A C T A
T H E R
V O L . X I I I , 3 2 : 511— 551.
I O L O G I C A
B IA Ł O W IE Ż A
K rz y sz to f
2 8 .X II .1 9 6 8
S W I E Ż Y Ń S K I
The Male R eproductive Organs of the European B ison
B tson iana X X X V i n
[W ith 3 F igs., 10 T a b le s a n d P la te s XV—X IX !
The dimensions and weight of the testicles, and also th eir shape and
position, w ere described on the basis of dissections of the cadavers of
20 European bison of d ifferent age. Corresponding data were ascertained
for the epididym is, ductus d e fe r e n s , genital accessory glands, urethra
and penis. U teru s 7nasculitius w as found to occur in m ale European
bison. No d istinct relation was found betw een the degree of developm ent
of the la tte r and the form ation of any of the other genital organs and
the reproductive capacity of the m ales of this species. A description is
given of the stru cture of the scrotum , prepuce and of m uscles topo­
graphically and fu nctionally connected w ith the genital organs. In com ­
parison w ith domestic ca ttle the testicles of adult European bison are
more spherical and th eir m ean weight is sm aller. The epididym is has
a strongly m arked, wide head and thickened tail set caudad. Am ong the
genital accessory glands the bulbourethral glands d iffer as to position,
and are sm aller than those in domestic bulls. The scrotum in the E u ro­
pean bison is situated m ore to the back, is relativ ely short, w ith fain tly
defined collum . The le ft part of the scrotum , together w ith the testicle,
protrudes slightly forw ards. The sigmoid flex u re of the penis is only
fa in tly m arked.
I.
II .
III.
IV .
V.
V I.
V II,
V III.
IX .
X.
X I.
Introduction
.......................................................................................................................612
M aterial and m e t h o d s ........................................................................................................512
The t e s t i c l e s ............................................................................................................................. 513
T h e epididym is, the ductus d e fe r e n s and sperm atic cord . . . .
516
The scrotum and covering structures of the testicles, the epididym is
and the sperm atic c o r d s ...................................................................................................... 510
The genital accessory glands
1. A m p u lla e d u ctu u m d e f e r e n t i u m ..........................................................................521
2 . V e sic u la e s e m i n a l e s ........................................................................................................522
3. The p r o s t a t e ...................................................................................................................... 523
.........................................................................................524
4. The bulbou rethral glands
The u te ru s m a s c u l i n u s ...................................................................................................... 525
T he pelvic part of the urogenital d u c t ....................................................................... 529
The p e n i s ..................................................................................................................................... 530
1 . Corpus p e n i s .......................................................................................................................530
2. The penile p art of the urogenital d u c t ...................................................................531
3. D im ension of the p e n i s ................................................................................................ 533
The p r e p u c e ..............................................................................................................................533
The preputial and the extren sic penile m uscles . . . . . . .
535
[5111
K . Św ieżyński
512
X I I . R esu lts and d i s c u s s i o n ............................................
1. T esticles, epididym is and scrotum .
.
.
2. The genital accessory g l a n d s ..............................
3. U teru s m a s c u l i n u s ....................................................
4. P e n i s ...........................................................................
5. T h e preputial and extren sic penile m uscle .
R eferences
.
.
.
Streszczenie .
.
.
E xplanation of plates
I.
537
537
541
542
545
546
547
549
550
IN TRO DU CTIO N
L ite ra tu re dealing w ith the anatom y of the m ale genital organs of the E uro­
pean bison is scanty. From the historical aspect m ention must firs t be made of
the w ork by B o j a n u s (1827), who gives some rem arks on the appearance of the
scrotum and testicles.
O w e n (1848) in describing the structure of the internal organs of the m ale
European bison lim its h im self to the laconic statem ent »...the scrotum and testes
were m uch sm aller than in the young dom estic bulls of the same age; the scrotum
is rugous, senile, not pendulous, w ith a constricted neck as in B o s tau ru s«. M ii I1 e r (1852) in his study »R em arks on European bison« describes in the section
devoted to anatom y the testicles, v a s d e fe r e n s , sem inal vesicules and u teru s m a s ­
cu lin u s o f »unusual stru ctu re not hitherto described in any species«. In fu rth er
discussion on the su b ject of this organ the author reached the conclusion that it
was a case not exh ibiting abnorm al ch aracteristics.
K r o l l i n g (1930) had a t his disposal p art of the genital organs of a m ale bison
which died in Schonbrunn Zoo. Using histological slides m ade from the organs of
the pelvic part of the m ale gen ital system , he described in detail the structure of
the term inal portions of the deferens ducts, v e s ic u la e se m in a les and u teru s m a scu lius, w hich occurred in th is individual also. R em arks on the appearance of the
extern al genitalia of the European bison are also to be found in the monograph
by W r ó b l e w s k i (1927).
The aim of the present study is to elaborate as com prehensively as possible the
anatom y of the m ale reproductive organs of the European bison. The author hopes
that the results of his investigations, in addition to their value as a source of
inform ation on the morphology, m ay also be of use in other studies on the species,
such as its physiology, in tergeneric hybrids or breeding operations.
I I . M A T E R IA L AND M ETHODS
The m aterial used consisted of the cadavers of 20 bison which reached the
Research C entre of A natom y of the European bison during the period from 1950—
1966. These were purebred European bison varying in age from one day to
seventeen years of age.
In order to give a fu lle r description of the anim als dissected some of the e x ­
ternal body dimensions have been included in the details of m aterial, thus m aking
it possible to com pare the size of the various individuals (T able 1). M easurem ents
w ere m ade on cadavers which had not yet been fixed . The m easurem ent points
and way in which the m easurem ents w ere made, and term s used for the m easur­
em ents, w ere the same as those hitherto used in the W arsaw C entre (P i e k o s et
a I., 1958).
L in ear m easurem ents w ere m ade by means of a soft surveying tape m easure,
ru ler w ith m illim etre scale, com passes, nonius and m icrom eter. The m easurem ent
513
The m ale reproductive organs of the European bison
points of the various organs are illustrated in Fig. 1. W here necessary the way in
which the m easurem ent w as made is described together w ith the description of
the appropriate organ.
The weight of the organs was found on a laboratory balance.
As the degree of bleeding or its absence and the method used fo r fixin g were
of necessity d ifferent, absolute figures fo r m easurem ents and w eights of organs
should be treated as approxim ate data.
Some dissections w ere made from fresh m aterial, but the g rea t m ajority from
fixed m aterial. F ix in g was carried out by storing the isolated organs in 7—3°/o
form alin w ater solution or by in tra a rteria l in jection of the w hole body by the
method used in the Centre ( P i l a r s k i et a I., 1967).
T able 1.
M aterial examined.
No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
Name
Pos
Pub
Pom
P la k a t
Pustybór
Poronin
P utn ar
Puck
Puzor
Pug II
Pozew
P uer
P usteln ik
Połam iec
Pokorny
Posusz
Pluszcz
P lam iec
Pom ruk
P lato
No. of
pedigree
book
**
*♦
1364
**
906
1459
1569
1151
1208
1087
1082
934
694
572
1077
984
785
789
816
575
Body
e xtern al m easurem ents
Age
Years
Months
Days
—
—
—
—
—
—
2
3
4
5
6
6
7
8
9
9
10
10
14
14
17
1
10
—
3
—
7
1
4
—
4
2
1
1
5
4
7
3
1
7
10
17
8
24
22
3
25
13
27
21
5
26
12
3
IS
—
28
A
116
114
107*
132
183
212
247
247
275
280*
250
—
269
260
283
273
248
262
—
280
D
_
86
—
104
135
156
171
179
185
184*
170
—
170
183
177
202
—
172
—
190
E
F
45
45
44*
67
105
95
128
112
130
125*
120
—
—
125
133
129
—
126
—
135
* — M easurem ent made after rem oving skin; ** — The pedigree
European bison born in these years have not yet been published.
53
62
75*
89
115
137
178
162
185
189*
190
—
—
—
220
206
—
180
—
160
books of
W here the state of the cadavers, on account either of advanced decomposition
or dam age caused by anatom ical and pathological dissection, prevented th eir being
used as a w hole, only certain o f the organs were taken for detailed exam ination.
The term used in this study »adult bison« refers, in accordance w ith th e results
of investigations made by R o s k o s z & E m p e l (1963), to individuals over 5 years
old.
III. TH E T E S T IC L E S
T h e te stic le s of th e m ale E u ro p e a n bison a re o v al in sh ap e. T h e long
a x is of th e te sticle s in ad u lt an im als is situ a te d d ecid ed ly v e r tic a lly . T h e
514
K . Sw iezynski
m edial s u rfa c e is fla tte r th a n th e la te ra l. T h e ep id id ym is in th e c a d a v e r
is ad h e re n t to th e ca u d o la te ra l su rfa c e of th e te ste s.
In tw o c a lv e s, one o n e -d a y old and one se v e n -d a y s old, th e te stic le s
w ere situ a te d close to th e e x te r n a l ingu in al rin g , th e long a x is b eing
alm o st h orizo n tal. T h e th re a d -lik e lig a m e n tu m
in g u in a le testis, a b o u t
Fig. 1. P lan of distribution of m easuring points.
A, B — testicles; a — la tera l su rface; b — cran ial surface. 1 — 3 length of
testicle; 2 — 4 breadth of testicle; 5 — 6 breadth of proxim al root of caput e p id id y m id is ; 7 — 8 breadth of d istal root of ca p u t e p id id y m id is ; 5 — 8 breadth of
base of ca p u t e p id id y m id is.
C — u teru s m a scu lin u s (dorsal surface); a — undivided p art; b, c — le ft and
right horn; d — co rp u s p r o s ta ta e . 1 — 2 length of le ft horn; 2 — 3 length of right
horn; 3 — 4 length of undivided part; 5 — 6 transverse m easurem ent of prostate.
D — bulbus urethrae and b ulbou rethral glands (caudal surface); a — bulbo­
urethral glands; b — bulbus u reth rae; c — r a d ix p en is. 1 — 2 tranverse m easure­
ment of bulbourethral glands; 1 — 3 longitudinal m easurem ent of bulbou rethral
gland; 4 — 6 length of bulbus u reth rae', 5 — 7 breadth of butbus u r e th r a e in
widest place.
E — m ale genital organs (seen from le ft side); a — u teru s m a s cu lin u s; b —
am pulla ductus d e fe r e n tis ; c — prostate; d — vesicula se m in a lis ; e — pars p elv in a
u r e th r a e ; f — m . b u lb o c a v e r n o s u s ; g — m. is c h io c a v e r n o s u s ; h — penis; i —
ductus d e fe r e n s . 1 — 3 length of ductus deferens; 2 — 3 length of a m p u lla ductus
d eferentis; 4 — 5 longitudinal m easurem ent of v es ic u la s e m in a lis ; 6 — 7 transverse
m easurem ent of vesicula s e m in a lis ; 8 — 9 total length of penis; 9 — 10 length of
p a rs p r e p u tia lis p en is.
The m ale reproductive organs of the European bison
515
3 cm . lon g, ru n s fro m th e tail of th e ep id id ym is of e a ch o f th e te sticle s
to th e c a u d o v e n tra l in n er s u rfa c e of th e s c ro tu m w a ll. In » P la k a t« , one
m o n th old, th e long a x is of th e te stic le s ra n obliqu ely. L iga m en to , in gu in a lia te s tic u lo ru m w e re f a r b ro a d e r th a n in th e p re v io u s te sticle s.
T ab le 2.
M easurem ents and w eight of testicles according to age groups (cf. tab le 1).
M easurem ent in m illim etres
No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
18
17
W eight in gram m es
Testicle
Testicle
Name
L eft
Right
R ight
L eft
12
16
15
1.55
2.00
3.00
1.75
1.50
3.00
26
18
2.17
2,07
14
19
34
36
16
18
3.10
8.00
3.50
7.50
36
17
35
17
5.55
5.50
Poronin
P utnar
86
91
53
50
92
90
62
44
120.00
128.00
106.00
100.00
Mean
89
51
91
53
124.00
103.00
Pug II
Pozew
Puer
P ustelnik
Połam iec
Pokorny
Posusz
Pluszcz
P lam iec
Plato
96
94
95
114
115
104
123
104
102
87
52
48
50
53
62
60
24
59
71
55
88
100
108
121
113
112
97
105
94
95
50
47
45
55
59
61
43
57
66
69
110.00
121.00
185.00
200.00
287.00
265.00
90-00
230.00
—
135 00
96.00
124.00
155.00
208.00
279.00
272.00
104.00
240.00
—
139.00
Mean
101
57
104
57
191 63
189.13
Length
Length
B readth
Pos
Pub
Pom
23
26
29
11
15
14
24
25
28
Mean
26
13
P la k a t
Pustybor
34
38
Mean
Breadth
T h e te sticle s o f th e 1 0 -m o n th s old » P u sty b o r« still re ta in e d th e tra n s ­
v e rs e position and th e ingu in al lig a m e n t w as only fa in tly d iffe re n tia te d
in th is bison. In all th e ab o v e-m e n tio n e d bison th e le ft te stic le p ro tru d ed
s lig h tly in fro n t of th e rig h t te sticle . D iffe re n ce s in th e size of th e
te s tic le s b etw een ind ivid u als in th is ag e g ro u p a re re la tiv e ly sm all, b ut
th e w eig h t o f th e te stic le s of 1 0 -m jn th s old » P u sty b o r« w as alm ost
tw ic e as g re a t as th a t of th e te sticle s of o n e -m o n th old » P la k a t« .
/
K . Sw iezyñski
516
The d a ta g iven in T ab le 2 show th a t th e d im ensions, and in p a rtic u la r
th e w eig h t of th e te stic le s of E u ro p e a n bison in th e o th e r ag e g ro u p s,
a re c h a ra c te riz e d b y co n sid erab le in d ivid u al v a ria tio n . T h e d im ension s
and w eig h t of th e te stic le s of » P u ck « , »P osu sz« and » P o m ru k « w e re n ot
tak en into co n sid eratio n w hen ca lc u la tin g m e a n v a lu e s as th e firs t e x ­
hib ited b ila te ra l an d th e o th ers rig h t-sid e d cry p to rch ism u s.
T u n ica a lb u g in e a te sticu lo ru m in a d u lt E u ro p e a n bison, to g e th e r w ith
th e e p io rch iu m , h as a m ean th ick n ess of 1.24 m m (m a x 1.46, m in 1.05
m m ). It is clo sely k n it and from it n u m ero u s tra b e cu la e of co n n e ctiv e
tissu e pass into th e in te rio r of th e glan d . No d istin ct in te rlo b u la r sep ta
can be seen m acro sco p ically .
M e d ia stin u m testis in th e fo rm of an a x ia l stra n d of co n n e ctiv e tissu e
fro m 1 to 4 m m w id e, e xten d s alon g th e lon g a x is fo r 3/4 of its len g th ,
sta rtin g fro m e x t re m it a s capitata. It is situ a te d a lm o st ce n tra lly w ith ,
a ten d en cy to a p p ro ach th e ep id id ym al s u rfa c e (P h o to 2 -B , C an d 3).
D u ctu li e f f e r e n t e s ru n n in g fro m th e m ed iastin u m testis, su rro u n d ed
b y a b u n d an t co n n ectiv e tissu e, le a v e th e te stic le n e a r the u p p er pole.
T h e y p e n e tra te into th e head of ep id id ym is on th e c o n v e x te s tic u la r
s u rfa c e of th e p ro x im a l a rm of th e h ead of ep id id ym is, n e a r its d o rso cau d al m arg in .
IV. E P ID ID Y M IS , DUCTUS D E F E R E N S , S P E R M A T IC CORD
T he epid id ym is (F ig . 2) begins w ith a s tro n g ly th ick en ed h ead (A — g;
B and C — e, f, g) ad h erin g to th e up p er pole of th e te sticle . T h e head
passed into th e b od y
(A — j) situ a te d
c a u d o la te ra lly
and th e n to th e
tail of th e ep id id ym is (A and B — k) situ a te d ca u d o v e n tra lly in re la tio n
to the lo w er pole of th e testicle .
T he head of th e epid id ym is is sim ilar in shape to a tria n g le w ith its
b ase d ire cte d to th e
b ase of
re a ch in g
th e
a lm o st
to
the
outlin e
sp e rm a tic
of
th e
cord (C— i), and th e a p e x
a n te rio -m e d ia l su rfa ce of th e
te stic le (B — e, C —- e). T h e h ead is fo rm e d of tw o ro o ts, th e p ro x im a l
an d d i s t a l J). T h e p ro x im a l ro o t (f) is co n n ected in th e w a y p re v io u sly
d escrib ed w ith th e te sticle . Th e b re a d th of th is ro o t in a d u lt in d ivid u als
is g re a te r th a n th e b re a d th of th e d istal ro o t (g). In th e m a te ria l e x a m in ­
ed this dim ension e x ce e d s 50% of th e b re a d th of th e b ase o f th e ep id i­
d ym is h ead (h). T h e d istal ro o t passed fro m th e a p e x of the epididym is
h ead a t an a cu te
40%
an gle to th e side
and b ack .
Its b re a d th fo rm s ab ou t
of th e b read th o f th e base. Th e b od y of th e ep id id ym is, slig h tly
‘) Term s introduced by O. R. S c h l u m p e r g e r
%
(1054).
The m ale reproductive organs of the European bison
517
w idened In th e in itial p a rt, re ta in s a lm o st id en tical d im en sion s th ro u g h ­
o u t its len g th (T ab le 3).
T he tail of th e ep id id ym is, also fo rm ed of tw o ro o ts clo se ly ad h erin g
to each o th e r, is a tta c h e d to th e te stic le c a u d o la te ra lly fro m its lo w e r
pole. Th e c la v a te a p e x o f th e ca u d a ep id id y m id is is d ire cte d b a ck w a rd s
and slig h tly d o w n w ard s. T h e p ro x im a l ro o t (w h ich is a co n tin u a tio n of
th e body) has a s m a lle r lum en th a n th e d istal ro o t (P h o to 3). L ig a m e n ­
tu m testis p ro p riu m is stro n g , an d sh o rt.
In th e in d ivid u als ex a m in e d th e re w ere no co n n e ctio n s b e tw e e n th e
fre e m arg in of th e ep ididym is and th e ep id id ym al su rf a c e of th e te stic le .
A s a re s u lt b u rs a te s tic u lo ru m is cap acio u s and is u n in te rru p te d ly co n ­
n ected w ith c a v u m scro ti.
T ab le 3.
M easurem ents of epididym is in mm.
B readth of head of
epididymis
No.
1
2
3
4
5
6
7
8
Name
Putn ar
Pug II
Pozew
Pustelnik
P ołam aniec
Pokorny
Pluszcz
Plato
B read th of corpus
Base
Proxim al
root
D istal
root
18
26
17
17
35
35
28
35
14
20
12
16
22
21
21
21
34
47
28
37
57
58
49
56
In itial
Midway
Term inal
9
15
9
15
17
18
12
11
8
9
9
12
16
16
10
9
10
3
10
17
17
16
8
10
D u ctu s d e f e r e n s ru n n in g fro m th e ta il of th e ep id id ym is and a tta c h e d
to m eso d eferen s lies f la t along th e ca u d o m ed ial s u rf a c e of th e te stic le .
P a r a lle l to th e d u c tu s d e f e r e n s and c ra n ia d fro m it a d is tin c tly th ick e n ­
ed b an d ca n be seen situ a te d alon g th e a n te rio r m a rg in of th e p lica d u c ­
tu s d e f e r e n t i s an d co v ered b y th is fold. It a cc o m p a n ie s th e d u c tu s d e ­
f e r e n s in th e reg io n of th e sp e rm a tic co rd and in g u in al ca n a l (cf. p. 518).
B eg in n in g fro m th e ep id id ym id al ta il, th e d u ctu s d e f e r e n s fo rm s a se rie s
o f tu rn s up to th e lev el of th e d istal
o f th e lon g a x is o f th e te stic le ,
th e n ru n s stra ig h t fo rw a rd an d e n te rs into th e co m p o sition of th e
s p e rm a tic co rd .
T h e s p e rm a tic co rd n e a r th e e x te r n a l ingu in al rin g is o v al in sh ap e in
th e c ro s s-se c tio n , and ro u n d a t th e lev el of co llu m s c ro ti (P h o to 2 -A ),
M . c r e m a s te r e x te rn u s , in th e reg io n of th e ingu in al c a n a l, lies on th e
la te r a l sid e of th e sp e rm a tic co rd . A f te r leavin g th e ca n a l th e m u scle
m e n tio n ed ab ov e is at firs t situ a te d ca u d a d , th e n a t th e level of th e
K , Sw iezynski
518
c o llu m sc ro ti c a u d o la te ra lly fro m th e sp e rm a tic co rd (P h o td 2— A — a).
In th is p a rt m . c r e m a s t e r e x t e r n u s su rro u n d s th e sp e rm a tic co rd o v e r
an a re a of V2 to V3 of its c irc u m fe re n ce . V e n tra d th is m u scle on th e
p o s te rio r side a tta in s th e le v e l of th e h ead of th e ep id id ym is. O n th e
la te r a l side it p ro d u ces s e v e ra l fla t fa scicle s en sh eath in g p ro ce ssu s v a ­
gin a lis below th e h ead of ep id id ym is also. A m on g th e E u ro p e a n bison
d issected , in tw o in d ivid u als th ese fa scicle s e x te n d to th e lev el of ca u d a
ep id id y m id is. In th e reg io n of th e sp e rm a tic co rd the d u c tu s d e f e r e n s ,
to g e th e r w ith th e acco m p a n y in g band (th e re sid u u m of th e co rre sp o n d ­
ing e m b ry o n ic M illerian d u ct), jo in tly a tta c h e d to a re la tiv e ly long
m eso d eferen s are lo c a te d behind th e v a s c u la r p a rt of th e sp e rm a tic co rd .
P h o to 2— A illu s tra te s th e re c ip ro ca l re la tio n of th ese e le m e n ts in th e
le ft an d rig h t sp e rm a tic co rd a t th e level of th e ep id id ym id al h ead .
A
B
C
Fig. 2. Plan of situation of epididymis in relation to testicle (le ft testicle),
a, b, c, d — su rfaces: cran ial, m edial, caudal and lateral of the te sticle ; e — apex;
f — proxim al root; g — d istal root of head of epididym is; h — bread th of base
of epididym al head; i — position of vascu lar part of the sperm atic cord; j — corpus;
k — tail of epididymis.
B etw e e n th e le ft an d rig h t sp e rm a tic co rd s, ab ove th e pen is, a m ass
of f a t and co n n ectiv e tissu e a cc u m u la te s in th e E u ro p e a n bison. In this
reg io n th e re a re also th e ingu in al su p e rficia l lym p h onod es, tw o of w hich
o ccu rre d on each side. C ra n ia lly th e y a r e situ a te d in fro n t of th e sp e r­
m a tic co rd s ab ov e th e penis. C a u d a lly behind th e sp e rm a tic co rd s, below
the sigm oid fle x u re of th e penis.
D u ctu s d e f e r e n s , a f t e r leav in g th e in te rn a l ingu in al rin g ru n s, a tta ch e d
b y th e d efe re n tia l fold , alon g th e in te rn a l su rfa ce of th e ab d om in al w all.
The m ale reproductive organs of the European bison
51 9
A fte r a lo n g er o r sh o rte r d istan ce (depending on th e d e g re e of d evelo p ­
m en t o f u t e r u s m a s c u lin u s ) th e d iffe re n tia l fold join s w ith th e fold of
the o th er side to fo rm p lica u ro g e n ita lis. T h e »band« a cco m p a n y in g th e
d u c tu s d e f e r e n s , a fte r re a c h in g th e a p e x of th e u te rin e h o rn of th e
u te ru s m a s c u lin u s , p e n e tra te s into its w all (cf. F ig . 3 an d P h o to 5). T h e
d u ctu s d e f e r e n s , on th e o th e r h an d , p asses in th e d ire ctio n of th e la te ra l
m arg in of th e h orn an d is situ a te d v e n tra lly along th is m a rg in . In som e
E u ro p ean bison th e w all of th e u te rin e h orn su rro u n d e d , th e d u c tu s d e ­
fe r e n s in su ch a w a y th a t it w as n e ce s sa ry to cu t open th e u te rin e w all
in o rd er to tra c e its co u rse. A t th e lev el of th e undivided p o rtio n o f th e
u te ru s m a s cu lin u s th e d u ctu s d e f e r e n t e s leav e th e l a tte r an d ru n in th e
u ro g en ital fold to w a rd s th e u re th ra . T h e end p a rts of d u ctu s d e f e r e n t e s
are m ark e d ly th ick en ed and e x h ib it s tru c tu r e ty p ica l of a m p u lla d u c t u s
d e fe re n tis .
In th e g ro u p of in d ivid u als up to ten m o n th s old d iffe re n ce s in th e
len g th of d u ctu s d e f e r e n t e s a re v e r y sligh t.
In E u ro p ean bison o v e r tw o y e a rs old th e len g th of d u c tu s d e f e r e n t e s
(F ig . 1 E : 1— 3) v a rie d w ith the ind ivid u al irre sp e c tiv e o f a g e an d size
of th e an im al (cf. T ab les 1 and 4).
D u ctu s d e f e r e n t e s end jo in tly w ith th e e x c r e to r y d u cts of th e v e s ic u la r
glan d s on th e co llicu lu s sem in a lis in openings sim ila r in a p p e a ra n ce to
slits (P h o to 14).
V.
TH E
SCRO TU M AND TH E COVERIN G
ST R U C T U R E S OF TH E T E S T IC L E S ,
O F TH E E P ID ID Y M IS AND O F TH E SP E R M A T IC
CORDS
T h e s c ro tu m in th e E u ro p e a n bison is situ a te d in th e in g u in al
on th e level of th e p ro x im a l cu rv e of th e sigm oid fle x u re of th e
In th e fixed sp ecim en s it is la te ra lly fla tte n e d w ith sh a llo w
m ark ed lon g itu d in al g ro o v es (P h o to 1). C o llu m sc ro ti is o n ly
re g io n
penis.
fa in tly
fa in tly
d iffe re n tia te d an d its cro s s-se c tio n below th e penis is a p p ro x im a te ly
o v al, w ith its long a x is situ a te d obliqu ely to th e fro n ta l p lan e.
T h e le ft sp e rm a tic co rd is situ a te d cra n ia d , and th e rig h t ca u d a d fro m
th is p lan e. T h e te stic le s a re sim ila rly situ a te d in th e s c ro ta l sa c , th e le ft
a d v a n c e d in fro n t of th e rig h t b y a p p ro x im a te ly h a lf th e tra n s v e rs e
d im en sion . In th e fix e d ca d a v e rs this fo rw a rd s sh ift is ev en m o re m a rk e d
(P h o to 2, B an d C ).
A t a d ista n ce of 10 to 15 cm to th e fro n t th e co llu m s c r o ti an d to th e
sid e fro m a m ed ian lin e th e re a re ru d im e n ta ry te a ts -p a p iia e u b e ris, tw o
a sid e (P h o to 1-c).
T h e skin o f th e s c ro tu m is co v e rd w ith sh o rte r and sc a n tie r h a ir th a n
th e h a ir of th e a d ja c e n t reg io n s. T h ick n ess of th e skin on th e lev el of
K. Św ieżyński
520
c o llu m scro ti is fro m 11 to 16 m m in ad u lt E u ro p e a n bison. H a lfw a y
alon g th e le n g th o f th e s c ro tu m th is m e a su re m e n t v a rie s fro m 4 to 6
m m , and at th e a p e x of th e s c ro tu m fro m 1 do 2.5 m m .
T u n ic a d arto s is th ick e st in th e reg io n of th e coliu m sc ro ti, and in th e
a re a of s e p tu m sc ro ti, w h e re th e th ick n ess of th e d a rto s is a s m u ch as
4— 5 m m . S t r a t u m s u b d a rto ic u m is fa irly dense in te x tu r e . T h e p resen ce
in th is la y e r of loose co n n e ctiv e tissu e is m o st d istin ct on th e a p e x ol
th e sc ro tu m . F a s cia s u p e rfic ia lis scro ti a d h e re s clo sely to th is stra tu m
(p a rtic u la rly on th e side o f s e p t u m s c ro ti), and ca n easily be sep arated
T ab le 4.
Length of d u ctu s d eferentes in m illim etres according to age
groups.
No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Name
Total length of
ductus d eferen tes
Including length of
ampullae
R ight
Left
Right
Left
Pos
Pub
Pom
P la k a t
202
272
265
275
225
280
230
292
23
20
20
35
26
21
25
33
M ean
254
257
25
25
Poronin
P u tn a r
840
610
743
600
66
£9
62
115
M ean
725
672
77
89
Pug II
Pozew
Puer
Połam aniec
Pokorny
703
710
—
—
570
740
795
720
690
630
148
121
11?
115
115
120
100
115
112
119
M ean
708
688
125
116
Posusz
Pluszcz
P lam iec
Pom ruk
505
712
792
631
712
745
709
736
87
90
92
86
96
82
82
101
Mean
712
726
89
SO
fro m fa scia p ro fu n d a scro ti. P ro fu se , m a in ly fa t, co n n e ctiv e tissue,
th ru s ts b etw een th e tw o fa s cia l la y e rs a t th e lev el of co llu m scro ti and
fo rm s a co n tin u atio n o f th e a ccu m u la tio n p re v io u sly d escrib ed if
co n n ectiv e tissu e se p a ra tin g th e sp e rm a tic co rd s.
F a scia c r e m a s t e r ic a can e a sily be se p a ra te d fro m fa scia tra n sv ersa .
Th e la tte r , on th e o th e r h an d , se p a ra te s fro m th e sero u s m e m b ra n e only
The m ale reproductive organs of the European bison
521
a t th e lev el of th e sp e rm a tic co rd . In th e re m a in d e r of th e a re a fa scia
tra n sv ersa ad h eres to th e su rfa ce of p ro cessu s va gin a lis, fo rm in g la m in a
fib ro s a of th e p a rie ta l la y e r
of
v a g in a l p ro cess — p e r io r c h iu m s.
th e
tu nica v agin a lis co m m u n is. Th e v is ce ra l la y e r of th e v a g in a l p ro ce ss —
e p io rc h iu m s. tu n ica v a gin a lis p ro p ria b e h av es ty p ic a lly in th e E u ro p e a n
bison, th a t is, it clo sely co v e rs tu n ica a lb u g in e a of th e te stic le a n d p a rti­
cip a tes in th e fo rm a tio n of m esoep idid ym is.
V I.
TH E G E N ITA L A C C E SSO R Y GLANDS
1. The a m p u lla e d u ctu u m d e fe r e n tiu m
T h e s tr u c tu r e of a m p u lla d u c tu s d e fe r e n t is
in
th e
E u ro p e a n
bison
ex a m in e d su g g e sts th a t th e th ick en in g of th is p a rt of th e d u ctu s d e f e r e n s
is due both to in cre a se in th e th ick n ess of th e w all and to in c re a s e in
th e d ia m e te r. T h e d ia m e te r of th e
d u ct is fo rm ed
-sh a p e d
(1 9 3 0 )
b ran ch in g s.
K r ol l i n g
found
b y se v e ra l
n u m ero u s
n ich e­
g la n d u la r
e lem en ts in th e m u co u s m em b ra n e o f th e a m p u lla e d u c t u u m d e f e r e n ­
tiu m in th e E u ro p e a n bison, w h ich p oin ts to th e g la n d u la r c h a r a c te r
of th e am p u lla.
In th e m a te ria l ex a m in e d fo u r g ro u p s, co rresp on d in g to th e ag e grou p s,
c a n be d istin guish ed in re s p e c t of le n g th of am p u lla d u ctu s d e fe r e n t is
(cf. T ab le 4). W h ile th e co n sid erab le d iffe re n ce in this d im ension b etw een
c a lv e s up to one m o n th old and o ld er in d ivid u als is of co u rse obviou s,
th e d ifferen ces am o n g
th e
re m a in in g
d im en sion d e cre a se s in a n im als o v e r
10
a n im a ls
y e a rs
a re
re m a rk a b le . This
old. A s
th e
le n g th
of
d u c t u s d e f e r e n t e s v a rie s co n sid e ra b ly in all th e g rou p s, irre s p e c tiv e of
th e b od y m e a su re m e n ts, calcu la tio n w as m ad e of the le n g th o f a m p u lla e
d u c t u s d e f e r e n t e s in th e fo rm of p e r ce n t of len g th of th e co rresp o n d in g
d u c t u u m d e f e r e n t i u m v a rie s co n sid e ra b ly in all th e g ro u p s, irre sp e c tiv e
of th e b od y m e a su re m e n ts, ca lc u la tio n w as m ad e of th e le n g th of a m p u l­
lae d u c t u u m d e fe r e n t iu m in th e fo rm of p e r ce n t of len g th of th e c o rr e s ­
p on d in g d u c tu s d e f e r e n s (T ab le 5). T h ese d a ta sh o w th a t th e p a rtic ip a tio n
of a m p u lla d u c tu s d e f e r e n t i s in th e le n g th of th e d u c tu s d e f e r e n s in c r e a ­
ses w ith ag e, atta in in g a m a x im u m a t 6— 10 y e a rs of age. In o ld er bison
th e v a lu e s of th is ra tio ag ain d e cre a se , a p p ro ach in g th e co rresp o n d in g
v a lu e s in an im als fro m 2— 3 y e a rs old. T h is m a y be co n n e cte d w ith th e
p ro c e s s of sen escen ce, a lth o u g h d a ta o b tain ed fro m p ed ig ree books do
n ot in d ica te th a t
th e
re p ro d u c tiv e
c a p a c ity
of
m ale
bison
m a rk e d ly
d e c re a s e d a t ag es o v e r te n y e a rs (» P la m ie c 7 89«, »P lish 229«, » P lu v iu s II
5 4 6 « , »P od b ip ig ta 738«),
K. Św ieżyński
522
2, V e s ic u la e se m in a les
T w o v e sicu la r glan d s o ccu r in th e E u ro p e a n bison (P h o to 5, 6, 7).
T h e y a re fo rm ed b y elo n g ated g la n d u la r p ack ets, su rrou n d ed b y a lay er
fro m 1.5— 2.5 m m th ick of co n n e ctiv e tissu e w ith ab u n d an t v essels, n erve
fib res an d sm o o th m u scle fib res. S itu a te d la te ra lly fro m a m p u lla e d uctu u m d e f e r e n t i u m th e se g lan d s a re hidden, to g e th e r w ith th e am pullae
T ab le 5.
Ratio of length of a m p u lla ductus d e je r e n tis
to length of d u ctu s d e fe r e n s in percentages
(according to age groups).
A m p u lla ductus d eferen tis
No.
1
2
3
4
5
6
7
8
9
10
Name
L eft %
Pos
Pub
Pom
P la k a t
11.40
7.35
7.55
12.73
11.60
7.50
9.13
11.30
M ean
9 75
8.34
Poronin
P utnar
7.85
14.59
8.34
19.16
Mean
11.22
13.75
Pug II
P uer
P ołam aniec
Pokorny
21.05
19.64
15.54
14.47
16.90
15.97
16.23
18.88
17.67
16.99
Mean
11
12
13
14
R ight %
Posusz
Pluszcz
P lam iec
Pom ruk
12.64
11.71
13.63
13.48
11.00
11.56
13.72
Mean
12.66
12.44
—
and u t e r u s m a s c u lin u s , in th e u ro g e n ita l fold. T h e y a re co v ered o v e r the
cra n ia l 3/* of th e ir len g th b y th is fold. T h e e x te rn a l su rfa ce of th e g laid s
is folded, w h ich is co n n e cte d w ith th e ir lo b u la r s tru c tu r e and gives the
w h o le a ra ce m o se a p p e a ra n ce (P h o to 6). O n th e p o ste rio r end of each of
th e g lan d s th e r e is a lobe, situ a te d m ed ially , w h ich is co n n ected v ith
a sim ila r lobe of th e glan d on th e opposite side b y co n n e ctiv e tissue.
T h e re is a so rt of tra n s v e rs e isth m u s th ru stin g b etw een th e am pullae
d u c t u u m d e f e r e n t i u m an d th e u re th r a (P h o to 7b). T h is com m issure, o f
The m ale reproductive organs of the European bison
523
a co n n ectiv e tissu e c h a r a c te r , is u n ited w ith the end p a rts of the d u c tu s
d e fe r e n t e s and u r e th r a b y th e su rro u n d in g co n n e ctiv e tissu e . T h e m ain
d e fe re n t d u ct ru n s fro m ea ch of th ese g lan d s an d ends on th e v e n tra l
w all of th e d u c tu s d e fe r e n s .
T h e m e a s u re m e n ts m ad e on th e m a te ria l a t o u r d isp osal show th a t
th e d im ension s of th e v e sicu la r g lan d s in cre a se m a rk e d ly up to th e age
of ab o u t 4 y e a rs (T ab le 6). T h e p red o m in an ce of d im ension s of one glan d
T ab le 6.
Measurements of v esicu lae seminales according to age groups.
Right vesicle
No.
1
2
3
4
5
6
7
8
9
10
U
12
13
14
15
16
17
Name
Longitudinal
m easurem ent
Pos
Pub
Pom
P la k a t
23
22
30
35
Mean
L eft vesicle
T ransverse
m easurem ent
7
Longitudinal
m easurem ent
Transverse
m easurem ent
5
6
26
31
28
33
28
6
39
8
Poronin
P utnar
Puck
54
86
83
17
32
28
52
90
63
15
17
21
Mean
74
26
68
18
100
120
83
90
112
112
95
91
92
111
28
30
19
35
41
44
51
31
39
36
95
102
78
110
110
108
101
85
98
119
30
35
25
38
35
35
37
38
18
41
101
35
101
33
Puzor
Pug II
Pozew
Puer
Połam aniec
Pokorny
Posusz
Pluszcz
P lam iec
Pom ruk
Mean
—
9
—
7
8
o v e r th e o th e r v a rie s w ith th e ind ivid u al. In E u ro p e a n b ison o v e r five
y e a r s old th e re is no re la tio n b e tw e e n th e size o f th e g lan d s and th e
b od y d im ension s (cf. »P u g II« , »P o z e w « , » P la m ie c« ).
3. T h e p ro state
T h e p ro sta te — co n sists of co rp u s p ro sta ta e and p a rs d issem in a ta .
T h e b od y of th e p ro s ta te is situ a te d on th e d o rsa l s u rfa c e of th e u re th ra
in th e im m e d ia te v ic in ity o f th e o stiu m s of d u ctu s d e f e r e n t e s and the
e x c r e t o r y d u cts of th e v e sicu la r g lan d s (P h o to 6).
K . Sw ieżyński
524
In th e m a te ria ł e x a m in e d th e body of th e p ro sta te (P h o to 6, d) w as
fo rm ed of tw o lob es in ca lv e s up to one m o n th old. In o n e -d a y old
»P os« an d »P u b «, an d in se v e n -d a y old » P o m «, th e lobes of th e b od y
of th e p ro sta te h a v e rem ain e d co m p le te ly in d ep en d en t. In sh ap e and size
th ey a re sim ilar to f l a x seed s a rra n g e d obliqu ely in re la tio n to e a ch
o th e r. In o n e-m o n th old » P la k a t« th e tw o lobes a re clo se r to e a ch o th e r
but ca n be se p a ra te d w ith o u t d ifficu lty . In bison o v e r tw o y e a r s old th e
body of th e p ro s ta te tak e s th e fo rm of a single cy lin d rica l fo rm a tio n .
T h e m o re or less d istin ct c la v a te ends a re th e » v estig e« of th e b ilo b u lar
s tr u c tu r e of th e g lan d . Th e tw o lobes ad h e re clo se ly to e a ch o th e r and
in ad u lt ind ivid u als th e b o rd e r b etw een th e m ca n n o t be d e te rm in e d
m acro sco p ica lly . T h e tra n s v e rs e dim ension of th e b o d y of th e p ro sta te
(F ig . 1, C , 5— 6) in a d u lt E u ro p e a n bison v a rie s fro m 23— 54 m m (m ean
33 m m ). T h e lon g itu d in al m e a su re m e n t of th e rig h t lobe is fro m 5—
8 m m (m ean 6 m m ) and th e le ft lobe fro m 5 to 9 m m (m ean 7 m m ,
n—
D ev elo p m en t of th e lobes in d iffe re n t in d ivid u als does n ot e x h ib it
a n y im p o rta n t d ifferen ces. It w as o n ly in th e bison »P o ła m a n ie c« th a t
th e rig h t lobe w as f a r less d eveloped th an th e left.
P a rs d issem in a ta p ro sta ta e begins a t a d ista n ce of ab ou t 0.5 cm ca u d a d
fro m co llicu lu s sem in a lis.
A t firs t th e a m o u n t o f g la n d u la r tissu e is sm all and is situ a te d on th e
d o rsal and d o rs o -la te ra l side of th e u re th ra (P h o to 16— 3). T h e fu rth e r
cau d ad , th e m o re th e m e a s u re m e n ts of th e g la n d u la r lobes in cre a se , so
th a t h a lfw a y alon g p a rs pe lv in a u r e t h r a e m a s cu lin a e, p a rs d issem in a ta
p ro sta ta e su rro u n d s th e u re th r a co m p le te ly fro m th e top an d sides,
leavin g only a n a rro w b an d fre e on the v e n tra l side (P h o to 16— 5, 6).
In this p a rt th e g la n d u la r la y e r on th e d orsal side ca n be as m u ch as
15 m m th ick , an d on th e la te r a l sides fro m 8 to 10 m m . In th e a re a of
th e p o ste rio r h a lf of th e p elv ic p o rtion of th e u re th r a th e g la n d u la r
p a re n ch y m a also th ru s ts u n d er th e u re th ra on th e v e n tra l side (P h o to
16— 8, 9, 10), g ra d u a lly b ecom in g th in n e r. T h e »cu ff« fo rm ed in th is w ay
w ith th e p a rs d is se m in a ta p ro sta ta e ends in a co n ical sh ap e a t a d ista n ce
of ab o u t 2 cm fro m th e tra n sitio n of p a rs p elv in a u r e t h r a e into pars
b u lb i u r e t h r a e .
The openings o f th e d efe re n t
d u cts of the
p ro sta te
a re
a rra n g e d in
fo u r ro w s on th e d o rsal w all of th e u r e th r a (P h o to 14).
4. The b ulb ou reth ral glands
T h e b u lb o u re th ra l g lan d s — g la n d u la e b u lb o u re t h r a le s — a re form ed
of th e lobes co n n ected b y p ro fu se co n n e ctiv e tissu e. A s a re s u lt a flat
h e a rt-sh a p e d fo rm a tio n is cre a te d , th e double s tr u c tu r e of w h ich is m a ­
The m ale reproductive organs of the European bison
525
n ifested b y th e c o n v e x a n te rio r u p p er m a rg in and th e sh allow g ro o v e
on th e p o sterio r su rfa ce . T h e d e ffe re n t d u cts of th e g lan d s join in one
com m on d u ct fro m 1.2 to 2 cm lon g w h ich ru n s along th e ca u d o -d o rsa l
su rfa ce of the u re th ra . The d u ct opens into a re ce ss situ a te d on th e
ca u d o -d o rsal w all of p a rs b u lb i u r e t h r a e . T h e re ce ss is bounded b y a fold
of m u co u s m em b ran e in th e fo rm of an in v e rte d le tte r V.
T h is o rg an , w ith
th e
s tru c tu re
b o u n d ary lin e b etw een th e p e lv ic
d escrib ed
and
ab ov e, is situ a te d on th e
b u lb ar
p o rtion of th e u re th ra
(P h o to 15).
In ad u lt bison th e lo n g itu d in al d im ension of th e g lan d {F ig . 1, D , 1— 3)
w as fro m 17 to 28 m m (m ean 21 m m ). T h e tra n sv e rse
(Fig. 1, D, 1— 2) v a rie s fro m 15— 33 m m (m ean 27 m m ).
m e a su re m e n t
S e v e ra l m u scles su rro u n d th e glan d s fro m all sides. T h e ca u d o v e n tra l
p a rt is co v ered b y th e p a rs b u lb i and to g e th e r w ith it, b y m. b u lb o c a v ern o su s.
of
T h e cau d o d o rsal s u rfa c e co v e rs a n in d iv id u ally v a ry in g (in re sp e ct
o c c u rre n c e and d irectio n in w h ich th e fib res ru n ) m u scle , th e
s tr u c tu r e and in sertio n s of w hich co rresp o n d to m . isch io b u lb o su s
d escrib ed in th e ra m b y B r a u e l l (1 8 6 8 ). M u scles su ch a s m . ischiou re th ra lis , m . u re th ro g la n d u la ris and p a rtly m . bulbogrlaiidularis th r u s t
b e tw een th e a n te rio v e n tra l su rfa c e of th e gland d escrib ed and th e
u re th r a . O f th ese m u scles, o n ly th e first o ccu rre d in all th e d issected
a n im als an d w a s ch a ra c te riz e d b y a co n tin u ou s co u rse (P h o to 6). Th e
p elv ic in sertio n of th is m u scle is situ a te d on th e in te rn a l su rfa ce of th e
t u b e r isch ii and is su rro u n d e d b y th e stro n g m . isch io c a v ern o s u s . Th e
th in sle n d e r b elly o f th e m u scle ru n s m ed ially to w a rd s th e u re th r a and
p asses in to an ap o n eu ro sis, w h ich th ru s ts u n d er gl. b u lb o u re th ra lis ,
su rro u n d s th e u r e th r a and u n ites w ith th e sam e m u scles of th e opposite
sid e. B e tw e e n th e co n n e ctiv e tissu e su rro u n d in g th e glan d an d th e a p o ­
n e u ro sis th e re is a m a c ro s co p ic a lly visib le e x ch a n g e of fib res. The
a d u re th ra l su rfa ce of th e b u lb o u re th ra l g lan d , w h e re n ot se p a ra te d b y
m u sc le s, ad h eres clo se ly to th e p e lv ic p a rt of th e u re th r a . In this p lace
m . u r e t h r a lis is th in and th e b u lb o u re th ra l glan d is se p a ra te d fro m th e
a p e x o f p a rs d issem in a ta p ro sta ta e by th e ap on eu ro sis of th e u re th ra l
m u sc le o nly.
V II. THE U T E R U S M A SC U LIN U S
A m o n g th e se v e n te e n m ale E u ro p e a n bison w h ich w e e x a m in e d in
th is re s p e c t th e u t e r u s m ascu lin u s o ccu rre d in six te e n cases. T h is organ
e x h ib ite d a v a ry in g d eg ree of d ev elo p m en t in d iffe re n t in d ivid u als and
s e v e r a l ty p e s w e re d istig u ished on this b asis (F ig . 3).
K . Sw iezynski
526
In th e case of n in e bison
th e
e x te rn a l
a p p e a ra n ce
is
d e ce p tiv e ly
sim ila r in ap p e a ra n ce to th e u te ru s in th e fem ale. P a ire d h o rn s an d
u n divid ed p a rt cou ld be d istin g u ish ed in it.
T h e h orn s, like th e
u te rin e
h orn s
of
th e
co w , w e re d ire cte d w ith
g r e a te r c u rv a tu r e u p w a rd s an d fo rw a rd s, and le sse r c u rv a tu r e d o w n ­
w a rd s and b ack w ard s. The tu r n of th e h orn s w as n o t h o w ev er co m p le te ,
an d th e apices of th e h o rn s e x te n d e d into »bands« a cco m p a n y in g th e
d u ctu s d e je r e n t e s . T h e u n d ivid ed p a r t, a t first w id e, b eco m es n a rro w e r,
th en w id en s o u t ag ain in th e te rm in a l p a rt (P h o to 5, 6, 7). T h e w hole
o rg a n w as d o rs o -v e n tra lly fla tte n e d , th is being m ost m ark ed in th e
Fig. 3. P lan of types of u teru s m a scu lin u s in the European bison. Numbers under
figures represen t the num ber of cases found.
te rm in a l p a rt, w h e re th ick n ess n e v e r ex ce e d e d 10 m m . P lic a u ro g e n italis co v e rs u t e r u s m a s cu lin u s to g e th e r w ith the fin al p a rts of d u c tu s
d e j e r e n t e s and th e v e sicu la r glan d s, b eh avin g in th e re g io n of th e p elv ic
c a v ity in a sim ilar w a y to th a t found in fem ales. In a d u lt E u ro p e a n
bison w ith th is ty p e of u te ru s th e dim ensions (F ig . 1, C) of its d iffe re n t
p a rts w ere as follow s:
(a) le n g th of rig h t h o rn : (1— 2) 115 to 260 m m , m ean 185 m m
(b) len g th of le ft h o rn : (2— 3) 100 to 270 m m , m ean 160 m m
(c) len g th of u n d ivid ed p a r t: (2— 4) 185 to 208 m m , m ean 198 m m
In fo u r bison u t e r u s m a s cu lin u s o ccu rre d in the fo rm o f an u n d ivid ­
ed co n ical d u ct p assin g to th e fro n t in a single »b an d «. On th e b o rd e r
of p lica u ro g e n ita lis th e »band« divided into tw o, w h ich b eh aved sim il­
a r ly in th e ir co n tin u ed co u rse to th a t in th e first ty p e . A m o n g th e th re e
bison possessing th is ty p e o f u te ru s th e re w e re in d ivid u als in w hich
The m ale reproductive organs of the European bison
g ro w th w as n ot co m p le te and it is th u s
dim ensions of th is o rg an in this grou p .
d ifficu lt
to
527
sp eak o f m e a n
A sim ila rly undivided u te ru s m a s cu lin u s w a s found in th e E u ro p e a n
bison »P o ła m a n ie c« . It w a s a m p u lla ry in sh ap e, in th e fo rm of a n o val
v esicle m easu rin g 15 m m X 20 m m . W e th e re fo re d iffe re n tia te d th is
u teru s into a se p a ra te ty p e (D) on th ese g rou n d s.
A lth o u g h
in
tw o
fu rth e r
E u ro p e a n
bison
u t e r u s m a scu lin u s
w as
sim ilar in s tr u c tu r e to th a t in th e first ty p e , it d iffe re d co n sid e ra b ly
in dim ensions (cf. in T ab le 7 — » P u ck « , »P u g II« ). T h e u te rin e h o rn s
of th ese u te ri e x te n d e d to th e in te rn a l ingu in al rin g s, an d in »P u g II«
even p e n e tra te d into th e a re a of th e ingu in al can al.
T able 7.
E xtern a l m easurem ents of u teru s m ascu lin u s in m illim etres.
No.
Name
T ype of
uterus
acc. to
Fig. 3
1
2
3
4
D
6
7
8
9
10
11
12
13
14
15
16
17
Pos
Pub
Pom
P la k a t
Poronin
P utnar
Puck
Puzor
Pug II
Pozew
Puer
Połam aniec
Pokorny
Posusz
Pluszcz
P lam iec
Pom ruk
A
B
B
A
B
A
C
A
C
(E)
A
D
A
B
A
A
A
Length of horns Length of
undi­
vided
R ight
Left
part
39
40
—
—
—
—
30
35
—
—
120
290
180
460
135
320
120
510
68
50
30
80
40
180
135
200
275
—
—
—
115
160
200
19
202
20
185
208
193
—
—
240
270
—
—
260
151
162
100
132
180
U te ru s m a s c u lin u s in all th e d issected E u ro p e a n bison, irre sp e c tiv e
of ty p e , had a lu m en , w h ich o n ly in » P la k a t« w as in te rru p te d in th e
un d ivid ed p a rt by ad h esion of th e w alls o v e r a len g th of a b o u t 5 m m .
T h e lu m en of th e u te ru s w a s co n n ected w ith th e lu m en of th e u ro g e n i­
ta l d u ct b y m ean s of th e opening lo ca te d on co llic u lu m sem in a lis
(P h o to 14).
T h e w a lls of u te ru s m ascu iitiu s of th e first and th ird ty p e s ex h ib ite d
d iffe re n t p a rts in re s p e c t of s tru c tu r e and co lou r of th e m u co u s m e m ­
b ran e.
T h e h o rn s w ere co v e re d b y d a rk co lo u red m u co u s m e m b ra n e n ot e x ­
h ib itin g fold in g. In so m e in d ivid u als sm all p ro m in en ces w e re ev id en t
52 8
K, SwiezyAski
in th e sh ap e of lo n g itu d in al folds, and in th re e th e y ap p eared as b e a d ­
-lik e th ick en in gs. T h e th ick n e ss of th e w alls of th e h o rn s, lik e th e
th ick n ess of th e w alls of th e o th e r p a rts of u t e r u s m a s c u lin u s , d iffe re d
in d iffe re n t in d ivid u als, v a ry in g fro m 1 to 1.37 m m . Th e »ban d s« ru n n ­
ing fro m th e h o rn s (m en tion ed p re v io u sly in d iscussing th e co u rse of
th e d u c tu s d e f e r e n t e s ) had a lu m en in th e a d u te rin e p a rts an d w e re
co v e re d w ith sim p le co lu m n a r e p ith eliu m . T h e e x te n t of th e lu m en in
a d ire ctio n to w a rd s th e te stic le s cou ld be d e te rm in e d m a c ro s co p ic a lly
o v e r sectio n s of d iffe re n t len g th .
T h e h orn s p assed in to th e re g io n
of th e undivided p a rt of u t e r u s
m a s c u lin u s , w h ich is w id est and situ a te d fu rth e s t cra n ia d (P h o to 6 and
7). It is co v ered b y m u co u s m e m b ra n e d evoid of m a c ro s co p ic a lly v isib le
folds and sim ila r in co lo u r to th e m u co u s m e m b ra n e of th e h orn s.
A m on g th e eleven E u ro p e a n bison (ty p es A + C) in sev en a le n g th -w a y s sep tu m , d ividin g th e lu m en into tw o h a lv e s, o ccu rre d on th e
w all of th e la st-m e n tio n e d sectio n . This sep tu m is th e sta rtin g p oin t of
th e division into h o rn s. T h e b o rd e r of th e sep tu m w as fo rm e d b y
m u scle tissu e co v e re d b y m u co u s m em b ran e (4 cases), o r w a s o n ly th e
doubling of th e m u co u s m e m b ra n e (3 cases). T h e th ick n ess of th e w alls
of th is p a rt of th e o rg a n v a rie d fro m 1.2 m m to 2.9 m m .
The n e x t se c tio n of th e le n g th fro m 20 to 60 m m w a s co v e re d b y
lig h t-co lo u re d m u co u s m em b ra n e fo rm in g c irc u la r folds v isib le u n d er
a m ag n ify in g g lass w ith 3 X m a g n ifica tio n . T h e w alls in th is se ctio n
th ick en d istin ctly , re a ch in g as m u ch as 2.30 to 4 .3 6 m m .
T h e n e x t sectio n , s tro n g ly n a rro w e d in th e m a jo rity of th e E u ro p e a n
bison (P h o to 7 d ) and fro m 80 to 120 m m long, w as co v e re d b y m u co u s
m em b ran e slig h tly d a rk e r in co lo u r th a n th e p reced in g , w ith a slig h t
y ellow ish o r b ro w n sh ad e. It w as co v e re d b y lon g itu d in al low fold s p e r­
p en d icu lar to th e folds p rev io u sly d escrib ed . T h e th ick n ess of th e w alls
in th e section v a rie d fro m 0.5 to 2 .3 0 m m .
The m u co u s m e m b ra n e in th e fin al re -e n la rg e d sectio n w as d ark in
co lou r. It w as co v e re d w ith d en sely a rra n g e d fold s fo rm in g a s o r t of
e x te n sio n to th e back o f th e folds of th e n eig h b ou rin g se ctio n . T h e
len g th of this a re a w as 5 to 17 m m .
H isto lo g ical slides m ad e fro m th e reg io n s e x h ib itin g m a c ro s co p ic d if­
fe ren ces u su ally co n firm th e e x iste n ce of th e se d ifferen ces.
The w all of th e h orn s (P h o to 8, 9) is lined w ith m u co u s
co v ered b y sim p le co lu m n a r ep ith eliu m . T h e tru e la y e r
m em b ran e co n tain s n u m ero u s b ra n ch e d tu b u la r glan d s an d
sels. C o lu m n ar ep ith eliu m lin es th e glan d s, w h ich re a c h to
m e m b ra n e
of m u co u s
blood v e s ­
th e m u scle
la y e r. The v a s c u la r la y e r is c le a rly d efined. S im ila r re la tio n s a re e x ­
h ib ited b y th e firs t tw o sectio n s of th e undivided p a rt, th e th ick n ess of
The m ale reproductive organs of the European bison
529
th e m u scle la y e r in creasin g cau d ad , and th e n u m b e r of g lan d s d e c re a s ­
ing.
The slides fro m th e n a rro w e d ca u d a l sectio n of th e u n d ivid ed p a rt
e xh ib it a d iffe re n t s tr u c tu r e (P h o to 10, 11). A lth o u g h th e m u co u s m e m ­
b ran e is also co v ered b y sim ple co lu m n a r e p ith eliu m it is lo w er th a n
in the p rev io u s sectio n s. T h e tru e la y e r o f m u co u s m e m b ra n e does n ot
co n tain g lan d s and e x h ib ite a la rg e n u m b e r of co n n e ctiv e tissu e fib res.
T he sh o rt b ro ad en ed fin al sectio n , w ith stro n g ly folded m u co u s m e m ­
b ran e, h as sim ple co lu m n a r ep ith eliu m (P h o to 12, 13). T h e tru e la y e r
fo rm s high tu b e rcle s u n d er w h ich th e re is a dense n e tw o rk of v en o u s
vessels. S c a tte re d glan d s ap p e a r again. S tra t u m m u s c u la r e is s tro n g ly
developed.
V III. TH E P E L V IC PO RTIO N OF TH E U R O G EN ITA L DUCT
C ollicu lu s sem in alis is situ a te d on th e d orsal w all of th e in ita l se ctio n
of th e p elv ic p o rtio n o f th e u ro g e n ita l can al (P h o to 14). A cco rd in g to
P i l a r s k i (1967) in th e E u ro p e a n bison co llicu lu s sem in alis is situ a te d
on th e b o u n d ary b etw een p a rs u ra lis and p a rs p ro sta tica of th e u re th r a .
It fo rm s a oval p ro m in en ce , th e len g th of w hich a tta in s a s m u ch as
2 cm in ad u lt bison. T h e p o ste rio r h ig h est p a rt of th e co llicu lu s w h ich
is a th ick en in g p assin g into a le n g th -w a y s fold ru n n in g alon g th e d o rsa l
w all o f th e u ro g e n ita l ca n a l (P h o to 14, d). T h e su rfa ce o f th e co lli­
cu lu s se m in a lis is d ep ressed like a c r a te r . T h e d ep ression is su rro u n d e d
by a rid g e -lik e p ro m in en ce fro m w h ich th e o pen in gs of th e e ja c u la to r y
d u cts a re situ a te d m ed ially. On th e fla tte n e d p o ste rio r su rfa ce of th e
co llicu lu s in th o se in d ivid u als in w h ich th e lum en of u t e r u s m a s c u lin u s
join s th e lum en of he u ro g e n ita l ca n a l th e re is an opening into u t e r u s
m a s c u lin u s (P h o to 14).
T h e p elv ic p o rtio n o f th e u ro g e n ita l ca n a l is su rro u n d e d b y co rp u s
c a v e rn o s u m u r e t h r a e , b egin n in g a t the h e ig h t of co llicu lu s sem in a lis.
A t firs t it is situ a te d v e n tra lly in re la tio n to th e u re th r a in th e fo rm
o f a n a rro w band of e re c tile tissu e (P h o to 16 — 1, 2). A t a d ista n ce of
a p p ro x im a te ly 1 cm behind co llicu lis se m in a lis th e e re c tile tissu e s u r ­
ro u n d s th e u re th r a on all sides (P h o to 16— 3). A s p a rs d is se m in a ta p ro statae d evelo p s th e e re c tile tissu e of th e u re th r a d e cre a se s in size, r e ­
tain in g th e th ic k e s t la y e r ag a in on th e v e n tra l side of th e u re th r a
(P h o to 16— 4, 5).
T h e e re c tile tissu e th ick en s and ag a in su rro u n d s th e u r e th r a in th e
reg io n of th e p o ste rio r h a lf of th e p elv ic p o rtio n of th e u ro g e n ita l ca n a l.
The lu m en o f th e ca v e rn o u s sinuses m a rk e d ly in cre a se s, w h ich in th e
cro s s-se c tio n g iv es th e a p p e a ra n ce ty p ica l of e re c tile tissu e (P h o to
16— 9, 10, 11).
530
K . Sw iezynski
IX . TH E P EN IS
In th e penis of th e m ale E u ro p e a n bison, as in o th e r m a m m a ls, it is
p ossible to d istin gu ish c o rp u s p e n is and p a rs p en is s. p ars
u r e t h r a e su rro u n d ed by th e c o rp u s c a v e rn o s u s u r e t h r a e 2).
extern a
I. C o rp u s p en is
T h e c o rp u s p e n is begins a t th e ischial a rc h b y m ean s of p aired c ru ra
p e n is , w h ich n e x t u n ite to fo rm th e u n p aired r a d ix p e n is a tta c h e d by
stro n g sh o rt lig am en ts to th e p o ste rio r p a rt o f s y m p h y s is iscJtii. In tu rn ,
r a d ix p e n is e x te n d s cra n ia d into t ru n c u s co rp o ris p en is , ending in a p e x
c o rp o ris p en is . T r u n c u s co rp o ris p e n is fo r ' / 3 of its len g th to w a rd s th e
p elv is fo rm s th e »sigm oid fle x u re « w h ich is less d istin ct th a n in th e
d o m estic bull. Th e fle x u re did n ot o ccu r in ca lv e s up to 7 d a y s old in
th e m a te ria l w e d issected .
C o rp u s c a v e rn o s u m co rp o ris p e n is in th e reg io n of th e tw o c r u r a is
su rro u n d ed by tu n ica a lb u g in e a . T h is is re in fo rc e d by fa scia e of c i r ­
c u la r fib res a t first loosely k n it, b u t w h ich , as th e d ista n ce to r a d ix
p e n is d e cre a s e s, b ecom e in c re a s in g ly dense (P h o to 17— 1, 2). T h e d ist­
an ce b etw een c r u r a d e cre a se s even m ore not only on a cco u n t of th e
in c re a s e in th e am o u n t of co n n e ctiv e tissu e in th e w alls of c r u r a b ut
ch ie fly ow ing to th e ir oblique co u rse.
W h en th e m ed ial w alls of th e tw o c r u r a m eet a sep tu m is fo rm ed
b etw een th e e re c tile tissu e (P h o to 22 — 3, 4) w h ich loses its c h a r a c te r
of sep tu m n e a r r a d ix p en is , and as a re s u lt w e h av e an a p p a re n tly
sin g le c o rp u s c a v e rn o s u m p e n is in the reg io n of ra d ix p en is (P h o to
— 5 , 6, 7). A t th e sam e tim e th e co n ten ts of th e v a s c u la r co m p o n en ts
of th e c o rp u s c a v e rn o s u m d e cre a se co n sid e ra b ly in fa v o u r of th e fib rou s
tra b e cu la e . In th e cro s s-se c tio n th e tra b e cu la e ru n in fan sh ap e fro m
th e fib rou s tissu e » ce n tru m « w h ich ex h ib its a dense, co m p a ct s tr u c tu re .
In this in itial se ctio n th e »ce n tru m « is situ a te d im m e d ia te ly a b o v e th e
u re th r a l g ro o v e (P h o to 17 — 7). W h en o b serv in g su ccessiv e c r o s s -s e c t­
ions of th e penis, in a d irectio n up to the a p e x , it ca n be seen th a t th e
» ce n tru m « d escrib ed ab ove in cre a se s and is situ a te d m ed ially in fro n t
o f th e p ro x im a l c u rv e of th e sigm oid fle x u re . H ere it h as a la te ra lly
fla tte n e d sh ap e, and th e tra b e c u la e ra d ia te o u tw a rd s fro m it (P h o to
17 — 8). In th e reg io n of th e sigm oid fle x u re th e » ce n tru m « is fla tte n ­
ed d o rs o -v e n tra lly , w ith a te n d e n c y to division into tw o ow ing to the
2 The term inology used in describing the penis is in accordance w ith that used
by B u c h h o l t z (1951) and P r e u s s (1954) in describing the penis in domestic
anim als.
T h e m ale reproductive organs of the European bison
m edial n a rro w in g (P h o to 17 —
531
10, 11, 12). T h is division is ev en m o re
clearly m a rk e d in th e in itial p a rt of th e a p e x (cro ss -s e ctio n 13). In
fu rth e r cro ss-se c tio n s o f th e a p e x th e » ce n tru m « h a s th e a p p e a ra n ce of
an in cre a sin g ly fla tte n e d la m in a (P h o to 17 — 14, 15, 16), d isap p earin g
at a d istan ce of ab o u t 5 m m fro m th e end of th e a p e x . In v iew o f th e
fact th a t th e d escrip tio n s g iv en ab ov e r e fe r to cro ss-se c tio n s is m u st be
said th a t in th e E u ro p e a n bison a c e n tra l a x ia l band of dense fib ro u s
tissue fo rm ed b y th e tra b e cu la e of co rp u s c a v e rn o s u m p e n is w h ich
crosses in th is p lace insides sp re a d s th e e re c tile tissu e of co rp u s p en is.
The a p p e a ra n ce and position of th e a x ia l b an d d iffe r in d iffe re n t s e c t­
ions of co rp u s p en is and d epend on th e sh ap e of th e penis.
T h e m ale u re th r a , su rro u n d e d b y co rp u s ca v ern o su m , u r e t h r a e , is
lo cated along th e v e n tra l su rfa ce of th e b od y o f th e p en is inside th e
u re th ra l g ro o v e.
2. T h e penile p a rt of the u rog en ital d u ct
T h e p ro x im a l end of th e p en ile p o rtio n of th e u ro g e n ita l d u ct is s u r­
rou nd ed b y b u lb u s ca v e rn o s u s u r e t h r a e .
B u lb u s c a v e rn o s u s u r e t h r a e in th e E u ro p e a n bison is a t firs t lo ca te d
d o rsally , th en also la te ra lly in re la tio n to th e d u ct. It e x te n d s b e tw e e n
the b u lb o u re th ra l glan d s and th e p la ce in w h ich c r u r a penis u n ite. T h is
fo rm ed g rou n d s fo r callin g th is sectio n of th e d u ct — p a rs b u lb i u r e ­
th ra e ( G r a b o w s k i , 1937). A fte r re m o v in g m . b u lb o c a v e rn o s u s th e
bulbus in th e fix e d sp ecim en s is sim ila r in sh ap e to a steep te tra h e d ra l
p y ram id w ith its a p e x tu rn e d in th e d ire ctio n of th e penis (P h o to 15).
On th e d o rsal side of th is o rg a n th e re is a sh arp c r is ta h ig h est a t th e
end d ire cte d to w a rd s th e p elv is and b ecom ing lo w er v e n tra lly . T h e
d o rso -c a u d a l s u rfa c e s situ a te d on th e sides of this c r is ta a re slig h tly
co n ca v e in u n in jected p re p a ra tio n s. T h e la te ra l su rfa ce s a re su rro u n d ­
ed b y a th ick la y e r o f co n n e ctiv e tissu e. N e a r ra d ix p en is , w h e re th e
bulbus n a rro w s , th e la y e r of co n n e ctiv e tissu e e x te n d s to 20 m m on
e a ch sid e an d fills th e sp aces b e tw e e n c r u r a and bulbus. T h e bulb u s
b u ilt in th is w a y is u su a lly a sin gle s tru c tu r e . I t is o n ly in th e b ro a d e n ­
ed end o f th e b u lbu s n e a r th e p elv is th a t th e sep tu m d ividin g co rp o ra
ca v ern o s a b u lb i into tw o h a lv e s ca n be seen in cro s s-se c tio n s . T h is
se p tu m d isap p ears a t th e le v e l of ab o u t 1U o f the p ro x im a l le n g th of
th e o rg an .
T h e d im ension s of bulb us v a r y in d iv id u a lly in th e E u ro p e a n bison.
In th e m a te ria l d issected th e le n g t of b u lb u s in ad u lt bison (m e a su re d
alon g th e d o rsal c r is ta — F ig . 1 D 4— 6) v a rie d fro m 55 to 105 m m .
B r e a d th in th e w id est p lace (F ig . 1, 6, 7— 5, in p ro je ctio n ) w as m o re
s ta b le an d v a rie d fro m 26 to 28 m m .
532
K . Swiezynski
A t firs t co rp u s c a v e rn o s u m u r e t r a e d iffe re n tia te s into tw o la y e rs .
T h e in te rn a l la y e r, w ith sm a ll ca v e rn o u s sp aces, ad h eres to th e w a ll
of th e u re th r a ; th e e x te rn a l la y e r , w ith la rg e r sp aces, is lo ca te d p e ri­
p h e ra lly fro m th e fo rm e r. In this initial p a rt th e u r e th r a is situ a te d
n e a re r th e c o rp u s p e n is . T h e e x te rn a l la y e r of th e e re c tile tissu e, s itu a t­
ed v e n tra lly in re la tio n to th e u re th r a , is th ic k e r (P h o to 17 — 5, 6, 7).
B eg in n in g w ith th e p ro x im a l cu rv e of th e sigm oid fle x u re of th e p en is
and fu rth e r in th e d irectio n of th e a p e x co rp u s c a v e r n o s u m u r e t h r a e
su rro u n d s the u re th ra ev e n ly . A s a re su lt th e position of th e u re th r a
is ce n tra l in c o rp u s c a v e rn o s u m in th is p a rt of th e penis (P h o to 17 —
10, 11, 12). In th e p re p u tia l p a rt of the penis co rp u s c a v e rn o s u m u r e ­
th ra e th ick en s in th e p a rt situ a te d d o rsally fro m th e u re th r a (P h o to 17
— 15, 16). It ceases to be m a cro sco p ica lly visible w h en th e u re th r a
p asses into th e u re th ra l p ro cess.
T h e penile p o rtion of th e u r e th r a and its co rp u s c a v e rn o s u m a re
su rro u n d ed d u rin g its co u rse th ro u g h th e u re th ra l g ro o v e , fro m th e
b ack and th en fro m th e b o tto m , b y an e x te rn a l la y e r of th e tu n ica a l­
b u g in e a co rp o ris p en is. A t first th e p a rt of the penis to w a rd s th e pelvis
up to th e p ro x im a l c u rv e of th e sigm oid fle x u re is fla tte n e d la te ra lly .
T h e stro n g ly d evelop ed v e n tra l p a rt of th e co rp u s c a v e rn o s u m p en is
p ro tru d e s ab ov e th e u re th r a l g ro o v e and can be seen th ro u g t th e
tu n ica. In th e reg io n o f th e fle x u re the penis b eco m es cy lin d rica l in
sh ap e. C o rp u s p e n is co m p le te ly su rrou n d s th e u re th ra and th e e x te rn a l
la y e r of tu n ica e a lb u g in e a e th ick en s. T h e u re th ra to g e th e r w ith co rp u s
c a v e rn o s u m b eco m es invisib le fro m th e e x te r io r (P h o to 17 — 10, 11,
12). On th e a p e x of th e penis th e u re th ra is visib le o n ly in th e fo rm
of a s h o rt u re th ra l p ro cess.
T h e a p e x of th e penis is co v e re d from th e e x te r io r b y th e p re p u ce
p ro p er. B e tw e e n th e p re p u ce p ro p er and th e d o rsal s u rfa c e of co rp u s
p e n is th e re is an a g g re g a tio n of dense co n n e ctiv e tissu e fib res. T h e y
ru n obliqu ely fro m th e p re p u tia l la y e r to a p e x co rp o ris p e n is in serted
on it a t th e lev el of su lcu s g a lea e apicis. In th e reg io n o f ga lea apicis
p e n is s. ga lea g la n d is th e re is a thin la y e r of co n n e ctiv e tissu e w ith
visib le blood v essels, fo rm in g th e b o rd er of ga lea a picis p en is .
G a lea a picis p e n is in th e E u ro p e a n bison is rou nd in sh ap e, w ith an
a p e x d irected b a ck w a rd s and to th e rig h t. T h e re is no d istin ct lin e of
d e m a rca tio n b etw een g a lea e p e n is and th e p o ste rio r p re p u tia l p a rt of
th e a p e x penis. A t th e lev el of th e beginning of th e u r e th r a l process
on th e opposite (rig h t) side of ra p h e p re p u tii th e r e is a p ro m in en ce,
w h ich
in the c a d a v e rs h ad sh a rp m arg in s and ad h e re s fla tly to th e
u re th ra l p ro cess fro m th e rig h t side. In th e o rg an s ta k e n fro m ca d a v e rs
The m ale reproductive organs of the European bison
533
fixed b y m ean s of in tr a a r te r ia l in je ctio n s th e su rfa ce of this p ro m i­
n en ce is sm oo th and co n v e x . T h e w hole h as th e a p p e a ra n ce of a low
tu b ercle ab ou t 3 m m in d ia m e te r.
3. Dimensions of the penis
T ab le 8 g iv es th e re s u lts of m e a su re m e n ts of th e le n g th of th e penis
in E u ro p e a n bison, m e a su re d alon g th e v e n tra l su rfa ce (F ig . 1, E , 8— 9).
It w ould seem th a t th e r e la tiv e ly la rg e d iffe re n ce in to ta l le n g th o f th e
penis and len g th of the p rep u tia l p a rt b etw een th e la st tw o ag e grou p s
is due to th e f a c t th a t th e re w e re only tw o in d ivid u als in th e th ird
grou p.
Table S.
M easurem ents of length of penis in the European bison (in mm).
Age lim its
1— 7 days
1— 10 months
2— 5 years
over 5 years
Length of p a rs p rep u tialis
p en is
Number
of
cases
Total length of penis
From
To
Mean
From
To
3
2
2
11
206
309
630
755
252
340
740
965
231
324
685
864
34
50
80
120
36
58
115
160
Mean
35
54
111
139*
*) — Num ber of cases fo r this m ean value = 10.
C alcu la tio n w as m ade of th e ra tio b etw een le n g th of penis and
m e a s u re m e n ts of body le n g th o f th e ind ivid u als d issected (T ab le 9 ). In
E u ro p e a n b ison aged fro m one d a y to te n m o n th s old th e se v a lu e s a re
d iffe re n t an d in th e m a jo rity of cases e x ce e d th e co rre sp o n d in g v a lu e
fo r bison o v e r th re e y e a rs old. In tu r n in th e re m a in in g m a le s th e ra tio
b e tw een p enis len g th and co rresp on d in g b od y m e a su re m e n ts is co n sta n t.
X . TH E P R E P U C E
T h e a p e x of th e penis, to g e th e r w ith ga lea apicis p e n is , is co v e re d by
th e p ra e p u t iu m (P h o to 21). T h e in te rn a l la y e r of th e p re p u ce b egin s
ro u n d o stiu m p ra e p u tia le w ith a n a rro w band of p ig m en ted skin. T h e
skin su rro u n d in g th e o stiu m is co v e re d w ith se ta ce o u s h a irs , p a rtic u la rly
d en se an d long on th e d orsal side (P h o to 19). It is s tr o n g ly th ick en ed
an d th e lu m en of th e o stiu m n a rro w (in fixed ca d a v e rs 1 to 1.5 cm in
d ia m e te r).
F ro m th e e x te r n a l su rfa c e of th e ab d om in al w all u n d er th e th ick e n e d
sk in o f th e p re p u tia l o stiu m a th ick
fla t
bundle
of
co n n e ctiv e tissu e
K. Św ieżyński
534
fib res ra d ia te s . It is lig a m e n ta l in c h a ra c te r and stre n g th e n s th e a tta c h ­
m en t of th e p rep u ce to th e w alls of the body. Inside th e p rep u ce, fro m
th e back to th e o stiu m , th e r e is a region fro m 120 to 150 m m long,
co v ered b y lon g itu d in al folds. On th e d o rso cra n ia l w all in five bison
th e re w as a d a rk ly p ig m en ted field co v ered w ith sc a n ty , s h o rt h airs.
Table 9.
R a tio of length of penis to extern a l body m easurem ents.
Ratio to m easurem ent
No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
Name
A
E
F
Pos
Pub
Pom
P la k a t
P ustybór
1:5.6
1:4.5
1:4.5
1:4.3
1:5.4
1:22
1:1.8
1:1.9
1:2.2
1:3.1
1:2.6
1:2.5
1:3.2
1:2.9
1:3.3
Average (age of 1 day to 7 months)
1:4.9
1:2.3
1:2.9
Poronin
P utnar
1:3.4
1:3.3
1:1.5
1:1.7
1:2.2
1:2.4
A verage (age of 2— 3 yrs.)
1:3.4
1:1.6
1:2.3
Puzor
Pug II
Pozew
P ołam aniec
Pokorny
1:3.2
1:2.9
1:3.3
1:3.1
1:2 9
1:1.5
1:1.4
1:1.6
1:1.5
1:1.5
1:2.1
1:2.1
1:2-5
A verage (age of 5— 10 yrs.)
1:3.1
1:1.5
1:2.2
Posusz
Pluszcz
P lam iec
P lato
1:3.3
1:3.4
1:3.2
1:3.0
1:1.5
—
1:1.5
1:1.4
1:2.5
1:2.2
1:1.7
A verage (age over 10 yrs.)
1:3.2
1:1.5
1:2.1
—
1:2.3
—
A n e x te n sio n o f th is reg io n cau d a d is a zone of c irc u la r folds fro m 80
to 100 m m lon g. T h e p re p u ce p ro p e r — la m in a p en is p ra ep u tii,
en sh eath es th e penis fo rm in g se v e ra l c irc u la r folds. R a p h e p ra ep u tii
ru n s alon g th e v e n tra l s u rf a c e o f th e a p e x penis, to w a rd s th e rig h t side,
and a t th e lev el of th e b ase of th e u re th ra l p ro cess d isap p ears, in th e
ca se of y o u n g in d ivid u als, in n u m ero u s lo n g itu d in al folds. In a d u lt bison
th e rap h e ends sin gly .
In E u ro p e a n bison up to te n m o n th s old p a rie ta l and v is ce ra l lam in ae
p ra e p u tii co h ered . In sev en m o n th s old »P om « and one m o n th old » P la -
535
T h e m ale reproductive organs of the European bison
k at« this ad h esion w as co m p lete. In te n m o n th s old » P u sty b o r« th e
dorsal su rfa ce of ga lea apicis penis w as a lre a d y fre e . T h e v e n tr a l su rfa ce
of th e g ale a w as still a tta ch e d to th e p a rie ta l la y e r of th e p re p u ce b y
n u m ero u s sm all b an d s (P h o to 18).
X I. TH E P R E P U T IA L M U SC LES AND TH E E X T R E N S IC P E N IL E M U SC L E S
S e v e ra l m u scles a re co n n ected to p o g ra p h ica lly and fu n ctio n a lly w ith
the g en ital o rg an s. S om e of th em , su ch as m . c r e m a s t e r e x t e r n u s , o r th e
grou p of m u scles situ a te d in th e reg io n of th e b u lb o u re th ra l glan d s,
h av e a lre a d y b een m en tion ed in th e a p p ro p ria te sectio n s of th is stu d y .
M . u re th ra lis w as d escrib ed in th e stu d y on the u rin a r y sy ste m of th e
E u ro p e a n bison (P i 1 a r s k i, 1 967). A m on g th e m u scles co n n e cte d w ith
th e p rep u ce an d penis of th e E u ro p e a n bison m en tio n m u st be m a d e of
th e p rep u tial m u scles, m . b u lb o c a v e rn o s u s , m . is c h io c a v e m o s u s an d m .
re t r a c t o r p en is.
M m . p ra e p u tia le s o ccu r in E u ro p e a n bison in th e fo rm o f p a ire d
c ra n ia l and cau d al m m . p ra e p u tia le s . T h e y a re co n n ected w ith th e skin
m u sc u la tu re o f th e tru n k .
M m . p ra e p u tia le s cra n ia les b egin a t th e level of lin ea a lb a cau d ad
fro m th e xip h o id p ro cess. T h e fla t bellies of th e m u scle s in y o u n g
in d ivid u als a t first a re in c o n ta c t w ith e a ch o th e r, su rro u n d th e u m b ilicu m an d ru n in th e d irectio n o f th e p re p u ce . In a d u lt E u ro p e a n bison
th e m u sc u la r b ellies a re s tro n g ly b ro ad en ed (up to 14 cm w id e in th e
b ro a d e st p a rt), w ith a b ow -sh ap ed co u rse d ire cte d to th e e x te r io r in
a la rg e cu rv e . T h e fla t term in a l ten d o n of ea ch of th e m u scle s e m b r a ­
ces, fro m th e e x te r io r , th e lig am e n t su sp end in g th e o stiu m of th e p re p u ce
an d p a r tly join s w ith th e ten d o n on th e o th e r side and p a rt of th e fib res
ra d ia tin g in to th e su b cu tan eo u s tissu e in th e a re a of th e o stiu m .
In so m e in d ivid u als, in add ition to th e bellies d escrib ed , th re e o r fo u r
b u n d les fro m cu tan eu s tru n c i
p re p u tia l o stiu m .
m u scle
ra d ia te d
into th e re g io n of th e
M m . p ra e p u tia le s ca u d a le s a re less stro n g ly fo rm ed th a n th e p re ce d ­
ing. T h e y begin in th e d eep fascia o f th e tru n k and on th e a p o n e u ro sis
of m . o b liq u u s a b d o m in is e x t e r n u s a t th e level of th e d ivision o f th e
p u d en d al e x te rn a l v essels. T h in m u scle bellies ru n along th e ab d om in al
w all an d ra d ia te b etw een th e e x te rn a l and p a rie ta l la y e rs of th e p re p u ce ,
5 to 10 cm cra n ia d fro m f u n d u s p ra e p u tii.
M . b u lb o s p o n g io s u s s. b u lb o c a v e rn o s u s
ru n s
fro m
th e
b asis
of
th e
b u lb o u re th ra l g lan d s and e x te n d s alon g th e d orsal su rfa ce of th e b u lb u s
of th e u ro g e n ia l can a l. On the sides th is m u scle is bou n d ed b y m m .
isc h io c a v e rn o s i. It fused w ith th e m alo n g th e w hole le n g th of c o n ta c t.
536
K . Sw iezynski
T h e p ro x im a l th ick e st and w id est p a rt o f m . b u lb o c a v e m o s u s (up to
6 cm w ide and 4 cm th ick ) p ro tru d e d in fro n t of th e a n te rio r m a rg in
of th e bulbus. T h is p a rt fo rm s a cy lin d rica l m u scu la r p ro m in en ce a b o v e
th e bulbus and b u lb o u re th ra l glan d s. In th e cra n io -d o rsa l p a rt th e
m u scle bundles, seen fro m th e b ack , ta k e a se m ic irc u la r co u rse. T o w a rd s
th e ra d ix p e n is th e m u scle b ecom es in cre a sin g n a rro w and th e b u n d les
of fib res ru n obliqu ely (P h o to 20). On th e d o rso -ca u d a l su rfa ce of th e
m u scle along th e m ed ian lin e th e re is a lo n g itu d in al sep tu m . In th e
lo w e r th re e q u a rte r s in th e e x te n sio n of th e sep tu m th e re is a lo n g i­
tu d in al g ro o v e. T h e d orsal c re s t of th e b u lb u s of th e u ro g e n ita l ca n a l
fo rm s the co n tin u atio n of th e se p tu m into th e in te rio r of th e m u scle.
M . b u lb o c a v e m o s u s ends, below th e te rm in a tio n of b u lb u s u r e t h r a e a t
a d istan ce of 4 .5 to 6.5 cm fro m th e lin e of th e lo w e r re a c h of th e m m .
isch io ca v ern o si. T h e len g th of m m . b u lb o c a v e m o s i v a rie s w ith th e
in d ivid u al and d ep en ds on th e le n g th of bulb u s
m u scle is a lw a y s lo n g e r th a n it.
In o u r m a te ria l th e m ean
10 .2 cm .
in th e sen se th a t th e
len g th of th e m u scle
in a d u lt bison w a s
M . isc h io c a v ern o s u s s. isch in sv o n g io su s is stro n g ly d evelop ed in th e
E u ro p e a n bison (P h o to 20). It begins on t u b e r ischii cau d ed an d c e n tra d .
It su rro u n d s c r u r a p e n is and is in serted to tu n ica a lb u g in e a of th e c r u r a
and p a rtly of ra d ix p en is.
A bundle of fib res, th e e x te r n a l a p p e a ra n ce and co u rse of w h ich
ju s tify th e ir b ein g tre a te d as in d ep en d en t m u scles, ru n fro m m . isc h io ­
ca v ern o s u s . A m o n g th e m is th e m u scle o ccu rrin g in all th e E u ro p e a n
bison w e d issected , m . is c h io u re th ra lis an d , in som e bison, m . isch io b ulbosus (b o th d escrib ed to g e th e r w ith th e b u lb o u re th ra l glan d s). In
tw o cases (» P u tn a r « , »P o k o rn y « ) th e bundle of fib res ru n n in g cra n ia d
and in serted in th e m ed ian line fro m th e d orsal s u rfa c e of a p o n eu ro sis
m . u re th ra lis se p a ra te fro m m . isch io u reth ra lis. On th e w a y th e y p a rti­
cip ate in th e fo rm a tio n of th e »m u scle sa c« fo r th e b u lb o u re th ra l glands.
M . re tra c to r p e n is s. re c t o p e n in u s begins w ith a v e rte b ra l in sertio n
on th e secon d , th ird o r b oth co ccy g e a l v e rte b ra e , la te ra lly fro m th e
v a s c u la r p ro cesses of th e v e rte b ra e . T h e fla t tend on of this m u scle
em erg es fro m u n d e r m . sa c ro co ccy p e u s v en tra lis la tera lis and e n te rs
u n d er m . lev a to r ani. T h is tend on is join ed by b u n dles of fib res fro m
m . lev a to r ani, m . s p h in c te r ani e x t e m u s and in one ca se (» P o m ru k « )
fro m m. iran su e rstts p e rin e i.
B elo w th e an u s b oth m m . re tra c to r e s p en is a re situ a te d in a g ro o v e
fo rm ed b y m . isch io cau ern o sits and m . b u lb o c a v e m o s u s (P h o to 20). B elow
th e a p e x of m . b u lb o c a v e m o s u s w h e re th e penis is m o st fla tte n e d
la te ra lly , th e tw o m m . re tra c to r e s p e n is ap p ro a ch e a ch o th e r. S ta rtin g
The m ale reproductive organs of the European bison
537
fro m th e p ro x im a l c u rv e of th e sigm oid fle x u re of th e p en is th e y a r e
su rrou n d ed to g e th e r b y fa scia and a re lo ca te d below th e penis (P h o to
21). A t th e level of th e d istal c u rv e , w h e re the p en is b eco m es cy lin d rica l
in shape, m m . re tra c to re s p e n is a re a tta c h e d to th e penis v e n tr o -la te r a lly
and fu rth e r ru n alon g th e la te ra l su rfa ce s of th e penis. In th e d ire ctio n
of the a p e x th ese m u scles b eco m e in cre a sin g ly th in , ag a in p ass on to
th e v e n tra l su rfa ce of th e penis and end in tu n ica e p en is n e a r th e p la ce
of tra n sitio n of th e p a rie ta l la y e r of th e p re p u ce into th e p re p u ce
p ro p er.
X I I . R E S U L T S AND D ISC U SSIO N
L ittle h as h ith e rto b een w ritte n on th e su b je ct o f th e m o rp h o lo g y of
the re p ro d u c tiv e o rg an s of th e E u ro p e a n bison, and in fa c t o u r k n ow ­
ledge of th is a sp e ct of th e A m e ric a n b ison ’s clo sest re la tiv e — Bison
bison L i n n a e u s , 1758 is sim ila rly lim ited . In co m p arin g th e re s u lts
of o u r ow n in v estig atio n s w e p e rfo rce m ad e use of d a ta co n ta in e d in
co rresp o n d in g lite r a tu r e on o th e r ru m in a n ts, ch ie fly c a ttle . T h is w ould
ap p ear to be ju stified on acco u n t of th e a b u n d an ce of lite r a tu r e on th is
sp ecies and its p h y lo g en etic re la tio n to th e E u ro p e a n bison.
1. T esticles, epididym is and scrotum
In o rd er to ch eck w h e th e r th e v iew s e x p re sse d b y O w e n (1 848) and
M u l l e r (1852) on th e size of th e te stic le s in th e E u ro p e a n bison a re
co n firm ed b y stu d ies o f a la r g e r a m o u n t of m a te ria l, co m p a riso n w as
m a d e of th e re s u lts of o u r ow n o b se rv a tio n s w ith d a ta on th e d o m e stic
T able 10.
Comparison of m easurem ents and w eight of testicles in European bison
and domestic rum inants.
Species
L en gth mm
B read th mm
W eight g
Bison bon asu s
d ex {r.
sinistr.
87— 115
4 8 -7 1
110—287
8 8 —121
45—69
96—279
Bos taurus
Ovis aries
1 4 0 -1 7 0
5 0 -6 0
250—300
100
50—60
200—300
Capra
93— 108
46 —54
145— If 0
bull (T ab le 10). D ata on th e E u ro p e a n bison w ere o b tain ed fro m fix e d
m a te r ia l, and th u s m a te ria l w ith re d u ce d d im ension s and w e ig h t. S c h u ­
b e r t (1909) sta te d th a t lin ear losses in th e c a rd ia c m u scles of c a ttle as
th e re s u lt of fix in g a re ab o u t 3 % . T h ese d iffe re n ce s u n d o u b ted ly v a r y
538
K . S w ieiy ń sk i
w ith d iffe re n t o rg an s. I t is n e v e rth e le ss possible to sta te t h a t the le n g th
of te s tic le s in a d u lt E u ro p e a n bison is f a r sm a lle r th a t th e le n g th o f
te stic le s in th e d o m e stic bull. T h e b re a d th of the te sticle in th e E u r o ­
p ean bison is, on th e o th e r h an d , sim ila r to th e corresp on d in g d im en sion
in c a ttle . R atio s b etw een le n g th an d b re a d th of te stic le s in th e E u r o ­
p ean b ison thu s d iffe r fro m th o se in d om estic bulls and a re u n e x p e c te d ly
clo se r to th e ra tio s fou nd in sm all d o m e stic ru m in an ts.
C lasses of v a ria tio n s in w eig h t of th e testicles in ad u lt E u ro p e a n b i­
son a re fa r w id er th a n those a cce p te d b y S c h w a r z (1939) fo r th e
d o m estic bull. It is p a rtic u la rly th e lo w e r lim it of v a ria tio n w h ich is
sh ifted in m in us, and as a re s u lt th e m ean w eig h ts of te sticle s in th e
bison a r e sm a lle r th a n in d o m e stic bulls and clo se r to th e ra tio s
o ccu rrin g in th e ra m .
In th e case of tw o E u ro p e a n bison w e w ere ab le to w eig h t th e
te stic le s b efo re an d a fte r fix in g . W e ig h t loss in th ese ca se s w as a b o u t
4.5% .T h e in clu sio n in o u r d iscu ssion s of this loss (alth ou g h th e f a c t t h a t
only tw o ca se s w e re exam in e d did not en title us to do so) did n o t in
p rin cip le a ffe c t th e co n clu sion s fo rm u la te d above.
A s c a n be seen fro m ta b le 2 th e dim ensions, and p a rtic u la r ly th e
w eig h t o f th e te s tic le s in ad u lt E u ro p e a n bison, e x h ib it co n sid e ra b le
v a ria tio n p er in d ivid u al. T h e m a te ria ] exam in ed leads to th e co n clu sion
th a t th e d ifferen ces a re n ot co n n ected w ith body m e a su re m e n ts (cf. d a ta
in ta b le s 1 and 2, re fe rrin g to » P u g II«, » P o łam an iec«, » P o k o rn y «) b u t
r a th e r w ith age. T h e re is n ot d istin ct p red om in an ce in size an d w eig h t
b etw een th e rig h t an d le ft te sticle s.
C o n n e ctiv e tissu e e lem e n ts in th e te sticle s of th e E u ro p e a n bison a re ,
fro m th e p oint of v ie w o f m a cro sco p ic s tru c tu re , m o d e ra te ly d eveloped .
T h e th ick n ess of tu n ic a a lb u g in e a v a rie d from 1.05 m m to 1.4 6 m m (in
c a ttle fro m 0.5 to 1.5 m m — K r o l l i n g & G r a u , 1960). T h e th ick n ess
of th e m e d ia s tin u m testis, th e m a cro sco p ic p ictu re of w hich d ep en ds on
th e size an d t e x tu r e of co n n e ctiv e tissu e, is far sm a lle r th a n in th e
te sticle s of th e d o m estic bull. M a x im u m th ick n ess of m e d ia s tin u m te stis
n e a r th e e x t re m it a s capitata in th e E u ro p e a n bison w e d issected w as
4 m m . In c a ttle th is m e a s u re m e n t is 6— 8 m m ( S c h w a r z , 1 9 3 9 ;
K r o l l i n g & G r a u , 1960). A lso th e sep ta of the te sticle ru n n in g fro m
m e d ia s tin u m testis a r e o n ly fa in tly visib le m acro sco p ica lly (P h o to 3).
T h e te stic le s in b o th th e o n e -d a y old bison ca lv e s d issected had
a lre a d y d escen ded below th e e x te r n a l inguinal rin g. This is evid en ce
th a t d e s c e n s u s te s tic u lo ru m o ccu rre d in principle d u rin g in tra u te rin e
d ev elo p m en t. T ak in g as an e x a m p le , u n fo rtu n a te ly th e only one in our
m a te ria l, th e te n -m o n th s old » P u sty b ó r« , it m igh t be co n sid ered th a t
th e fin al lo catio n o f th e te sticle s in th e scro tu m o ccu rs a t a n ag e o v e r
The m ale reproductive organs of the European bison
ten m on th s. A cco rd in g to V I o t e n
539
(1 9 2 7 ) this ta k e s p la ce f a r e a rlie r
in d om estic bu lls, a t th e ag e of a b o u t 3 m o n th s. In th e E u ro p e a n bison,
as a w ild livin g sp ecies, p ro ce sse s co n n ected w ith th e d e sce n t o f th e
te sticle s w ould ap p e a r to la st lo n g e r. It m u st h o w e v e r be p oin ted o u t
th a t o u r o b serv atio n s in th is re s p e c t r e f e r to one in d ivid u al only.
The ab ove p ro viso is th e m o re n e ce s sa ry in v ie w of th e f a c t th a t in
th re e E u ro p e a n bison d is tu rb a n c e s w e re found in th e p ro ce ss of d e sce n t
of the testicles. F o u r - y e a r old »P u ck « exh ib ited b ila te ra l c r y p to o rc h is m u s, th e te stic le s being situ a te d w ith in th e ab d om in al c a v ity , th e rig h t
a t a d istan ce of ab ou t 5 cm and th e left ju st n e a r th e in te rn a l in g u in al
rin g. T h e long a x is of th e te stic le s w as lo ca te d h o riz o n ta lly . Di­
m ensions — len g th of rig h t te stic le 90 m m an d le ft 70 m m — a r e sim ila r
to th e co rresp o n d in g v a lu e s in n o rm a lly d eveloping te stic le s . The
te sticle s co v ered b y th e p e rito n e u m w e re d ire cte d w ith th e ep id id ym al
b o rd er to th e ab d om in al w all, and th e fre e b ord er in w a rd s. P lica d u c tu s
d e fe r e n t is e x te n d e d to th e e x t r e m it a s ca u d a ta and th e p lica v a scu la ris
to the e x t re m it a s capitata of th e te sticle . T h e b road m , c r e m a s t e r e x te rn u s join s th e co v e rin g s tr u c tu r e s of th e te sticle s in th is sa m e p lace.
T h e v a g in a l p ro cess p e n e tra te d to th e b ila te ra l p a te n t in g u in al ca n a ls
an d ended a t th e lev el of th e e x te r n a l in g u in al rin g.
In te n -y e a r old »P osu sz« th e p ro cess of d escen t of th e rig h t te stic le
w as a rre s te d a t th e level of th e in g u in al ca n a l. T h e te stic le h a d r e m a in ­
ed in th e ca n a l and w as r e la tiv e ly long and
d ev iatio n s in s tru c tu re w e re found.
The rig h t te s tic le of fo u rte e n -y e a r old
ingu in al can al b u t h ad n ot d escen d ed into
s c ro ta l c a v ity on th e rig h t side. T h e fu n d u s
n a rro w (T ab le 2). No o th e r
» P om ru k « w a s o u tsid e th e
th e sc ro tu m . T h e re w a s no
of th e rig h t v a g in a l p ro cess
re a c h e d th e level of th e c o llu m scro ti. T h e rig h t te s tic le w a s irre g u la r
in sh ap e and th e d im ension s o f th e rig h t ep id id ym is e n o rm o u s. B o th
»P osu sz« and »P o m ru k « co v e re d fe m a le bison and had p ro g e n y .
If w e do not ta k e th e f ir s t fiv e E u ro p e a n bison (T ab le 1) in to c o n ­
sid e ra tio n on a c c o u n t of th e u n fin ish ed p ro cess of d e sce n t o f th e te stic le s
fou nd in th em , th e n th e fig u re of th r e e d e v elo p m en tal d is tu rb a n c e s of
th is kind o u t of fifteen E u ro p e a n bison m u st be co n sid ered a s a co n ­
sid erab le p e rce n ta g e . C ry p to o rch ism u s h a s been re co rd e d in all sp ecies
of d o m estic an im als and in m a n y w ild sp ecies ( Z e d e n o v , 1965).
A m o n g ru m in a n ts th is o ften o c cu rs in ra m s ( E c k s t e i n & Z u c k e r ­
m a n n , 1956), re a ch in g as m u ch as 12.5% in som e b re e d s ( W e n d t
et al., 1960).
T h e ep id id ym is d efines th e ep id id ym al b o rd e r of th e te s tic le on th e
ca u d o la te ra l side. T h is s ta te m e n t ap p lies to th e re la tio n s o b se rv e d in
c a d a v e rs. T h e te stic le s of ru m in a n ts , on a cc o u n t of th e ir sh ap e, v e rtic a l
K . Sw iezynski
540
position and w a y in w h ich th e y a re a tta c h e d , a re e asily rev o lv ed rou nd
th e lon g a x is. T h is m ay also o ccu r as th e re s u lt of p o stm o rtal ch an g es
(e .g . d iffe re n ce in tension of m . c r e m a s t e r e x t . o r a position of th e body
d iffe re n t fro m th e p h ysio lo g ical one). T h e d e scrip tio n of th e position
of th e te sticle s and ep id id ym is in th e sc ro tu m of the d om estic bull based
on o b serv atio n s m ad e in th e d issectio n ro o m (re p e a te d o v e r a long
p eriod of y e a rs in a n a to m ica l p ap ers an d textb o o k s) had to be co rre c te d
a fte r co m p ariso n w ith th e re s u lts of e x a m in a tio n s m ad e on livin g
an im als. F o r obvious rea so n s it w a s im possible to c a r r y o u t su ch
e x a m in a tio n s on livin g E u ro p e a n b ison and w e w e re th e re fo re obliged
to c o n te n t o u rselv es w ith a n a to m ica l la b o ra to ry m a te ria l.
T h e b re a d th s of d ifferen t p a rts of th e epid id ym is in c a ttle is g iv en
by S c h l u m p e r g e r (19 5 4 ). C om p arison o f th ese d a ta show th a t th e
h ead of th e ep ididym is is fa r b ro a d e r in th e E u ro p e a n bison. The
b re a d th of th e co rp u s e p id id y m is is sim ila r, b u t th e c le a rly defined
n arro w in g of th e m ed ian p a rt o f the co rp u s p re se n t in c a ttle does not
o ccu r in E u ro p ean bison. A m o n g o th e r d ifferen ces is th e a p p e a ra n ce of
th e tail of th e ep id id ym is. In c a ttle it is sim ilar in shape to a hood
p ro tru d in g in th e e xten sio n of th e long a x is of th e co rp u s ( S c h w a r z e ,
1939) w hile in th e E u ro p ea n bison it is set m o re to th e back (F ig . 2).
B o th S c h w a r z e (1939) and S c h l u m p e r g e r (1954) co n sid er th e
p osition o f m . c r e m a s te r e x t. in d o m e stic an im als a s a fe a tu re co n sta n t
to th e species. In th e d o m e stic bull th e m u scle co v e rs th e ca u d a l and
m ed ial su rfa ce s of th e sp e rm a tic co rd ( S c h l u m p e r g e r , I.e.). M u scle
fib res end ab ov e th e h ead of th e ep id id ym is ( S c h w a r z e , I.e.). In th e
ra m th e m ed ian su rfa ce s of th e co rd s a re n ot co v e re d ( S c h l u m p e r ­
g e r , i.e.). Th e m u scle ends a t th e level of th e h ead of th e ep id id ym is
(Schwarze,
I.e.).
In th e E u ro p e a n bison re la tio n s a re d iffe re n t. M. cre m a s te r e x t . co v e rs
th e sp e rm a tic co rd s c a u d o la te ra lly (P h o to 2, A ), leav in g an u n co v e re d
c ra n ia l and m ed ian su rfa ce s . T h e position of th e o th e r co m p o n en t
e lem en ts of th e co rd s and th e p re v io u sly discussed situ a tio n of th e
ep id id ym is in re la tio n to th e te sticle s c re a te s a situ a tio n in th e ca se of
th e E u ro p e a n bison in w h ich th e te sticle s to g e th e r w ith th e sp e rm a tic
co rd s a re re v o lv e d b y ab o u t 9 0 ° in co m p ariso n w ith c a ttle . T h e sw ing
on th e le ft side took p lace to th e rig h t, and on th e rig h t side to th e le ft
rou nd th e long a x is. D oes th is co rresp o n d to th e situ a tio n of th e te stic le s
and sp e rm a tic co rd s in liv in g E u ro p e a n bison? A n a rg u m e n t ag a in st
accep tin g th e ab o v e-m en tio n e d situ a tio n as p h ysio lo g ical is fo rm e d b y
th e fa c t th a t it is sim ilar to th e e a rlie r d escrip tio n of re la tio n s in c a ttle
( M a r t i n - S c h a u d e r , 1 938; S c h w a r z e , 1 9 3 9 ; E l l e n b e r g e r - B a u m , 1943) w h ich p ro v ed to be d iffe re n t fro m th a t estab lish ed by
The male reproductive organs of the European bison
541
exam in atio n s o f liv e a n im als ( S c h l u m p e r g e r , 1 954; N i c k e l et
al., 1960). It m a y th e re fo re be co n sid ered b y a n a lo g y th a t th e c a s e is
sim ilar in th e E u ro p e a n bison. O n th e o th e r side th e follow in g a rg u e in
fa v o u r of th e position of the te stic le s in th e E u ro p e a n bison b eing
d ifferen t fro m th a t in th e d o m e stic b u ll: a) th e fa c t th a t th e le ft te stic le
is sh ifted fo rw a rd s in fro n t of th e rig h t and th e d iffe re n ce s in th e shape
of th e sc ro tu m , b) th e d ifferen ce in th e e x te n t of m . c r e m a s te r e x t e r n u s .
T he s c ro tu m in E u ro p e a n bison is se t fu rth e r cau d ad in co m p ariso n
w ith th e re la tio n found in d o m estic bu lls. It is also s h o rte r th a n in
bulls, w h ich w ou ld a g re e w ith th e g e n e ra l p rin cip le in th e g ro u p of
an im als w ith a »p arap en ial« ty p e of sc ro tu m , a cco rd in g to w h ich th e
len g th of th e s c ro tu m in cre a se s w ith in cre a sin g d ista n ce fro m th e an u s
( Z e d e n o v , 1965). In E u ro p e a n bison up to th e ag e of th re e th e
scro tu m is v e r y sh o rt, w ith fa in tly m a rk e d sh ift fo rw a rd s of th e le ft
p a rt. In ad u lt E u ro p e a n bison it is m o re pendulous an d th e fo rw a rd s
sh ift of th e le ft p a rt is m o re d istin ct. A s a re s u lt th e lo n g itu d in al
g ro o v es along r a p h e scro ti are located not c ra n ia lly and ca u d a lly as in
the d o m estic b u ll, b u t th e a n te rio r g ro o v e d e v ia te s to th e rig h t an d th e
p o ste rio r to th e le ft. T h e w hole gives th e im p ression of la te ra l fla tte n in g ,
w hich is due on th e one han d to th e ca u d a l situ a tio n o f th e sc ro tu m
itself, and on th e o th e r to th e n a rro w sp acin g of th e E u ro p e a n b ison ’s
hind lim bs.
T h is lo catio n of th e s c ro tu m w ould seem to be of a d v a n ta g e u n d er
n a tu ra l livin g co n d ition s. T h e high su spension re d u ce s th e d a n g e r of
in ju ry w h en th e an im al m ak es its w a y th ro u g h d en se u n d e rg ro w th ,
w hile th e legs fo rm som e p ro te ctio n for th e scro tu m fro m th e sides.
2. The gen ital accesso ry glands
O u r d escrip tio n of th e s tru c tu r e of th e a m p u lla d u ctu s d e fe r e n t is in
th e E u ro p e a n bison in p rin cip le co rre sp o n d s to th e d e scrip tio n g iv e n by
K r 6 1 l i n g (1 9 3 0 ), w ith th e e x ce p tio n of th e re m a rk s as to th e size of
th e a m p u lla. T h e in d ivid u al d issected b y K r o 11 i n g ( i . e . ) as fo u r­
- y e a r s old b elo n ged to an im als w ith in com p lete g ro w th p rocess an d th e
7 0 m m len g th o f th e am p u lla does in fa c t co m e w ith in th e lim its
a cc e p te d fo r th e ra m (6 0 — 80 m m — S c h u m m e r & N i c k e l , 1960),
In o u r m a te ria l a m p u lla e d u c t u u m d e f e r e n t i u m w ith sim ila r m e a s u re ­
m e n ts w e re fou nd in E u ro p e a n bison tw o and th re e y e a rs old. In E u r o ­
p e a n bison fro m fiv e to te n y e a rs old th is d im ension is g r e a te r and
co m e s w ith in th e lim its acce p te d fo r c a ttle (80— 150 m m — N i c k e l ,
1 9 5 4 ). T h u s n o t o n ly th e s tru c tu re b u t also th e len g th of a m p u lla e in
th e E u ro p e a n b ison is sim ila r to th a t in d o m estic c a ttle .
542
K . Sw iezynski
V e s ic u la e se m in a les do n o t d iffe r in e ith e r s tru c tu r e o r position fro m
those in c a ttle . T h e sligh t d ev elo p m en t of th ese g lan d s in th e E u ro p e a n
bison w h ich M ü l l e r (1852) d escrib ed ca n n o t in a n y case be co n sid ered
as a c h a r a c te r of th e sp ecies. K r ö l l i n g (1 9 3 0 ) also adopted th is
stan d p o in t. T h e d im ensions h e g iv e s (len g th 100 m m , b read th 25 m m )
are sim ila r to th e m ean v a lu e s w h ich w e o b tain ed (cf. T ab le 6) an d
com e w ith in th e lim its acce p te d fo r c a ttle (len g th 70— 120 m m — N i c ­
k e l , 1954). T h e ab sen ce in th e E u ro p e a n bison of th e fle x u re s of th e
cau d al p a rts of th e se g lan d s ty p ica l o f c a ttle re p o rte d b y K r ö l l i n g
(i. c.) is due to th e f a c t t h a t th e E u ro p e a n bison w h ich he d issected w as
n ot fu lly g ro w n . In bison o v e r five y e a rs old th e fle x u re s on th e e x ­
te rio r of th e te rm in a l p a rts of th e glands a re visib le (P h o to 6, 7).
Th e p ro sta te in th e E u ro p e a n bison is fo rm ed b oth in th e e x te rn a l
p a rt — th e co rp u s and in th e in te rn a l — p a rs d issem in a ta p ro sta ta e.
T h e m ean tra n s v e rs e
m e a s u re m e n t
of th e
co rp u s
is sim ilar, and th e
lo n g itu d in al m e a su re m e n ts slig h tly sm a lle r, th a n th e dim ensions g iv en
b y N i c k e l (1954) fo r th e se g lan d s in th e d om estic bull (tra n s v e rs e
ab ou t 30 m m , lon g itu din al a b o u t 10 m m ). T h e re su lts of our o b serv atio n s
on th e e x te n t an d m e a s u re m e n ts of p a rs d issem in a ta p rostatae in th e
E u ro p ean bison co rresp o n d to th e re la tio n s in b u lls d escrib ed by
G r a-
b o w s k i (1937).
T h e E u ro p e a n b ison ’s b u lb o u re th ra l glands a re situ a te d on the d o rsa l
side of th e m ale u re th r a , n e a r th e m edial line, and th u s a re sim ila r
to th o se in sm all d o m estic ru m in a n ts , p a rtic u la rly th e ra m ( G r a b o w ­
s k i, I. c.). In th e m ale g o a t th e se glands a re sh ifted m o re to th e side,
and th is is ev en m o re m ark e d in th e d o m estic bull ( G r a b o w s k i , I. c.).
In o u r m a te ria l th e m ean le n g th of th e g lan d in ad u lt E u ro p e a n bison
w as 21 m m and b re a d th 27 m m . A cco rd in g to N i c k e 1 (1954) th e len g th
of th e b u lb o u re th ra l g lan d in c a ttle v a rie d fro m 40 to 50 m m and
w id th ab ou t 25 m m . M e a su re m e n ts of b re a d th , on a cc o u n t of th e d if­
fe re n c e s in th e situ a tio n an d th e fusion of th e tw o lobes, and in co n ­
seq u en ce in th e w a y in w h ich th e m e a su re m e n t is m ad e (cf. F ig . 1-D ),
can n o t be d ire c tly co m p ared . On th e basis of co m p a ra b le d a ta on le n g th
of th e glan d s, b earin g in m ind th e fa c t t h a t th e b re a d th w e g iv e applies
to b oth lobes, w e can s ta te th a t th e b u lb o u re th ra l g lan d s a re fa r sm a lle r
in th e E u ro p e a n bison th an in c a ttle .
3. U teru s m ascu lin u s
T h e u t e r u s m a s c u lin u s in E u ro p e a n bison w as d escrib ed fo r th e first
tim e b y M ü l l e r (1852). T h e undivided p a r t of th e o rg a n w as »4V2
in ch es lon g« and th e stro n g ly developed horns re a c h e d alm ost to th e
T h e m ale reproductive organs of the European bison
543
testicles. T h e lu m en of th e u te ru s w as co n n ected w ith th e lu m en o f th e
u ro g en ital ca n a l th ro u g h an o stiu m on coZZicuZus sem in a lis.
L e i s e r i n g (1890) also fou nd th e u t e r u s m a s cu lin u s in an E u ro p e a n
bison ca lf. In th e tw o ind ivid u als w h ich he d issected one h ad this o rg a n
h ig h ly d eveloped , w h ile th e o th e r ex h ib ite d re la tio n s sim ila r to c a ttle .
The u t e r u s m a s cu lin u s d escrib ed in E u ro p e a n bison b y K r ö l l i n g
(1930) d iffered fro m th e case d escrib ed b y M ü l l e r (1 852) p rim a rily
as to d im ensions, ab sen ce of horns an d blind c ra n ia l end. K r ö l l i n g
did not find a n y recessi o r p ro cesses w h ich w ould fo rm ev id e n ce of th e
v estig e o f p aired M ü llerian d u cts. D etailed e x a m in a tio n o f th e m i­
cro sco p ic s tr u c tu r e of se ria l sectio n s of th e o rg an , w h ich w a s 105 m m
in len g th , en ab led th e a u th o r to d istin gu ish se v e ra l p a rts . T h e y c o r ­
respond to p a rs ce ru icis u te ri, p a rs u te rin a v a g in a e, v a g in a p ro p ria et
pars v e s tib u la ris v a gin a e. U sin g d a ta tak en fro m stu d ies b y M ü l l e r
(I. c.) a nd L e i s e r i n g (Z. c.), and on th e basis of his ow n in v e stig a tio n ,
K r ö l l i n g ( i . e . ) d iscu sses in g re a te r d etail th e q u estio n o f th e
o ccu rre n c e of u t e r u s m a s cu lin u s in th e E u ro p e a n bison. In co n clu sion
he sta te s th a t »o u t of th e fo u r m ale E u ro p e a n bison so fa r e x a m in e d in
th re e th e re w as d istin ct m ix in g of se x u a l c h a ra c te rs d ifferin g fro m
sim ila r an o m alies in o th e r m a m m a ls. T h e y
d e g en erativ e c h a ra c te rs due to in -b re e d in g « .
m u st
be
co n sid ered
as
O u r o b serv atio n s in cre a se th e n u m b er of E u ro p e a n bison in w h ich th e
u te ru s m a s c u lin u s has been found so f a r to tw e n ty tw o 3), w h ich am o n g
the tw e n ty fo u r e x a m in e d in th is re s p e c t fo rm s 92% of all cases.
A cco rd in g to S c h m a l t z (1 9 1 1 ) and L e n k (1913) u te ru s m a s c u ­
lin u s does n o t o ccu r in d o m e stic ru m in a n ts, o r a t le a st h a s n ot been
found, a lth o u g h
D isselhorst
(1 9 0 4 ) co n sid ers th e g o a t an d sta g as
sp ecies in w h ich th is o rg a n m a y a tta in its g re a te s t d e v e lo p m e n t. S k o ­
d a (1 9 1 7 ), in ch eck in g w h ich of th e v iew s is c o r re c t, found th e p re se n ce
of v e stig e s of M ü llerian d u cts in 71% of th e d o m estic b u lls e x a m in e d ,
all th e b u ffalo ex a m in e d an d in five ra m s. He co n sid ers th e v e s ic u la r
fo rm of th e u t e r u s m a s c u lin u s th e m o st fra q u e n t. In his in tro d u c tio n to
his stu d y S k o d a (Z. c.) p olem ises w ith M ü l l e r (1852) re g a rd in g th e
c h a r a c te r of u t e r u s m a s c u lin u s in th e E u ro p e a n bison, co n sid e rin g th e
case d escrib ed b y M ü l l e r (1 8 5 2 ) a s an a n o m a ly of th e g y n e m a s ty
k ind. S w o b o d a
(1921) co n firm s th e re s u lts of S k o d a ’s stu d ies
(Z. c.) su p p lem en tin g th e m w ith th e e ffe c t of ag e an d c a s tra tio n on th e
o c c u rre n c e ,
ru m in a n ts .
size
and
a p p e a ra n ce
of
u teru s
m a s cu lin u s
in
d o m e stic
3) The presence of u teru s m a sc u lin u s was also noted in »Storper 879«, »Zona 1454«
and »Pudzik«, the organs of which were not included in the m aterial elaborated
in this study.
K . Swiezyrtski
544
It is cle a r
fro m
th e
d a ta
g iv en
ab ove
th a t
th e
o c c u rr e n c e of the
resid u a of M ü llerian d u cts in th e g e n ita l sy ste m of m ale B o v id a e is n ot
a r a r e p h en om en on. T h e E u ro p e a n bison w ou ld th u s n o t fo rm a sp ecial
ex ce p tio n if it w e re n o t a lm o st a ru le and if th e d egree of fo rm a tio n of
th ese o rg an s w h ich in o th e r sp ecies a re ru d im e n ta ry , w e re n ot so
co n sid erab le.
T h e la rg e d im ension s of u te ru s m a s c u lin u s and its co n n e ctio n w ith
ca n alis u ro g e n ita lis re su lte d in K r ö l l i n g (1 9 3 0 ) te rm in g th e case
d escrib ed b y M ü l l e r (185 2 ) as »a ty p ica l h e rm a p h ro d ite « . T h e d ra w ­
ing an d d escrip tio n of th e u te ru s g iven in M ü 11 e r ’s stu d y (1. c.)
sh o w s th a t it ca n be a llo ca te d to the th ird of th e ty p e s w e h a v e
d istin g u ish ed (F ig . 3— C).
W e th u s h a v e g rou n d s fo r sta tin g th a t resid u a of e m b ry o n ic M ü l­
leria n d u cts o c c u rre d , an d o v e r a co n sid erab le d ista n ce , in th e g re a t
m a jo rity of th e m ale E u ro p e a n bison so f a r e x a m in e d . T h e d e g re e of
th e ir co n tin u ed d ev elo p m en t m a y d iffe r up to th e sta g e , w h ich , fro m
th e a sp ect of e x te r n a l sh ap e, is sim ila r in a p p e a ra n ce to th e in te rn a l
g e n ita l o rgan s o f th e fem ale. T h is a d v an ced d e v elo p m en t, w h en w e can
d istin gu ish m a cro sco p ica lly p a rts co rresp o n d in g to th e v a g in a , tw o ­
-h o rn e d u te ru s an d ev en th e o v id u ct, is an e q u a lly fre q u e n t phen om en on
(cf. F ig . 3— A , C ).
W e h a v e no g ro u n d s fo r d e lib e ra tin g on w h e th e r th e E u ro p e a n bison
ca lf d escrib ed b y L e i s e r i n g (1 8 9 0 ) w as fu lly ca p a b le of siring
p ro g en y , b u t K r ö l l i n g
(1 930) co n sid ers th e m ale E u ro p e a n bison
w h ich h e d issected a s h avin g a tta in e d re p ro d u ctiv e fu n ctio n . A ll th e
E u ro p e a n bison w h ich w e e x a m in e d ca m e fro m an im als k ep t in a re s e rv e
and th u s th e ir se x u a l a c tiv ity w as of n e ce ssity lim ited (and in som e
ca se s co m p le te ly elim in ated ), n e v e rth e le ss se v e ra l of th em sired p ro g en y .
A m o n g m ales w ith ty p e A u te ru s th e follow in g th re e bison sire d p ro ­
g e n y : » P o k o rn y « , » P la m ie c« and »P o m ru k « . O ne bison in e a ch grou p
w ith ty p e B and C u te r u s : »P osu sz« and » P u g II«. E u ro p e a n bison w ith
typ e D and E u te ru s did not h a v e a n y p ro gen y. In v iew of th e sim ila rity
in th e s tru c tu re of g en ital o rg a n s in »M ü ller’s E u ro p e a n bison« and
»P u g II« an d th e fe rtility of th e l a tte r w e m a y assu m e th a t »M ü lle r’s
E u ro p e a n bison« w as not a h e rm a p h ro d ite as K r ö l l i n g (1 9 3 0 ) ju d ged
it to be.
T h u s ev en a d v an ced
d ev elo p m en t
in
m ale
E u ro p e a n
bison of th e
d é riv â te s of M ü llerian d u cts did n ot sig n ifica n tly a f f e c t th e ir fe rtility .
In o rd er to obtain a co m p le te p ic tu re of th e co n n ectio n s betw een
u t e r u s m a s cu lin u s an d th e fe rtility of th e m ale E u ro p e a n bison it w ould
ap p e a r essen tial to d ra w a tte n tio n to th e ab sen ce of p ro g e n y in in ­
d iv id u als o f w h ich on e h ad a n only v e r y slig h tly d eveloped u te ru s and
The m ale reproductive organs of the European bison
545
th e secon d had none a t all (F ig . 3— D, E ; T ab le 7). T h is w as co n n e cte d
w ith cau ses of a d iffe re n t n a tu re . » P o ła m a n ie c« d isclo sed p ath o lo g ica l
ch an ges in th e hip jo in t ( P i l a r s k i , 1956) w h ich m ad e it im possible
fo r it to ju m p w hen co v erin g a fem ale. »P ozew « w as tra n s fe rre d to th e
re s e rv e a t S m a rd z e w ice w hen tw o y e a rs old and h ad no c o n ta c t w ith
fem ale bison.
K r ö l l i n g (1930) co n sid ers th e e ffe ct o f in -b re e d in g as th e ca u se
of th e m ix in g of se x u a l c h a ra c te rs . T h is fa c to r u n d o u b ted ly o p e ra te d a t
th e tim e F . M ü l l e r a nd L e i s e r i n g
w ro te th e ir stu d ies and b e ca m e
ev en m o re sig n ifican t a fte r th e first w orld w a r, w h e n th e n u m b e r of
E u ro p ean bison su rv iv in g in th e w h o le w orld did n ot e x c e e d 50 in d ivid u als
(K r y s i a k, 1967). On th e o th e r h and, h o w e v e r, th e p h en om en o n of
o c c u rre n c e in m ales of d e riv a te s of M ü llerian d u cts of a d v an ced d e v e ­
lop m en t w a s found in g re a t n u m b ers o f cases in su ch m a m m a ls a s th e
E u ro p e a n b e a v e r (F r e y e, 1953) and C an ad ian b e a v e r (C o n a v a y ,
1958). It w ou ld be d ifficu lt to a scrib e th is to th e e ffe c t of c irc u m s ta n c e s
sim ila r to in -b reed in g in th e ca se of e ith e r species.
4. Penis
T he p en is of th e E u ro p e a n bison, as in o th e r u n g u la te s, is o f th e
fib ro -e la s tic ty p e . O u r o b serv a tio n s co n firm th e re m a rk s m ad e by
M üller
(1852) th a t th e sigm oid fle x u re of th e p en is is o n ly fa in tly
m ark ed in th e E u ro p e a n bison. T h is applies p a r tic u la rly to th e d istal
c u rv e of th e sigm oid fle x u re . T h e s tr u c tu r e of co rp u s penis in re s p e c t
of m a c ro s co p ic a l s tru c tu re s co rresp on d s to th e re la tio n s d escrib ed b y
Preuss
(1954) in d o m estic ca ttle .
B u lb u s ca n a lis u ro g e n ita lis in th e E u ro p e a n bison is m o re co m p a ct in
s tr u c tu r e th a n w ou ld ap p e a r to be th e case fro m th e d e scrip tio n g iv en
of th e b u lb u s in th e d o m estic b u ll ( G r a b o w s k i , 1 9 3 7 ). In th e
E u ro p e a n bison th e re is no m ed ian g ro o v e on th e d o rsa l su rfa ce of th e
b u lb u s, div idin g it in to tw o c r u r a . On th e c o n tr a ry th e r e is a sh a rp
c r is ta in th is p lace w h ich d orsad p asses into a se p tu m dividing th e
b ila te ra l m m . b u lb o c a v ern o si.
G r a b o w s k i (I. c.)
d escrib es an d
illu s tra te s th is sh a rp ly b o rd e re d sh ap e of th e bulbus, a s ty p ica l o f th e
m ale g o at.
T h e position of th e p a rs p e n is ca n alis u ro g e n ita lis to g e th e r w ith co rp u s
c a v e rn o s u m in re la tio n to co rp u s penis is sim ila r in th e E u ro p e a n bison
to th a t in c a ttle .
A p e x p e n is on th e d o rsal side, and n e a r th e g a lea gla n d is on th e
d o rs o la te ra l side (le ft) h as a dense b u n dle of co n n e ctiv e tissu e fib re s.
In th e d o m e stic bull it w as defined by M ä d e r (1 9 0 7 ) an d S c h m a l t z
K. Sw iezynski
546
(1919) as a co m p o n en t p a rt of tu n ic a penis. B y P r e u s s (1954) it w as
term ed lig a m e n tu m a p ica le p e n is and included in th e com p osition of
tu n ic a a lb u g in e a p e n is . T h e position of this p a rt of th e penis in E u r o ­
p ean bison is sim ilar to th a t in th e d om estic b u ll, e x c e p t t h a t th e
v a ria tio n s d escrib ed in th e l a tte r b y P r e u s (i. c.) w ere n ot found.
T h e e x te r n a l a p p e a ra n ce of a p e x p e n is in the E u ro p e a n bison p e rm its
of includ in g it in G e r h a r d’s p lan (1933) classify in g th e penis of
ru m in a n ts in th e ty p e clo se st to th a t accep ted for th e g en u s Bos.
T ab le 9 sh o w s th a t in E u ro p e a n bison ca lv e s fro m one d a y to te n ­
-m o n th s old th e penis is re la tiv e ly sh o rt. This is m o st c le a rly seen in
th e te n -m o n th s old » P u sty b o r« . In o ld er E u ro p e a n bison th e situ a tio n
ch an g es in fa v o u r of th e penis and is m ain tain ed on a sim ila r level in
all E u ro p e a n bison re g a rd le s s of age. It m a y th e re fo re be co n clu d ed
t h a t w hile th e in crease in b od y m e a su re m e n ts is at first m o re ra p id
th a n th e in crease in len g th of th e penis, a fte r th e ag e of ten m o n th s th e
p en is elo n g ates. Its len g th th en re m a in s in a ce rta in a p p ro x im a te ly
co n sta n t ra tio to th e e x te rn a l b od y m e a su re m e n ts (cf. T ab le 9).
T h is applies b oth
to
th e
ra tio
of
len g th
of penis to d im ension F ,
w h ich in a ce rta in sense co rre sp o n d s to the le n g th of th e ab d om in al
in teg u m en t. T he p ro p o rtio n a te ly sh o rt penis in y ou n g bison m a y th e r e ­
fo re be th e cau se (a p a rt fro m adhesion of the la y e rs of th e p rep u ce) of
th e penis n o t p ro tru d in g fro m th e p rep u ce d u rin g d e fa e ca tio n , w h ich
is a c h a ra c te ris tic of ad u lt bison ( J a c z e w s k i , 1958).
In his stu d ies of th e p o stn a ta l d evelo p m en t of penis and p re p u ce in
d o m estic bu lls A s h d o w n (1960) distin guish ed five su cce ssiv e sta g e s
of th is d evelo p m en t. C om p lete se p a ra tio n of th e la y e rs of th e p rep u ce
an d co m p lete elon g ation of th e penis, acco rd in g to this a u th o r, ta k e s
p la ce in you ng b u lls b etw ee n th e n in th and elev en th m on th of life. Th e
ap p e a ra n ce of a p e x penis in te n -m o n th s old » P u sty b o r« (P h o to 18)
co rresp o n d s to th e sta g e d escrib ed fo r fou r to six -m o n th s old d om estic
y ou n g bulls. T h is m a y fo rm e v id en ce of th e slo w e r d e v e lo p m e n t of th e
E u ro p e a n bison in th is re s p e c t also.
In o u r m a te ria l th e m e a n le n g th of th e penis in ad u lt E u ro p e a n bison
w a s 86.4 cm . In d om estic c a ttle th is dim ension v a rie s fro m 90 to 100
c m ( S c h u m m e r & N i c k e l , 1960). B e a rin g in m ind th e e ffe ct of
fix in g it m ay be said th a t th e le n g th of penis in E u ro p e a n bison co m es
w ith in th e lim its accep ted fo r ca ttle .
5. The p rep utial an d e x tre n sic penile m u scles
T h e p rep u tial m u scles co rresp o n d in re s p e c t of b oth in sertio n s and
co u rse
to
th e
re s p e c tiv e
m u scles d escribed b y
Brauell
(1868) in
The m ale reproductive organs of the European bison
d om estic ru m in an ts. Th e m a rk e d
547
b ro ad en in g and se m ic irc u la r co u rse
o f m m . p re p u tia e cra n ia le s sim ila r to th a t w h ich w e d e scrib e d in E u r o ­
pean bison w as found b y B r a u e 11 ( i . e .) o n ly in th e c a s e o f th e ra m .
M . b u lb o c a v e rn o s u s is in p rin cip le sim ila r in s tr u c tu re to th a t in
d om estic ru m in a n ts. It is s h o rte r th a n in th e d om estic b u ll, in w h ich
it a tta in s len g th s up to 18 cm ( H e i n e m a n n , 1 9 3 7 ), b u t is b ro a d e r
and th ick e r. T h is is u n d o u b ted ly co n n ected w ith th e d im en sion s an d
s tru c tu re of th e b u lb u s u re th ra lis , p re v io u sly m en tio n ed .
T he m m . isch io c a v ern o s i, stro n g ly d eveloped in th e E u ro p e a n bison,
su rro u n d
m. b u lb o c a v e rn o s u s
fro m
th e
sides,
fo rm in g
tw o
oblique
g ro o v es ru n n in g len g th w a y s. T h e b ila te ra l m m . r e t r a c t o r e s p e n is a re
situ ated in th ese g ro o v es (P h o to 20). T h e y a re se p a ra te d fro m e a ch
o th e r at th is lev el. T h is a rra n g e m e n t, su ch as d escrib ed ab ov e in th e
E u ro p e a n bison o f th e v a rio u s m u scles of th is region in re la tio n to e a ch
o th e r, w as d escrib ed in th e ca se of d om estic ru m in a n ts b y H e i n e ­
m a n n (1 9 3 7 ) in th e ca se of th e m a le g o at. A cco rd in g to H e i n e m a n n (i. c.) in th e m ale g o a t th e b ila te ra l m m . r e t r a c t o r e s p e n is re ta in
th e ir in d ep en d en ce in th e ir fu rth e r co u rse also, w hile in th e E u ro p e a n
bison b elo w ra d ix p e n is th ese m u scles b eh ave as th e y do in d o m e stic
bu lls.
REFEREN CES
1. A c k e r k n e c h t E., 1943: Das Eingew eidesystem . [In: E llen b erg er-B au m Hand­
buch der vergl. A n at. der H au stiere]: 522—544. Springer V erlag, B erlin .
2. A s h d o w n R ., 1960: D evelopm ent o f penis and sheath in the bull ca lf. J .
A gricu lt. Science, 54, 3: 340— 352. Cam bridge Univ. Press.
3. B o j a n u s L . H , 1827: D e u ro n o s tr a te e iu s q u e s c e le to . Nov. A cta A kad. L eo­
pold. Carol., 13: 411— 478. Bonnae.
4. B r a u e l l H., 1868: Myologie der m ännlichen G enitalien. V jsch r. wiss. V eterinärkd ., 29: 1—27. Wien.
5. B u c h h o l t z O., 1951: Zum B au des Peniskörpers beim P ferd e und seine
funktionelle Bedeutung. D iss. Hannover.
6. C o n a v a y C., 1958: The uterus Tnasculinus of C a sto r c a n a d en s is. J . M am m ,
39: 97— 108.
7. D i s s e l h o r s t R ., 1904: A u sführapparat und Anhangsdrüsen der m ännlichen
G eschlechtsorgane. [In: »Oppels Leh rbuch der m icroskop. A nat. W ierb eltieren«], 4 (acc. to W a l t e n , 1965).
8. E c k s t e i n P. & Z u c k e r m a n n J ., 1956: Morphology of the reproductive
tra ct. [In: »M arsh all’s physiology of reproduction«], 1, 1: 43— 174. Longm ans
G reen and Corp. London—N. Y ork,
9. F r e y e H., 1953: Bem erkungen zum G enitalsystem des B ib ers. W iss. Z. M ar­
tin L u th er U niversität., 2: 911—915.
10. G e r h a r d t U., 1933: K loake und Begattungsorgane. [In: »Handbuch der
vergl. Anat. der W ierbeltiere«], 6: 314— 347. U rban & Schw arzeberg. B erlin —
W ien.
548
K . Sw ieżyński
11. G r a b o w s k i K., 1937: Uber Sehw ellräum e der H arnröhre der m ännlichen
H aussäugetiere. Diss., H annover.
12. H e i n e m a n n K ., 1937: Einigen M uskeln des m ännlichen G eschlechtsappara­
tes der H aussäugetiere — M. buibocauernosus, m. is ch io c a v ern o su s , m . r e tr a c to r
p en is. D iss. Hannover.
13. J a c z e w s k i Z., 1958: Reproduction of the European bison, B ison b o n a su s
(L.), in reserves. A cta th eriol., 1, 9: 333— 376.
14. K r ö l l i n g O., 1930: Uber U teru s m a scu lin u s, sowie einige A rtm erkm ale im
m ännlichen G enitalapp arat des europäischen W isent (Bison b o n a su s L.).
Ztschr. f. m ikroskop-anat. Forsch., 20, 3/4: 557—583. Leipzig.
15. K r ö l l i n g O. & G r a u H., 1960: Lehrbuch der Histologie und vergleichen­
den m ikroskopischen A natom ie der H austiere: 315—354, P arey , B erlin .
16. K r y s i a k K ., 1967: T h e history of the European bison in the Białow ieża
forest and the results of its protection. Acta theriol. 12, 19: 323— 331.
17. L e i s e r i n g A. G. T., 1890: Eingew eidelehre [In: »G urlts Handb. der vergl.
A natom ie der H aussäugetiere«], 7 A ufl. (acc. to K r ö l l i n g , 1930).
18. L e n k H. J . ( 1913: Zur A natom ie und Histologie der H arnblase und P ars pelv in a der H arnröhre der H aussäugetiere. Diss. Dresden.
19. M a r t i n P. & S c h a u d e r W., 1938: Anatomie der H austiere. 3: 280— 301.
S chick hard t und Ebner Verl., S tu ttg art.
20. M ä d e r E., 1907: Zur A natom ie der Giarts p en is der H austiere. A rchiv f.
wiss. u. prakt. Tierh eilkd e, 33: 137— 188; 281— 311. V erlag Hirschwald, B erlin .
21. M ü l l e r F., 1852: B em erkungen über den Auerochsen... V jschr. wiss. V eterinärkde, 2: 110— 133. B erlin .
22. N i c k e l R ., 1954: Zur Topographie der akcessorischen Geschlechtsdrüsen bei
Schw ein, Rind und P ferd . T ierärztl. Um schau, 9, 21/22: 1— 12. Konstanz a. B.
23. O w e n R ., 1848: Notes on the anatom y of the m ale Auerochs (Bison eu r o p eu s). P roc. zool. Soc. Lond., 16: 126— 133.
24. P i ę k o ś M. , P i l a r s k i W. & R o s k o s z T., 1958: O bserw acje nad długością
je lita u żubra (Bison bonasus L.). Folia morphol., 9, 1: 69—79. W arszawa.
25. P i l a r s k i W., 1956: D eform acje szkieletu obserwowane na sześciu okazach
żubra, B is o n b o n a su s (L.). Folia morphol., 7: 301—306. W arszawa.
26. P i l a r s k i W. , S e r w a t k a S., S w i e ż y ń s k i K. & W ę g r z y n M., 1967:
New attem ps at fixin g anatom ical m aterial of large m am m als. A cta theriol.,
12, 31: 453—458.
27. P i l a r s k i W., 1967: Układ moczowy żubra, B iso n b o n asu s (L i n n a e u s
1758), D iss. W arszawa.
28. P r e u s s F., 1954: Die T u n ica a lb u g in e a p en is und ihre T rab ek el bei Pferd
und Rind. A nat. Anz., 101: 64— 83. Jen a.
29. R o s k o s z T. & E m p e l W., 1963: Das S k elett der G liedm assen des W isents,
B iso n b o n a su s ( L i n n a e u s , 1758). A cta theriol., 7, 13: 259— 300.
30. S c h l u m p e r g e r O. R., 1954; Der Nebenhoden und seine Lage zum Hoden.
Diss., Hannover.
31. S c h m a l t z
R., 1909: P enis. B erlin er tierärtzl. W schr., 25: 455— 458.
32. S c h m a l t z R., 1912: Die G eschlechtsorgane. [In: »E llenberger — Hdb.
vergl. m ikroskop. Anatom ie der Haustiere«], Zweite A ufl., B erlin , (acc. to
S k o d a , 1917).
33. S c h u b e r t F , 1909: B eitra g e zur Anatom ie des Herzens der H aussäugetiere.
Diss., D resden.
Układ płciowy samczy żubra
549
34. S c h u m m e r A. & N i c k e l E , I960: Lehrbuch der A natom ie der H austiere,
2: 338— 345. P . P arey , B erlin — Hamburg.
35. S c h w a r z e E., 1939: Hoden und Nebenhoden. T ierärztl. W schr., 47, 19: 391—
298. W ien.
36. S k o d a K., 1917: U ntersuchungen über das Vorkom m en Uterus m asculinus
bei einigen W iederkäuern (Bas tau ru s, B a s bubeius und O vis aries). Anat.
A n z , SO: 111— 120. Je n a .
37. S w o b o d a A., 1929: B eitra g zur K enntnis des U tricu lu s m a scu lin u s der
H austiere. Ztschr. Anat. E ntw icklugs-gesch., 89: 494— 512.
38. V 1 o t e n J . G. S., 1927: Entw icklungs des Hodens und das U rogenitalverbindungs des Rindes. Diss., U trecht (acc. to M a r t i n & S c h a u d e r , 1938).
39. W a 1 1 o n A., 1965: T he m ale genital tract. [In: »M arshall's physiol, of repro­
duction«] 1, 2: 133— 142. Longm ann G reen Co., London.
40. W e n d t K. , P o h l J . & M r o s k H., 1960: K riptochism us des Schafes und
seine w irtsch aftlich e Bedeutung. A rch. Tierzucht., 3, 5: 440—458. B erlin .
41. Z e d e n o v V. H., 1965: A natom ia domaänih zivotnih., 2: 311— 360. Izd. Vyzäaja
Skola. Moskva.
R eceived, J u n e 25, 7968.
W arsaw A gricu ltu ral U niversity,
D epartm ent of A nim al Anatom y,
W arszawa, G rochow ska 272.
Krzysztof Ś W IE Ż Y Ń S K I
U K ŁA D P Ł C IO W Y SA M C Z Y Ż U BR A
Streszczenie
P raca dotyczy badań nad anatom ią narządów układu płciowego samczego żu­
bra — B ison bonasus L . w oparciu o preparacie 20 zwłok osobników różnego w ie­
ku (Tabela 1). O kreślono w ym iary i ciężar jąd er (Tabela 2) oraz ich kształt i po­
łożenie (Fot. 2 do 4). Ustalono odpowiednie dane odnośnie najądrzy, nasieniow odów, gruczołów płciow ych dodatkowych, cew ki moczowopłciowej i prącia (Tabele
3 do 6 i 8 do 10 oraz F o t. 14 do 18). Stw ierdzono występow anie u samców żubra
u ierus m a scu lin u s (T ab. 7, Ryc. 3 oraz Fot. 5 do 13). B ra k widocznej zależności
między stopniem rozwoju te j ostatniej a w ykształceniem któregokolw iek z pozo­
stałych narządów układu płciowego oraz zdolnością rozrodczą sam ców om awiane­
go gatunku. Opisano budowę moszny, napletka i m ięśni topograficznie i czynnoś­
ciowo związanych z układem rozrodczym (Fot. 1, 19—21).
W porów naniu z bydłem domowym ją d ra żubrów dorosłych są bardziej kuliste
a ich średni ciężar je st niższy. N ajądrza m ają silnie w yrażoną, szeroką głowę
i zgrubiały odsądzony ku tyłow i ogon. Wśród gruczołów płciow ych dodatkowych
gruczoły opuszkow o-cewkowe różnią się położeniem i są m niejsze od takow ych
u bydła. Moszna żubra usytuow ana je s t bard ziej tylnie, je s t stosunkowo krótka
o słabo zaznaczonej szyjce. Lew a część moszny wraz z jąd rem w ysunięta je s t le k ­
ko ku przodowi. Zagięcie esow ate prącia je s t sła b iej zaznaczone.
550
K . Swiezyrtski
E X P L A N A T IO N OF P L A T E S X V —X I X
P la te X V .
Photo 1. Position of the scrotum in the European bison (»Pom ruk«, about 9 years
old). View from le ft side.
a — te stis sin istr.; b — testis d ex tr.; c — p a p illa e u b er is.
Photo 2. C ross-sections of scrotum and testicles of the European bison (»Pomruk«,
about 9 years old).
A — at level of neck: B — m id -length: C — near a p e x scro ti, a — m . c r e m a s te r
ex t,; b — ca p u t e p id id y m id is ; c — d u ctu s d e fe r e n s ; d — sep tu m s cr o ti; e ■
—■ m e d ia stin u m te s tis ; f — corpus e p id id y m id is ; g — cauda ep id id y m id is.
Photo 3. Longitudinal section of le ft testis of European bison (»Pug II« , about
6 years old).
a — c a p u t e p id id y m id is ; b — ca u d a e p id id y m id is ; c — m ed ia stin u m testis.
Photo 4. C ross-section of cauda e p id id y m id is of the European bison (»Pustelnik«,
about 8 years old).
P la te X V I.
Photo 5. U rogenital organs of the European bison (»Puzon«, about 5 years old).
P ars p elv in a of the system seen from ven tral side: bladder deflected, a — testis;
b — pars vascu laris fu nicu li sperm atiei; c — d u ctu s d e fe r e n s ; d — »band« con­
sisting of the restidua of the cranial part of M ullerian duct; e — co rn u ; f — p a rs
in d iv isa u te ri m asculini; g — a m p u lla d u ctu s d e fe r e n tis ; f — gl. v es ic u la ris :
i — p lic a u ro g e n ita lis (excised); j — v e s ic a u rin a ria ; k — pars p elv in a u r e th r a e ;
1 — cru s p e n is ; m — tru n cu s p en is (artificially bent in order to include the whole
in th e photograph).
Photo 6. P ars p e lv in a of the urogenital system of the European bison in situ
(»Pokorny«, about 9 years old). View from dorsal side.
A — os iliu m ; B — sp in a is c h ta d tc a : C — iu b er ischiadicum ; a — pars tndivisa
u teri m a s c u lin i; b — a m p u lla d u ctu s d e fe r e n t is d e x t r a ; c — gl. v es ic u la ris sin istra ;
d — co rp u s p r o s ta ta e ; e — pars p elv in a u r e th r a e ; f — m u s cle s o f th e b u lb o ­
u r e th r a l g la n d s; g — m . is c h io u r e th r a lis ; h — m . is c h io c a v ern o su s ; i — m. bulbocavernosus.
Photo 7. Isolated u teru s m a scu lin u s of the European bison (»Pomruk«)
together w ith neighbouring organs.
View from : A — dorsal side; B — ventral side; a — »band* — residua of cran ial
part of M ullerian duct; b — corn u u te r i sin istr.; c, d, e — cranial, median and
caudal parts of pars in d iv is a u teri; f — d u ctu s d e fe r e n s d ex tr.; g — gl. v e s ic u la ris d e x t r .; h — u r e th e r d e x t r .; i — vesica u rin a ria.
P late X V II,
Photo 8. Cross-s( ;tion of intern al lay er of corn u u te ri m a s cu lin i of the European
bison. Magn. approx. 70 X .
Photo 9. C ross-section of mucous m em brane gland of u teru s m u scu ltn u s of the
European bison. Magn. approx. 140 X .
Photo 10. C ross-section through internal layers of median part of u teru s m a s c u ­
lin u s o f the European bison. Magn. approx, 70X ,
The m ale reproductive organs of the European bison
551
Photo 11. Epithelium of the part of u teru s m a scu lin u s shown in Photo 10 — magn.
approx. 140 X .
Photo 12. Cross-section through intern al layers of term inal p art of u teru s m a s c u lin u s of the European bison. Magn. approx. 40 X ,
Photo 13. Epithelium and true mucous m em brane of the part shown in photo 12.
Magn. approx. 100X ,
P la te X V III.
Photo 14. Cranial part of dorsal w all of pars p e lv in a c a n a lis u ro g e n ita lis of the
European bison (»Połam aniec«, about 9 years old).
Upper pins indicate site of o stia d u ctu u m d e fe r e n tiu m : low er ostium u te r i m a scu lin i. a — colliculus se m in a lis ; b — the opening of excreto ry ducts of prostate;
c — u re th ra p r o p r ia ; d — p a rs d is ta lis c r is ta e u re th r a lis .
Photo 15. U rogenital organs in the European bison (»Pluszcz«, about 10 years old)
region of arcu s is c h ia d icu s m . b u lb o c a v e m o s u s rem oved, also c ru ra p en is together
w ith m . is c h io c a v e r n o s u s . View from right side.
a — term inal part of pars p elv in a u r e t h r a e ; b — g l. bulbo u r e th r a le s ; c — dorsal
and la tera l cristae of bulbus u rethralis; e — r a d ix p en is.
Photo 16. Cross-sections of pars p e lv in a canalis urogenitalis in the European bison
(»Pomruk«, about 14 years old). D istan ce betw een sections about 1 cm.
a — excretory ducts of the vesicu lar glans; b — corpus p r o s ta ta e ; c — colliculus
sem inalis; d — pars d is sem in a ta p r o s ta ta e ; e — c o r p u s c a v e r n o s u m u r e th r a e ; f —
m . u re th ra lis.
P la te X IX .
Photo 17. Cross-sections: A — of c ru ra and r a d ix ; B — truncus; C — a p e x p en is
in the European bison (»Pom ruk«, about 14 years old). D istance betw een sections
— from 1 to 7 about 1 cm, 8—9 about 12 cm, 11 to 13 — about 6 cm, 14 to 17
about 2 cm, 18 and 19 about 0.5 cm .
a — crus penis; b — co rp u s c a v e r n o s u m p en is; c — pars penis u r e th r a e su r­
rounded by corpus c a v e r n o s u m ; d, e — extern al and internal layer of tu n ica a l­
b u g in e a ; f — trabeculae co rp o ru m c a v e r n o s u m p en is form a »centrum «; g — lig.
a p tc a le p en is.
Photo 18. A p e x p en is in a 10-m onth old European bison (»Pustybór«). Prepuce
p a rtia lly pushed back.
a — g a le a grlandis; b — the connections betw een preputial layers.
Photo 19. A p e x p en is of the European bison (»Pom ruk«, about 14 years old).
Prepuce transected along ven tral w all.
A — r a p h e p r a e p u tii; e — o rific iu m u r e th r a e e x tern u m .
P hoto 20. The extren sic penile m uscles in the European bison (»Pokorny«, about
15 years old).
V iew from posterior: a — m . r e tr a c t o r p en is; b — m . is c h io c a v e r n o s u s ; c — m.
b u lb o c a v ern o s u s.
Photo 21. Isolated penis of the European bison (»Pokorny«, about 14 years old).
a — pars p e lv in a u r e th r a e ;
b — m. ischiocauernosus;
d — ap ex p en is.
c — m . r e tr a c t o r p en is;
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ACTA TH ERIO LO G ICA Vol. X I I I , 32.
K . Sw iezyhski
P la te XV
M. W^grzyn phot.
ACTA TH ER IO LO G IC A Vol. X I I I , 32,
K. Swiezyiiski
Plate XVI.
S. P ytel (7) &. M. W^grzyn phot.
ACTA TH ERIO LO G ICA Vol. X I I I, 32.
K. Sw iezyhski
P la te X V II.
S. P ytel (9) & M. W^grzyn phot.
ACTA THERIOLOG1CA Vol. X I I I , 32.
K. S wi e ż y ńs k j
Platę XVIII
M. W ęgrzyn phcot.