complexity, the threads gradually
becoming coarser (fig. 14). At the same time the threads
begin t o draw away from the periphery of the nucleus and
form a tangled mass, from which fine chromatin threads project out t o the wall of the nucleus (fig. 15). A little later the
tangled mass of chromatin threads moves to that side of the
nucleus against which lies the idiozome. At this time synapsis
occurs. The details of the pairing could not be determined,
but when entirely polarized, the strands of chromatin are thick
and coarse (fig. 16). During these changes, the achromatic
nucleolus is visible between the strands of the fine network,
and later when the chromatin has drawn t o one,side, it may
be seen in the clear area of the nucleus.
Now follows a process of unravelling so that the chromatin
comes to occupy the entire nuclear space in what appears to
be a continuous strand (fig. 17). Occasionally the thread is
split, indicating its double nature.
By the frequency with which it is seen, the pachytene stage
appears t o persist for some time. Eventually, however, the
chromatin strands lengthen and become thinner. The nuclear
space is now filled with the intricately interwoven thread
which in extreme cases approximates a network. Occasionally
this thin thread seems t o be split. The achromatic nucleolus
persists. This stage seems t o be a feebly accentuated diffuse
stage (fig. 18). Diiring this period the central body divides
into two which remain close together.
The fine chromatin thread now rapidly contracts. The
single thread splits into two, which are twisted about each
other in a strepsinema (fig. 19). As contraction continues,
the coiled thread breaks up into short lengths which become
the tetrads of diakinesis (fig. 20). The bivalent tetrads now
coalesce and thicken to form the prophase chromosomes of the
first spermatocyte division. The central bodies remain close
13
14
15
16
17
18
0
20
19
Fig. 13 Resting apermatoeyte.
Fig. 14 Early spireme.
Fig. 15 Synaptene stage.
Fig. 16 Bouquet stage.
Fig. 17 Pachytene stage.
Fig. 18 Diffuse stage.
Fig. 19 Strepsinema breaking up into tetrads.
Fig. 20 Diakinesis.
Fig. 21 Primary spermatocyte division normal male, side view.
Camera lucida drawings. X 2760.
171
172
RICHARD AVERY NILLER
together until just before divisioii and then rapidly separate.
The metapliase spindle is iiai-1-ow, aiid the central bodies relatively farther apart than in gonial divisions (fig. 21). During
these changes the hctcrochromosome is not distinguishable by
pycnosis or any peculiarities of beharior.
Sperm at ocyt e ch Y’ o m o s o ~nP S
I n figures 9, 10 and 40 are shown the first spermatocyte
chromosomes of the female, and in figures 11 xiid 1 2 those of
tlie male. The resemblance between the spermatocyte and
spcrmatogonial chromosomes is obvious. The bivalcnt spormatocyte tetrads are much heavier and f o r that reason the finer
structural details, such as the knob-like enlargement of the
third sperniatogonial chromosome, are not apparent.
The fifth chromosome requires special comment. I n male
spermatocytes it is a thick tetrad only slightly bent. On the
contrary in the metaphase plates OP female material, the fifth
chromosome closely reseinbles tlie fifth spermatogonial elemcnt. I t is slender, whereas the other cliromosonies are thick,
indicating that these are the result of conjugation of homologous goiiial elements. Since the fifth chromosome is unpaired
iu the female, it retains its gonial appearance.
In the female, the behavior of the fifth chromosome during
the first spermatocyte division is significant. This chromosome passes uiidivided to one pole of t h e spindle. T t has never
been seen to divide. Furthermore it is riot paired on the
equatorial plate with any of the smaller chromosomes. It may
regularly be seen migrating to one pole of the spindle while
the rest of the field remains in the iiietaphase condition (figs.
22, 23, 41). In extreme cases it has moved to a position betmeeii the central body aiid tlie cell wall while the autosomes
are just beginning their anaphase movement (fig. 24). Ti?
nialc spermatocytes the fifth tetrad segrcgates in the usual
manner.
The primary speriiiatocpte chromosomes in the ovariot omized female were especially favorable for counting. Good
preparations in which tlie small elements are well stained
SPERMlTOGENESIS I N THE DOMESTIC FOWL
173
regularly show forty chromosomes. Occasioiially thirty-eight
or thirty-nine are found. I n these cases it is probable that one
o r two of tlie small elements are concealed by the large ones.
I n male spermatocptes, where for some reason the chromosomes were not as well spread as in the female, counts gave
accordingly smaller numbers.
Secondary spe,rmatoc?jtes
Resting secondary spermatocytcs are slightly smaller than
spermatogonia. The nucleus shows several scattered clumps
of chromatin (fig. 2 5 ) . The central body lies in the idiozome.
Figs. 22 t o 24 Side views of primary sperrriatocytc divisions in the ovariotomized
female ; shows characteristic appearance of thc fifth chroinosorrie as it passes
uiidividetl t o oiie pole of the spiiidle ahead of the autosomes. Camera lucida
drawings. X 2500.
The resting stage is of short duration, followcd by the seconda r y spermaiocyte division. Cliromosomes in the inetaphase
plate are closely crowded. Five to seven large chromosomes
and several smaller elements may be seen in the better preparations (fig-. 26). The spindle is long and narrow (fig. 27).
There is no evidence of a second pairing as reported by Guyer
( '16).
Spermiogenesis
The nucleus of a resting spermatid shows one t o three
chroniatiii spheres aiid a scattering of granules along tlie
nuclear membrane. A small achromatic iiucleolus is present.
The central body lies in a small indistinct idiozome (fig. 28).
174
R IC HAR D AVERY MILLER
The first iiuclear change in spermiogenesis is a spinning
out of the chromatin spheres into a network and an accurnulation of chromatin along the nuclear membrane (fig. 30). The
network of chromatiii material then begins to contract
laterally. At first there is only a slight elongation iii the
anterior-posterior direction, which steadily increases as contraction diminishes. At this time the sperm head is seen
lying in a clear space surrounded by a membrane (fig. 31).
Shortly after contraction ceases the sperm head becomes attached t o a Scrtoli cell. The head grows enormously long and
tapers t o a fine point. The chromatin network becomes more
and more compact and eventually the sperm head stains homogeneously (fig. 32). At about the time the head has achieved
its maximum length, the membrane becomes closely applied to
the sperm liead which shortens (fig. 33). Shortly afterward,
spermatozoa are released into the lumen of the seminiferous
tubule.
Only a few of the details in the formation of the tail could
be observed. In the resting spermatid a central body lies in
a small idiozome. Later as the chromatin spins out into a
network, the central body elongates into a rod which will
eventually form the tail filament (figs. 28, 29). At first the
rod lies tangentially to the nucleus but later turns and moves
toward it until one end touches the nuclear membrane. A
little later the end appears to have entered the nucleus (fig.
30). The rod increases in length, the posterior end passing
out of the cell. As it elongates, it becomes thinner and stains
less and less intensely so that in later stages it may be seen
only with great difficulty. At the time when the membrane
enclosing the sperm head appears there is seen a fine thread
~
_ __
_
~
~
Fig. 25 Resting secondary spermatocyte.
Fig. 26 Secondary spermatocytc division, polar view.
Fig.27 Same, equatorial view.
Fig. 28 Resting spermatid.
Figs. 29 to 33 Several stages in the trailsformation of the spermatid into the
spermatozoon. (In figs. 28 and 29 oiily part of the tail filament is represented.)
Fig. 34 Spermatozoon free in lumen of a seminiferous tubule.
Camera lucida drawings. X 2760.
SPERMATOGENESIS IN THE DOMESTIC F O W L
25
1'75
27
26
28
32
31
2
33
Figures 25 t o 34
176
RICHARD AVERY MILLER
which leads away from the sperm head, and passing through
the enclosing membrane appears to be continuous with the tail
filament (fig. 31). At this point on the membrane there is a
small mass of darli-staining material whose origin could not
be determined (fig. 31). Later this material is 110 longer
visible. About the time the mcmbraiic flattens against the
sperm head, the tail becomes encased in a sheath (fig. 33).
The acrosome is formed from itliozome material wliich
migrates around the nucleus lo the anterior end before eloiigation begins (fig. 30). Spermatozoa ih the seminiferous tubules
show acrosomc, head and tail (fig. 34).
DISCUSSION
Breeding experiments on the inheritance of sex-linked charis of the ZZ-ZO
acters iii birds prove that sex deteri~~ination
or ZZ-ZW type. Until recently the cytological evidence for
this conclusion has not been satisfactory. I n fowl hoth the
largest and the fifth largest chromosomes have been designated as thc sex chromosome and both conclusions have been
repeatedly corroborated. I t is significant that tliosc who helieve that the first chromosome is unpaired in the female have
not reported a consistent individual morphology f o r the
largest elements. If the pairiiig of the chromosomes is based
upon the determination of their length, the well recognized
difficulty of accurate measurement may account for what now
appears t o be an erroneous conclusion.
Sokolotv and Trofimov ( ' 3 3 ) have described and their
figures show a consistent individual morphology for the sereii
largest chromosomes. With this important advance t h e sex
chromosome was identified as the fifth largest element in tlie
complex.
IVhile the investigations of the Russian workers is admirable
one must still admit that in material as difficult as avian chromosomes, there remains tlie danger of individual interpretation in making up series of paired chromosomes. For a
complete case, observation of the appearance and behavior
of the heterochromosome during maturation divisions is highly
SPERMATOQENESIS I N TIIE DOMESTIC FOWL
177
desirable. Obviously oiily such evidence could prove the
presence or absence of a W chromosome. I n species with the
more common male heterogamety this would be a simple
matter, but in avian material the technical difficulties of observation are almost insurmountable in the normal female.
Fortunately in fowl there exists the unusual opportunity in
the potentiality for male differentiatioli of the vestigial right
gonad of the female.
Witschi (’32) has shown in amphibians that the cortex and
medulla of the gonad are sources of substances inducing respectively male and female differentiation of the germ cells.
I n addition, cortical inductor inhibits male differentiation,
while medullary inductor snppresses female differenti at‘ion.
In fowl XTitschi ( ’35) bas concluded that thew is a “priniaqr
hereditarily fixed deficiency in the right cortical inductor. ? ’
This deficiency is manifested in the infrequency with which
the germ cells are attracted to the cortex at the time the germ
cells are di stribntecl during embryonic development. As a
~ e s u lthe
t right gonad is primarily a medullary structure with
occasionally a cortical rudiment. It is elcar that the presence
of the left ovary is iiistrumental in suppressing development
of the right gonad, siiice upon ovariotomy the right gonad
hypertrophics into a testis-like structure. This compensatory
growth of the right gonad recalls the reverse result obtained
in the experiments of Harms ( ’23) and Ponse (’24) on Bidder’s organ in toads. I n all cases following extirpation of the
testis, Bidder’s organ (a cortical rudiment) hypertrophies
into a functioiial ovary in the male.
I n the fowl any germ cells present iii the right gonad at the
time of sinistral ovariotomy then undergo male differentiation.
Rrocle (’28) believes that germ cells persist in the riglit gonad
up t o 3 weeks after hatching aiid then degenerate. However,
Padoa (’34) reports that riglit gonads showing spermatogenesis occurred with equal frcqnency in birds ovariotomized
between 3 and 6 months of age and in those operated between
18 and 23 days after hatching.
178
RICHBED RVEEY MILLER
Save for the presence of a well-developed cortex, the left
ovary is essentially similar to the right gonad. If the cortex
is surgically removed, the medulla is able to respond with
testicular development. Benoit ('23)' Zawadowsky ('26 b),
Domm ('27), and Padoa ('34) have noted in ovariotomized
fowl that there is a development of the remaining medulla
into a sterile testis.
We have shown that these germ cells which are differentiating into spermatozoa still retain the female chromosome complex. Obviously differentiation in the male line is not accompanied by any change in the female chromosome pattern.
Such evidence proves that differentiation of the germ cells
in these cases is controlled by the somatic medulla and not
directly by their chromosomal makeup.
Similar considerations have been derived by Witschi ( '29)
from his genetical analysis of adult hermaphrodites of the
grass frog, Rana temporaria. He artificially fertilized eggs
from hermaphrodites with normal sperm, and normal eggs
with sperm of the hermaphrodites. The sex ratio of the
progeny demonstrated that both eggs and sperm of the
hermaphrodite were genetically female. Crew ( '21) also
gave proof that the testes of adult hermaphrodites were
genetically female. On cytological examination Witschi ( '24)
reports a diploid number of twenty-six in spermatogonia and
oogonia of the hermaphrodites and a haploid number of
thirteen. Stohler ('28) also found that Bidder's organs and
the functional gonads of both sexes have the same chromosome complexes.
As yet no genetical analysis of the ovariotomized females
has been made. Domm ( '30) artificially inseminated several
normal females with motile sperm obtained from the right
gonad of ovariotomized fowl. All the eggs laid were infertile.
His technique was successful when sperm of normal males
was used. Domm suggests as an explanation for the negative
results that since the sperm were taken from the gonad they
may not have been physiologically mature. Other workers
have reported that spermatogenesis is atypical in the right
SPERMATOGENESIS IN THE DOMESTIC F O W L
179
gonad and that the spermatozoa are not normal. However,
Zawadowsky and Zubina ( '26) have a drawing of spermatozoa
in the seminiferous tubules of their hen Vega which appear
normal. In our material some of tlic spermatozoa are structurally quite normal.
It may be concluded that sex reversal in ovariotomized
fowl, as in adult hermaphrodite frogs, does not involve a
change in genetic constitution.
SUMMARY AND CONCLUSIONS
1. A Buff Orpington female sinistrally ovariotomized 4
days after hatching developed a testis-like structure on the
right side. The bird was stimulated with gonadotropic hormones and was killed during the night, when mitotic divisions
are most apt to occur in the testis. The appearance and behavior of the chromosomes during maturation are compared
in this ovariotomized female and in testes from siniilarly
stimulated males.
2. A study of spermatogenesis in normal males and in the
ovariotomized female is reported.
3. The form of the large chromosomes is described in
spermatogonial and spermatocyle divisions. The fifth chromosome in order of size, a V-shaped element with median spindle
fiber attachment, is the sex chromosome. I n male spermatogonia the element is paired; in female spermatogonia it is
single.
4. The fifth spermatocyte tetrad in males is a heavy element
formed by the pairing of homologous spermatogonial cliromosomes. In the female the corresponding element is a slender
chromosome resembling the fifth spermatogonial chromosome.
5. The fifth spermatocyte chromosome in the female passes
without dividing to one pole of the spindle ahead of the other
chromosomes. I n males the fifth tetrad segregates, one chromosome moving to each pole.
6. The exact iiumber of chromosomes could not be determined. Spei-matogonial counts vary between fifty-one and
sixty. Spermatocyte chromosomes are more easily- counted.
180
RICHARD AVERY MILLER
I n good preparations thc number is between thirty-eight and
forty. Thc number forty is found most frequently and probably represents the true monoploid chromosome number.
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T H E A X A T O X I C A L RECOBD, \ . 0 1 ~ . ‘70,N O . 2 A N D SUPPLHMEYP X O . 2
PLhTE 1
EXPLA4NA4T1OXOF FI GU R E S
35 Thick~nedcortex in the right gonad of the owriotomizpd female.
36 Section of seminiferous tubule of the right gonad, showiiig spermatozoa.
37 Section of seminiferous tubule in the right gonad shoring sereral primary
spcrmatocyte divisions aiid degenerated cells in the center of the tubule.
Photomicrograph. X 310.
184
SPERMATOGENESIS I N T H E DOMESTIC FOWL
RICHARD AVEBY MILLER
185
PLATE 1
PLATE 2
EXPLANATIOX OF FIGURES
38 Sterile seminiferous tubules i n the right gonad of the ovariotomized female.
Photomicrograph. X 130.
39 Seminiferoiis tubule of the Brown Leghorn rooster, showing inany of the
stages in maturation. Photomicrograph. X 210.
40 Photomicrograph of a primary spcrinatocyte division from the ovariotomized
female. The samc cell is reprcsentcd in figure 9. X 1930.
41 Side view of a primary spcrniatoeyte division iii the ovariotomized female.
The saiiie cell is rcprcseiited in figure 22. Photomicrograph. x 1300.
186
SPERYATOGEKESIS IN THE DOMESTIC FOWL
RLCHARD AVLBY MIT.LER
187
PLATE 2
?LATE 3
EXPLANATION O F FIGURES
We are indebted t o Ilr. Max Poser of the Bausch and Lomh Optic a1 C o . f o r tlic
photugraphs on this plate. They were not receired until a f t e r the engrsvingr had
been made a n d f o r this reason figures 40 and 44 are duplicates.
42 Polar view of a spermatogonial dil-ision i n the testis-like gonad of the
orariotoinized female. The same cell is represented in figure 6.
4:3 Polar Tiew of a spermatogonial division in the testis of a young Huff
Orpington male. Kepresented in figure 8.
44 Polar %iewof a primary sperinatocytc diTision in the ovariotomized female.
Keprcsented in figure 9.
45 Side view of a primary sperniatoq te division in the oLariotornizrd femalc.
Represented in figure 24.
Photomicrogrxphs, X 4500.
188
SPERAfATORESESIS IN THE DOMESTIC FOWL
RICHARD AYERY JIILLER
PLATE 3
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sex, reverse, fowl, malen, female, spermatogenesis, domestic, norman, gallus
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