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A C T A T H E R IO L O G IC A
V ol. 21, 4: 37— 100, 1976
BISO N IAN A.
LXI
Joints and Ligaments of Hind-Limbs of the European Bison
in Its Postnatal Development
Franciszek K O B R Y Ń C ZU K
K ob ryń czu k F., 1976: Joints and ligam ents o f h in d -lim b s o f the E uropean
bison in its postnatal developm ent. A cta theriol., 21, 4: 37— 100.
[With 16 T ables & 32 Figs.]
Using m aterial o f the h ind-lim bs o f 119 European bison (B ison bonasus
L i n n a e u s , 1758) (74 males, 45 fem ales) the author studied the join ts
in course o f postnatal developm ent. W hen the opportunity occu rred, the
observations w ere also m ade for com parative purposes on oth er species
o f B ovin ae. The capacity o f articular cavities and linear m easurem ents
o f ligam ents, articular cartilages, m enisci and articular su rfaces w ere
m easured. T h e angles o f the curvature o f the articular su rfaces and
the range o f m ovem en ts for flex ion and extension o f join ts (ex cept
sacroiliac and hip joints) w ere also estimated. The results con cern in g
the dim ension o f articular surfaces and m enisci and the capacity o f
the acetabulum w ere analysed statistically fo r groups, consistin g o f m ore
than 4 specim ens.
[Inst. A n im al P hysiol., A gric. A cad., 03-849 W arszaw a, G roch ow ska
272, Poland]
.............................................................................................................................37
I. Introduction
II. M aterial and m e t h o d ...............................................................................
.
. #.
38
III. O wn o b s e r v a t i o n s
*.
42
1. Joints o f the p elv ic g i r d l e ..............................................................................................42
1.1. P elvic s y m p h y s i s ...................................................................................................... 42
1.2. S a croiliac j o i n t ............................................................................................................. 43
1.3. O btu rator m e m b r a n e ...............................................................................................45
1.4. Other joints o f the pelvic g i r d l e ...................................................................45
2. H ip j o i n t ..................................................................................................................................45
3. S tifle j o i n t ...........................................................................................................................52
3.1. F em orotibial j o i n t ...................................................................................................... 52
3.2. F em oropatellar j o i n t ............................................................................................... 65
4. Fibrous band o f l e g ............................................................................................................ 68
5. H ock j o i n t ...........................................................................................................................69
6. M etatarsophalangeal j o i n t s .......................................................................................81
7. P roxim al interphalangeal j o i n t s ................................................................................87
8. Distal interph alageal j o i n t s ..............................................................................................88
9. Joints o f a ccessory digits II and V
...........................................................................89
IV. D i s c u s s i o n ................................................................................................................................... 90
R e f e r e n c e s ................................................................................................................................... 98
S t r e s z c z e n i e ................................................................................................................................. 100
I. IN TR OD U C TION
From among the studies on the structure of joints in the European
bison K r y s i a k ’ s (1951) paper concerning the m orphology of the nuchal
ligament should be mentioned here. E m p e 1 (1962) published a study
I
[371
F. Kobryńczuk
38
on the suturae betw een the bones of the skull of this animal and W ęg r z y n & S e r w a t k a (1961) their detailed observations on the sacrosciatic ligam ent in the European bison and dom estic cattle. R a d o m s k i
(1972) investigated the structure of the joints and ligaments of the fo re ­
lim bs o f the bison in its postnatal developm ent on rich materials. Taking
the opportunity o f their osteological studies, W r ó b l e w s k i (1927),
K o c h (1932), P o l e i n e r (1932), J a n i c k i (1938), J u ś k o
(1953),
E m p e l & R o s k o s z (1963), S o k o l o v (1972) and K o b r y ń (1973)
made detached observation on the articulations of bones. In discussing
the volum e of the autopodial bones in the European bison, K o b r y ń ­
c z u k & K o b r y ń (1973) provided remarks useful for the subject con­
cerned and S w i e ż y ń s k i (1962) gave a short m orphological description
of the m iddle interosseous muscle of this animal. P i l a r s k i & R o s ­
k o s z (1957) studied the problem of the ankylosis of the lum bosacral
joint in the process of sacralization o f the last lumbar vertebra in the
European bison and R a d o m s k i & K o b r y ń (1969) described a case
of the dislocation o f the shoulder joint. A case of the occurrence of a
pseudoarthrosis in the cervical vertebrae has been reported by K o b r y ń
& K o b r y ń c z u k (1973).
The lack of a system atic study o f the joints in the hind-lim bs of the
European bison in the course of postnatal developm ent has induced the
author to take up this problem.
—
II. M A TE R IA L AN D M ETH OD
Th e present study w as carried out on the h in d -lim bs o f 71 E uropean bisons,
47 m ales and 24 fem ales. Further observations o f articular surfaces w ere m ade
on the skeletons o f 48 E uropean bisons (27 m ales and 21 fem ales) belongin g to the
collection o f the E uropean Bison A natom ical R esearch Centre, A gricu ltural A ca d e ­
my, in W arsaw . T h e carcasses o f animals w ere obtained from breedin g stations
and zoolog ica l gardens o f this country. The age o f specim ens was established on
the basis o f the E uropean Bison Pedigree B ook ( Ż a b i ń s k i ,
1947— 1965; Ż a b i ń ­
s k i & R a c z y ń s k i , 1972). The division o f the m aterial into age groups, sepa­
rately fo r either sex, is given in Table 1.
W h en the opportu n ity occurred, detached observations w ere also m ade fo r com ­
parative pu rposes on oth er species o f the B ovin ae, like Bison bison L., Bison b o ­
nasus— B ison bison hybrids, Bison bonasus— dom estic cattle hybrids, dom estic cattle,
W atussi cattle, b u ffa loes, zebus and yaks. This m aterial is presented in T able 1,
the age o f the E uropean bison— dom estic cattle h ybrids being given after P v t e l
& K r a s i ń s k a (1971). The dom estic cattle and other species derived from zoolo­
gical gardens w ere considered to be adult on the basis o f the state o f ossification
o f the epiphyseal cartilages. Measurements o f the capacity of articular cavities
w ere m ade on fresh carcasses by injecting them w ith w arm gelatin colloid by
K o s t y r a ’ s (1962) m ethod. This made it possible to get to k n ow the shape o f the
articu lar cavities and th eir com m unications, if th ere w ere any, w ith the n eigh bou r­
ing sy n ovia l bursae. L inear measurem ents (to an accuracy o f 0.1 or 1.0 mm) o f
ligam ents, articu lar cartilages, menisci, articular surfaces, etc. w ere m ade on dissect­
Joints of hind-limbs of the European bison
39
ed carcasses, fix e d by the m ethod o f P i 1 a r s k i e t al. (1967). Fresh preparations
o f the lim bs, separated from the w h ole o f bod y w ere fix e d in a 4 % solution o f
form alin . T h e volum e o f the m enisci (in cu. cm ) w as m easured by im m ersin g them
in a liqu id in a graduated m easuring-glass. The capacity o f the acetabu lu m o f the
hip bon e w as determ ined by fillin g it w ith shot, after its rim had been evened
T ab le 1
M aterial.
n — in all exam inations; nj — on ly in exam ination s o f articu lar su r­
fa ces; n2 — in all exam inations o f articular surfaces, 1 — fo r sp eci­
m ens used in all exam inations, 2 — for specim ens used in exam ination s
o f articu lar surfaces.
E uropean bison
G rou p
A ge range
n
ni
0
I
II
III
IV
V
0— 7 days
1— 12 m onth
1— 4 years
4— 8 years
8— 12 years
over 12 year
8
5
11
8
8
7
0
I
II
III
IV
V
0— 8 days
1— 12 m onth
1— 4 years
4— 8 years
8— 12 years
over 12 year
Total
4
1
6
1
3
9
71
n2
A v g. age,
1
years
2
8
5
16
17
15
13
0.00
0.33
2.11
6.01
10.27
14.60
0.00
0.38
2.34
5.77
9.91
15.85
4
1
9
6
7
18
119
0.00
0.42
1.92
4.57
8.65
18.45
0.00
0.42
2.24
6.14
9.82
17.20
M ales
5
9
7
6
Fem ales
3
5
4
9
48
Other B ovin ae
Species
Bison bison L.
B. bonasus X B. bison
B. bonasus X cattle
Sex
n
F
M
M
4
1
Fest
Felon
F ellach
Festyn
Fey
Fez
Farad
F elly
Fela
•
Fam a
-
TT
D om estic cattle
W atussi cattle
Bubalus bubalus L.
Z ebu
Y ak
<
M
F
M
F
M
F
F
Nam e
2
10
1
1
1
1
1
Age
adult
adult
0.5 years
0.5 years
2 years
2.5 years
4.1 years
4.5 years
4.5 years
1.3 years
2.6 years
9.0 years
adult
adult
adult
adult
adult
adult
adult
40
F. Kobrynczuk
w ith plasticine. F or m easuring its thickness, the articular cartilage was rem oved
by m aking a cut w ith a scalpel along a chord o f the articular surface.
A u x ilia ry angles o f the curvatu re o f articular surfaces w ere m easured in sagittal
planes on carcasses and digested skeletons from m useum collections. A lth ough the
shape o f articular surfaces corresponds on ly in approxim ation w ith geom etrical
figures, it proves h elpfu l to represent them for com parison as regular solids. This
p roced u re w as recom m en ded and adopted, am ong other w orkers, by D u e r s t (1926),
Poplewski
(1927), V o k k e n et al. (1961) and K o s t y r a
(1962). H ere, such
studies h ave been carried out only for joints w ith articular surfaces described
m ore or less on the same radius. In the European bison these articular su rface
in clude the trochlea and condyles o f the fem ur and the surfaces of the tarsocrural,
p rox im a l intertarsal and m etatarsophalangeal joints.
In ord er to determ ine the au xiliary angle (a) o f the sagittal curvatu re o f an
articu lar surface, the length o f the chord a (BC) was m easured (Fig. 1 — an
ex am p le o f the calculation o f this curvature in the m etatarsophalangeal joint),
then a p oin t A w as chosen so that it w as approxim ately the highest point o f the
arc in relation to its chord BC, and the lines A B and A C w ere m easured (au xilia ry
m easurem ents). T he use o f trigonom etrical form u lae for the triangle A B C w ith
sides a, b and c m ade it possible to w ork out the value o f the au xiliary angle /?i.
Then, it w as easy to fix the angles ai and a. The follow in g w orkin g form u la is
used to calculate the angle
cos a/2 = (a2—b2—c2)/± 2 b c
The valu e obtain ed fo r the angle a should be m arked in the appropriate quarter
o f the rectan gle o f coordinates.
Then, the angle a being kn ow n , the radius o f curvature (r) w as calculated from
the equation
r=a /(2sin a /2 )
The angle /3 o f a con cav e articular surface (Fig. 1 sh ow s as an exam ple the
p rox im a l articu lar su rface o f the proxim al phalanx) w as obtained from the equation
sin (3/2 = l/a(sina/2)
w here 1 is the chord (DE) o f the concave articular surface, the other sym bols being
the sam e as p reviou sly.
^
The range o f m ovem en ts fo r the flex ion and extension o f the tarsocrural, p r o x i­
m al intertarsal and m etatarsophalangeal joints w as calculated by subtracting the
valu es o f the angles o f the ad join in g articular surfaces from each other:
(p = a —(3
q>={3—a
w hen a > (3
w hen a < /?
The ridges o f the trochlea o f the fem u r lie in the planes w h ich form a dihedral
angle a b elow the knee. This angle w as calculated from the form u la
tga/2 = (b —c)/2a
w here a is the length o f the chord o f the lateral ridge o f the trochlea, and b and c
are the p rox im a l and distal w idths o f the trochlea.
The troch lea o f the fem u r is inclined to a side, form in g an angle a w ith the
plane pa ra llel to the lon g axis o f the fem ur and passing through the backm ost
points o f the con dyles o f this bone. The equation used to calculate this angle w as
t g a = c /(a —b)
The lengths o f the lines a, b and c w ere obtained w ith the help o f a special table,
designed b y E m p e l & R o s k o s z (1963) and illustrated diagram m atically in Fig. 7.
T he angles o f inclination o f such elem ents as the head and condyles o " the fem ur
*
Joints of hind-limbs of the European bison
41
and the trochleae o f M t 1 I I + I V in relation to the lon g axis o f the bone w ere
determ ined trig on om etrically as angles o f suitably chosen triangles.
The angles form ed in the sagittal plan by the long bon es m eetin g in the joints
w ere determ ined in a sim ilar w ay. A b ou t 100 ph otographs o f E uropean bisons in
side view w ere used fo r this purpose. The prom in en ces o f the skeleton and central
points o f the join ts m anifesting them selves th rough the skin served as m easuring
points. T h e c o x o fe m o r a l, fem orotibia l and tibiom etatarsal angles w e re m easured
b y this m ethod. The accuracy o f m easurem ents is given in decim als o f a degree.
The results con cern in g all the dim ensions o f articular su rfaces and m en isci and
the capacity o f the acetabulum w ere analysed statistically (s and v) for groups
Fig. 1. D eterm in ation o f the angle o f con vex and concave curvatures o f an articular
su rface exem p lified by the m etatarsophalangeal joint,
cat — angle o f cu rva tu re o f the trochlea o f M tI I I + I V ,
— angle o f curvatu re o f
th e proxim al articu lar su rface o f the p roxim al phalanx, r — radius o f curvature
o f trochlea and articu lar surface o f the p roxim a l phalanx, ax, /h — central angle
an d angle in scribed in a circle w ith the centre 0, a (BC ) — chord o f trochlea,
I (DE) — chord o f articular surface o f proxim al phalanx, I — bones o f M tI I I + I V ,
II — proxim al phalanx.
consisting o f m ore than 4 specim ens. This proced u re w as not applied fo r the d i­
m ensions o f ligam ents, since the data obtained, under d ifferen t condition s o f p re ­
paration , fo r these structures, readily undergoing decay, w ere approxim ate.
The grow th co e fficie n t is, according do D a v l e t o v a
(1960), the quotient o f
m easu rem ents in groups V and 0. Bisons belongin g to grou ps III, IV and V
a r e regarded as adult specim ens. The Latin n om en clatu re is based on the
N om ina A n atóm ica V eterin aria (1973), on tex t-b ook s by P o p l e w s k i
(1948), E 11e n b e r g e r & B a u m (1943), M a r t i n & S c h a u d e r (1938) and papers by
N i c k e l & g a n g e r (1953) and K o s t y r a (1962).
42
F. Kobrynczuk
III. O W N O B SE R V A TIO N S
1. Joints of the Pelvic Girdle
1.1. P e lv ic Sym physis
The fibrocartilage of the pelvic sym physis form s tw o centres, of w hich
the anterior (Fig. 2— 1) unites the pubic bones and the posterior
(Fig. 2— 2) the ischia of the opposite sides. The tw o centres are con­
nected by a narrow isthmus of the same tissue. In the youngest E uro-
Fig. 2. H orizontal section through p elv ic sym physis (16-m on th -old male).
1, 2 — anterior and posterior cartilaginous centres, 3 — ischiatic arch, 4 — edge
o f the obturator foram en.
pean bisons the sym physeal fibrocartilage passes into the epiphyseal
cartilage of the ischial tuberosity and form s the cartilaginous bordering
o f the ischiatic arch (Fig. 2— 3). A small number of fibres run trans­
versely on the dorsal surface o f the symphysis, their nuipber being
Joints of hind-limbs of the European bison
43
greater on the ventral side; they m eet in the m idline to form a high
fibrous crest. In the anterior portion of the sym physis these fibres
extend obliqu ely cephalad and in the posterior portion obliquely caudad.
The pelvic sym physis is strengthened anteriorly b y the anterior pubic
ligam ent (Fig. 3— 1). This ligam ent is in the shape of an isosceles
triangle, the base of w hich extends betw een tw o iliopectineal eminences
and the vertex points to the front. This vertex is the point of attachment
for the straight abdom inal m uscle ( S w i e z y ñ s k i ,
1962). In adult
European bisons the length o f the branches of this ligament is, on the
average, 165 m m and its thickness 4 mm.
Fig. 3. P elv ic girdle, seen from the ventral side (14 -yea rs-old male).
1 — an terior p u bic ligam ent, 2 — iliolu m bar ligam ent, 3 — quadrate m uscle of
loins.
It should be mentioned, as a supplem ent to the study carried out by
E m p e l & R o s k o s z (1963), that the pelvic sym physis begins to ossify
at the age of 8 years in males and a year earlier in fem ales, the m ere
process o f ossification taking 21 months in males and 18 months in
females.
1.2. S a croiliac Joint
In foetuses, just before birth, and in new -born bisons the auricular
surfaces o f the sacrum and ilium are united by a cartilaginous insertion,
4 mm thick. In cross-sections this insertion shows brow n points arranged
\
44
F. Kobryñczuk
along the future articular cavity of the sacroiliac joint. In calves up to
year of age these points are replaced b y detached or com bined spaces
filled w ith the synovial fluid. Their num ber ranges from 15 to 25. Such
a m ultiple cavity of this joint occurs in 2-year-old European bisons.
In specimens aged 2— 9 or 11 years there is a single articular cavity
in the form o f a fissure. In males from 9 or 11 to 14 years o f age it
becom es obliterated in favou r of a fibrous joint and this, at the age
above 14 years, changes into a synostosis, w hich in old males is so
perfect that the boundary betw een the sacrum and ilium becom es effaced.
In fem ales the fibrous joint, appearing at the same tim e as it does in
males, remains unossified throughout lifetim e.
Table 2
D im ensions o f obturator foram en (in mm).
G roup
Length
n
W idth
X
s
V
8
5
16
17
15
13
co e ff.
13.1
54.4
93.9
108.0
109.9
111.9
2.57
1.0
6.2
12.4
5.7
6.1
3.2
2.3
11.4
13.2
5.3
5.6
2.8
0
4
I
1
II
9
III
6
IV
7
V
18
G row th co e ff.
36.0
X
s
V
21.9
32.2
57.8
68.6
70.0
73.4
3.35
0.6
4.8
6.2
3.7
3.7
5.2
2.7
14.8
10.7
5.4
5.3
7.1
Length: w idth
ratio
Males
0
I
II
III
IV
V
G row th
1:0.51
1:0.60
1:0.62
1:0.64
1:0.64
1:0.64
Females
17.0
86.8
98.2
96.8
98.4
2.73
1:0.51
—
—
9.3
4.2
3.0
2.0
10.7
4.2
3.1
2.0
53.6
61.8
65.4
65.7
3.81
—
4.7
1.0
4.0
2.4
8.8
1.6
6.1
3.6
1:0.62
1:0.68
1:0.68
1:0.66
The fibrous m em brane o f the capsule o f the sacroiliac joint is a hard
sw ollen lip surrounding the articular fissure. In young specimens it is
richly im pregnated with fibrocartilage. On the ventral side of the joint
this m em brane gives rise to the ventral sacroiliac ligament, from which
silvery periosteous fibres run off. They extend m edially and m ingle
with the fibrous ring o f the intervertebral disc of the lum bosacral joint.
M oreover, on the ventral side this joint capsule is strengthened by the
insertion of the quadrate m uscle of loins (Fig. 3— 3).
The interosseous sacroiliac ligament of the European bison consists
of a fairly large num ber o f detached fibres stretched transversely in the
gap betw een the w ings of the sacrum and ilium.
The other ligam ents in the sacroiliac joint o f the European bison have
been described b y W ^ g r z y n & S e r w a t k a (1961).
Joints of hind-limbs of the European bison
45
1.3. O bturator M em brane
In European bisons this m em brane is very thin, hardly 0.1— 0.2 mm
in thickness in adult specimens. It arises at the anterosuperior edge of
the obturator foram en and extends posterom edially. B etw een this m em ­
brane and the free edges of the obturator foram en there is a crescent
aperture, through w hich the internal obturator m uscle com es out o f the
pelvic cavity. The obturator foram en is finally furnished w ith both
obturator muscles. In the obturator m embrane, close to the anterior
edge of the foram en is the obturator canal, the lumen of w hich is equal
to an index finger in diameter.
The obturator foram en is elliptic in shape. Its dimensions (Table 2)
suggest that it attains its almost ultimate size in 4-year-old bisons. The
grow th coefficien t is greater fo r the width than length, w hich indicates
that the foram en becom es m ore and m ore rounded w ith age. Sexual
dim orphism is revealed by the fact that the grow th coefficients of this
foram en are greater in fem ales.
1.4. Other Joints o f the P elv ic G ird le
The ventral surface o f the ilium is united w ith the transverse process
of the last lum bar vertebra b y the iliolum bar ligam ent (Fig. 3— 2).
Its length averages 80 mm and the thickness 3.5 mm in adult specimens.
An additional fibrous structure connecting the transverse process of the
last lum bar vertebra to the hip tuberosity may appear in old European
bisons.
2. Hip Joint
The external outline of the acetabulum of the European bison is
interm ediate betw een a circle and an isosceles triangle. The rim of the
acetabulum is undulate in relation to its bottom and shows three eminen­
ces and three notches, an ilioischiatic, an iliopubic and an ischiopubic.
This last is term ed the acetabular notch proper. In absolute values the
half o f its w idth in the youngest specim ens (Table 3). This w idth is, in
addition, characterized by great individual variation.
On account of the ossification o f the acetabular lip in this place, the
iliopubic notch in the margin of the acetabulum m ay close to form an
additional foram en for the passage o f appropriate vessels.
The acetabular lip (Figs. 4, 5, 6— 1), triangular in cross-section, is
based on the edge of the acetabulum . Externally it passes into the
periosteous fibres and on the side o f the acetabular cavity unites with
its articular cartilage. It is best developed in the ilioschiatic notch, where
in the youngest specim ens it reaches 7 mm in thickness and raises the
edge of the acetabulum by 8 mm. In adults it is respectively 12 mm
46
F. Kobrynczuk
Fig. 4. C ross-section through the left hip join t, seen from behind (m ale foetus).
Fig. 5. C ross-section through the right hip join t, seen from behind (5 -y ea r-old male).
Fig. 6. A n terom edial aspect o f the hip join t (8-y ea rs-old fem ale) (joint capsule
filled w ith gelatin — black in colou r in Figs. 4 and 5).
1 — acetabular lip, 2 — acetabular fossa, 3, 4 — articular cartilage o f acetabulum
and fem ora l head, 5 — epiphyseal cartilage o f fem ora l head, 6 — greater trochanter,
7 — articular capsule, 8 — ilioisch iofem oral ligam ent, 9 — ligam ent o f fem oral
head, 10 — trochanteric bursa o f biceps m uscle o f thigh, 11 — p elv ic bone, 12 —
fem ur.
У
OJ
Table 3
Dim ensions o f p elv ic acetabulum (in m m and cu. cm).
G.C. — grow th coefficien t.
D im ension
G roup
n
V ertical
x
s
0
I
II
III
IV
V
G.C.
8
5
16
17
15
13
36.1
48.5
62.7
63.9
65.0
64.9
1.80
0
I
II
III
IV
V
G.C.
4
1
9
6
7
18
31.2
47.0
55.3
58.0
58.0
58.4
1.88
2.0
3.7
1.7
3.3
3.5
1.4
V
5.5
7.6
2.7
5.2
5.4
2.2
X
38.4
54.0
67.0
69.7
70.3
71.6
1.86
H orizon tal
s
3.0
3.5
2.2
1.4
2.4
3.5
V
7.8
6.5
3.3
2.0
3.4
4.9
X
M ales
20.3
26.6
42.3
43.8
45.1
45.2
2.23
D epth
s
5.0
4.0
1.7
2.6
2.4
4.9
V
24.6
15.0
4.0
5.9
5.3
10.8
X
12.2
32.2
72.4
79.0
84.9
88.6
7.26
Capacity
s
1.2
12.7
8.1
8.8
7.0
6.3
V
9.8
39.4
11.2
11.1
8.2
7.1
W idth o f notch
x
s
v
10.3
8.4
7.5
5.8
4.9
4.9
0.48
1.0
1.4
2.9
1.6
1.7
1.3
9.7
16.7
38.7
27.6
34.7
26.5
2.1
2.3
2.0
1.4
29.6
41.1
32.8
27.4
Fem ales
2.4
3.3
1.4
3.0
4.3
5.7
2.4
5.1
34.7
48.0
58.6
62.5
63.4
63.5
1.83
2.8
2.4
2.0
1.7
4.8
3.8
3.2
2.7
17.6
26.4
34.7
37.4
38.8
39.7
2.26
2.2
-2.1
1.7
3.7
6.3
5.6
4.4
9.3
9.5
31.0
51.5
58.3
61.5
66.2
6.97
8.1
4.7
3.2
6.1
15.7
8.1
5.2
9.2
10.5
6.0
7.1
5.6
6.1
5.1
0.48
Joints of hind-limbs of the European bison
49
thick and 15 mm w ide. At the height of the acetabular notch the lip
passes into the transverse acetabular ligam ent (see further).
The outline of the lunate articular surface of the acetabulum is
8-shaped in the youngest European bisons and consistent w ith its name
in adults. Its pars m ajor is divided from the pars m inor b y a rough
surface having the structure of synovial fossae.
The acetabular fossa (Figs. 4, 5— 2) is approxim ately tw ice as large
in area in the youngest European bisons as it is in adults. In foetuses
and newborns the m eeting point of the three bones of the pelvic girdle
is visible at the centre o f the fossa as a small triangular field built m erely
of a thin m em brane of connective tissue. Three lines of synchondroses
radiate from the central point of the acetabular fossa and divide the
circum ference into three equal parts.
The capacity of the osseous acetabular socket (Table 3), i.e. the socket
w ithout its lip, attains its almost final value in the fou r-y ea r-old bison.
The grow th coefficien t of the depth o f this socket (Table 3) exceeds
this coefficien t fo r its vertical and horizontal dimensions, w hich indicates
that in foetuses and newborns the sockets are relatively shallow er than
they are in adults. The proportions of the three basic linear dimensions
show that on e-year-old European bisons already have their sockets ulti­
m ately shaped.
The sockets w ith the lip preserved are deeper than those w ithout the
lip by 5.5 mm in foetuses and new borns and by 8 mm in adults.
In foetuses and newborns the articular cartilage of the acetabulum
(Figs. 4, 5— 3) is 1.8 m m thick in the pars m inor of the acetabular
surface and up to 2.4 mm in the pars m ajor. In adults the thickness of
this cartilage is the same in both parts, about 0.9 mm. In the youngest
specim ens the articular cartilage passes into the synchondroses of the
three bones o f the pelvic girdle m eeting in the socket.
G eom etrically, the head o f the fem ur o f European bisons is a com bina­
tion of a cone and a hemisphere. M ounted on the neck o f the fem ur, it
form s w ith the shaft an angle of 130°, w hich opens ventrom edially.
The grow th coefficients for the vertical and horizontal dimensions of
the head (Table 4) exceed these coefficients for the respective dim en­
sions of the acetabulum (Table 3). Thus, it m ay be inferred indirectly
that in postnatal life the volum e of the fem oral head of the European
bison increases m ore than does the capacity of the acetabulum.
The articular cartilage of the fem oral head (Figs. 4, 5— 4) blends
peripherally w ith the epiphyseal cartilage (Fig. 4— 5) in foetuses and
new borns. In the same specimens the articular cartilage covers also the
surface o f the neck, m erging into the cartilaginous structure o f the
greater trochanter on the outside (Fig. 4— 6). In adults this cartilage
F. Kobryñczuk
50
covers only the articular surface of the head except for its fovea. The
thickness of this cartilage in the youngest specimens is 4 m m close to
the depression of the head and 5— 6 m m in the periphery. In adults
it is m ore or less uniform , 1.0— 1.5 mm.
In foetuses and new borns the capsule of the hip joint (Figs. 4, 5, 6— 7)
is attached by m ean of its fibrous m em brane to the fem ur just below
the line o f epiphyseal cartilage and extends as far as the greater tro­
chanter on the dorsal side. Its other attachm ent occurs on the acetabular
lip. In adult European bisons the attachm ent of the joint capsule to the
fem ur is shifted in com parison w ith its place in the youngest specimens
beyond the line of the form er epiphyseal cartilage on the dorsal side
T able 4
D im ensions o f fem ora l head (in mm).
G roup
n
V ertica l dim ension
X
0
8
I
5
II
16
III
17
15
IV
V
13
G row th c o e ff.
27.4
38.7
52.2
55.8
58.3
58.8
2.14
4
0
I
1
9
II
6
III
7
IV
18
V
G row th co e ff.
23.7
37.5
47.0
50.4
51.9
52.1
2.20
s
1.3
4.5
1.7
1.5
1.0
1.4
V
M ales
4.7
11.6
3.2
2.7
1.7
2.4
H orizontal dim ension
X
s
V
3.1
3.0
1.5
2.6
3.3
1.9
11.0
7.1
4.4
2.6
5.5
3.1
6.7
2.0
2.0
2.0
13.4
3.7
3.6
3.6
28.3
42.4
56.8
58.8
60.3
60.9
2.15
F em ales
1.8
1.4
2.0
1.4
3.8
2.1
3.8
2.7
24.5
43.3
50.0
53.6
£4 8
55.3
2.26
and extends to the place w here the neck passes into the shaft of the
bone on the ventral side. Thus, in both cases in adult specimens the
fibrous m em brane o f the capsule of this joint is attached to the fem oral
head at a distance of about 2 cm from the edge of the articular surface
to the outside. A s in the youngest specim ens, its other attachment
is on the acetabulum.
T w o parts of the synovial m em brane can be distinguished in the hip
join t o f the European bison. One o f them participates in the form ation
of the joint capsule, the other is connected w ith the ligament of the
fem oral head (see further).
In foetuses and new borns the synovial membrane, w hich goes to
the making of the joint capsule accom panies the fibrous membrane
Joints of hind-limbs of the European bison
51
from one attachment to the other. In adults a similar situation is
observed as regards the attachments of both m em branes to the aceta­
bulum, whereas on the fem ur the synovial m em brane is attached to
the edges of the articular cartilage, from w here it goes on to the
circum ference, to the attachments o f the fibrous m embrane, thus covering
the neck stripped of articular cartilage. It has how ever been fou n d that
the synovial mem brane covering the fem oral neck in m iddle-aged Euro­
pean bisons disappears in old specimens.
The capsule of the hip joint is m oderately tense. It is thickest on the
dorsal side. On the posterior side of the joint it form s a recess the size
o f a cherry, w hich is pressed into the trochanteric fossa as the synovial
bursa of the obturator muscles.
The capacity of the cavity of the hip joint is 11— 17 ml in age group
0, 20— 29 ml in group I, 27— 40 ml in group II and on the average
36— 50 ml in adults. The low er limits refer to fem ales, the upper ones
to males. The amount of the synovial fluid that I managed to obtain
from the cavity of this joint was 6— 8 ml in adults.
Ligaments of the Hip Joint
The ilioischiofem oral ligam ent (Fig. 6— 8) begins antero-inferiorly at
the base of the greater trochanter and ends, split into tw o portions,
in the region of the ischiatic spine. Its undivided part is 20 m m w ide
and 2.5 m m thick in foetuses and newborns and, respectively, 45 and
5 mm in adults.
The transverse acetabular ligament connects the margins of the
acetabular notch and at the same time bounds the acetabular foram en,
through w hich run the extra-acetabular portion o f the ligam ent o f the
fem oral head and ram ifications of the synovial m em brane w ith small
vessels. The length of this ligam ent corresponds to the w idth o f the
acetabular notch (Table 3). Its width is 8.2 mm and the thickness 3.0 mm
in the youngest European bisons and, respectively, 17.6 and 7.3 m m in
adults. The m edial edge of the transverse acetabular ligam ent gives
attachm ent to a part of the ligament of the fem oral head.
The ligam ents o f the fem oral head (Figs. 4, 5— 9) arises in the fovea
o f the fem oral head and runs towards the acetabular notch. Here, it
divides into a part that ends in the periosteum of this notch and a part
that mingles w ith the transverse acetabular ligament. A band branches
o f f the latter part and passes out of the acetabular cavity through the
acetabular foram en to terminate in a special groove of the ischium.
T his band is an extra-acetabular portion of the ligam ent of the ligament
o f the fem oral head.
.
Starting from its attachment to the fem ur the ligam ent discussed is
52
F. Kobrynczuk
accom panied by a fold o f the synovial membrane, w hich enwraps it. In
the region of the acetabular fossa the fold w idens ca lix-lik e and is
attached to the edges of this fossa w ithout com ing into contact w ith
its bottom . A part o f the synovia m em brane enters the acetabular
foram en to accom pany the extra-acetabular portion of the ligament
under discussion.
The length of the ligam ent of the fem oral head is 25 m m in foetuses
and new borns of both sexes, 35 mm in adult males and 31 m m in
adult fem ales. The ratio of the length of this ligam ent to the horizontal
dim ension o f the acetabulum is as 0.65:1 in the group of the youngest
males and on the average as 0.50:1 fo r adult males. In fem ales this ratio
is as 0.72:1 in the youngest group and as 0.49:1 for adults. If in turn
the length of this ligam ent is com pared w ith the horizontal dimension
o f the fem oral head, the ratio is, respectively, as 0.88:1 and 0.59:1 for
the youngest and adult males and as 1.0:1 and 0.57:1 for the youngest
and adult fem ales. These com parisons show that the ligam ent of the
fem oral head is relatively longer and, what is more, considerably longer
m the first days o f life o f European bisons than it is in adult specimens.
As regards the thickness and w idth of the ligam ent concerned, indi­
vidual variation is very great. In group 0 of males the w idth averages
4.7 mm, ranging from 3.6 to 6.0 mm, whereas in adult males the mean
is 13.5 mm and the range 8.4— 19.0 mm. These data in fem ales are x —
6.7 m m (3.5— 8.8 m m ) fo r the youngest specimens and ¿c = 11.5 mm
(5.0— 17.0 mm) fo r adults. Som ew hat smaller individual variation is ob­
served in the thickness o f this ligament. The mean thickness is 3.0 mm
(2.0— 3.5 mm) fo r groups 0 of both sexes and, respectively, 5.5 mm
(3.0— 8.0 mm) and 7.5 m m (5.0— 10.0 mm) for adult males and females.
3. Stifle Joint
3.1. F em orotibial Joint
The condyles o f the fem ur and the long axis of this bone form an
angle of 110°, open to the rear. The length, or the chord, and the width
of one condyle approxim ate the corresponding dimensions of the other
(Table 5). The length:w idth ratio of the condyles changes w ith age in
favour of the first dim ension (Table 5). The angle of the curvature of
the m edial condyle is som ewhat greater than that of the lateral condyle
(Table 5) or, in other w ords, the m edial con dyle is m ore convex than
the lateral. The length of the curvature radius of the lateral condyle
(Table 5) is m ostly som ewhat greater than that of the m edial. The lateral
condyle lies in a sagittal plane, whereas the m edial one deflects medially
Joints of hind-limbs of the European bison
53
from this plane at an angle of 20°. The medial condyle is, in addition,
pushed farther to the back and, as a result, the rotation axis in the
fem orotibial joint runs obliquely to the sagittal plane, w ith w hich it
form s an angle of 77°.
T ab le 6
D im ensions o f distal end o f fem u r (in mm ).
A — w idth ratio o f distal end o f fem u r to in tercon d y lar fossa.
G rou p
n
Lenght
x
s
W idth
x
v
s
v
W idth o f in te r co n ­
d ylar fossa
X
s
V
A
E uropean bison, males
0
I
II
III
IV
V
G. C.
8
5
16
17
15
13
77.3
102.5
134.7
144.2
144.6
144.3
1.78
0
I
II
III
IV
V
G. C.
4
1
9
6
7
18
79.5
103.0
117.8
128.0
131.0
136.2
1.72
4
126.9
7.2
2.8
8.2
8.1
5.7
6.1
9.3
2.7
6.1
5.6
3.9
4.2
67.3
85.0
105.2
114.9
117.3
120.8
1.79
3.2
5.6
5.6
4.0
3.9
7.2
4.8
6.6
5.3
3.5
3.3
6.0
20.8
19.5
18.1
17.6
17.1
17.1
0.82
2.4
1.4
0.6
1.0
2.0
1.0
11.5
7.2
3.3
5.7
11.7
5.8
1:0.31
1:0.23
1:0.17
1:0.15
1:0.14
1:0.14
6.0
9.2
7.9
10.3
1:0.29
1:0.24
1:0.19
1:0.17
1:0.17
1:0.16
European bison, fem ales
9.0
4.0
2.6
6.7
7.6
3.1
2.0
4.9
64.0
82.0
94.4
103.6
104.5
109.0
1.70
6.2
3.0
7.4
3.6
6.6
2.9
7.1
3.3
18.7
20.0
18.2
17.4
17.8
17.5
0.94
1.1
1.6
1.4
1.8
Bison bison L., fem ales
1
147.0
93.6
14.2
1:0.15
Bison bonasus — Bison bison hybrid, m ale
126.0
13.8
1:0.11
D om estic cattle, m ales
2
159.0
131.8
15.4
1:0.12
D om estic cattle, fem ales
10
135.4
1
140.5
10.2
7.5
100.8
1.4
1.4
15.2
2.0
7.9
1:0.15
W atussi cattle, m ale
124.5
11.1
1:0.09
11.2
1:0.11
17.2
1:0.15
13.6
1:0.14
6.5
1:0.8
W atussi cattle, fem ale
1
131.0
106.5
Bubalus bubalus L., male
1
135.5
113.0
Zebu, fem ale
1
119.0
94.5
Yak, .fem ale
1
91.5
81.0
54
F. Kobrynczuk
The intercondylar fossa becomes narrow er with age (Table 6). C on­
sequently, the w idth ratio o f the distal end of the fem ur to this fossa
changes during postnatal life (Table 6). A s the length of the distal end
o f the fem ur hardly differs from the w idth in growth coefficient (Ta­
ble 6), its com parison w ith the width o f the intercondylar fossa gives
a similar ratio.
The articular cartilage o f the fem oral condyles (Fig. 8— 1) changes its
colour in course o f life. In foetuses and newborns it is pale blue, in
animals above one year of age cream, and in the oldest ones almost
white w ith patches in brick-red colour. In some males, m ore than 15
Fig. 7. The angle o f in clin ation (a) of the fem ora l trochlea in the foetus (left)
and the adult E uropean bison (right),
a, b, c — a u x ilia ry lines used to calculate the angle a.
years old, and in fem ales 2— 3 years older, degenerative changes appear
in this cartilage, affecting circular areas, 1— 2 cm in diameter, adjacent
to the intercondylar fossa. In males the thickness of the articular carti­
lage on the lateral condyle is 5.7 m m in group 0, 3.0 mm in group I
and 1.3 mm, w ith small deviations, in the other groups, and in females
it is, respectively, 4.4, 1.4 and 1.3 mm. On the medial condyle the thick­
ness of this cartilage is 6.3 mm in group 0, 2.7 mm in group I and 1.3 mm
in the remaining groups in males and, respectively, 3.9, 1.2 and 1.3 mm
in females.
The articular surfaces o f the tibial condyles resemble saddles in shape.
Their articular cartilage (Figs. 8, 9, 10— 2) is characterized by physical
Joints of hind-limbs of the European bison
55
features similar to those of the cartilage described above, not excluding
pathological changes, w hich on both condyles occur in the vicinity of the
appropriate intercondylar tubercles. The thickness of this cartilage on
both condyles is 2.6 mm in the youngest males and 1.7— 1.8 mm in the
remaining ones, whereas in fem ales it is, respectively, 2.2 and 1.3— 1.4
mm. In foetuses and new borns the articular cartilage of the condyles
of the tibia passes into its epiphyseal cartilage (Fig. 10— 3).
Menisci
The m enisci of European bisons, the lateral (Figs. 8, 11, 12, 13, 15,
16— 4) and the m edial (Figs. 11, 12, 14, 15, 16— 5), resem ble the conchae
of human ear in outline. In m ost cases the lateral m eniscus is slightly
longer and broader than the m edial one (Table 7). Both these fibrocartilages are thinnest in the m iddle o f their length, the lateral meniscus
being thickest at the back and the m edial one in the front. In the course
of postnatal life the relative thickness at the back o f the lateral
meniscus increases considerably, becom ing sm aller in the m iddle
portion. The shape o f the m edial m eniscus hardly changes during
life. The thickness given in Table 7 is a mean value from its three
measurements analysed. Sexual dim orphism is m anifested by the fact
that in males the grow th coefficien t of the thickness is greater for the
medial meniscus in contradistinction to the situation in fem ales. In both
sexes this thickness is m uch greater in the lateral m eniscus. As regards
volum e, the lateral meniscus exceeds the medial, but not strikingly
(Table 7). In analysing the increm ents in this parameter, one can see
that the menisci of males attain their final size earlier than those of
females. In the oldest specimens the m enisci sometimes undergo de­
form ation. Their peripheral margins lose their characteristic sem icircular
line and the fibrous processes their m orphological distinctness, the axial
margins look like unhem m ed fabric and the notches becom e lengthened
and deepened. The menisci are cream in colour, w ith brick-red stripes.
N
M eniscal Ligam ents
The anterior tibial ligam ent of the lateral meniscus (Fig. 11— 7) arises
from the anterior end of the meniscus and ends betw een the layers of
the anterior cruciate ligam ent in the m edial intercondylar area.
The anterior tibial ligam ent of the m edial meniscus (Fig. 11— 8) is the
extension of the anterior end of the meniscus. It ends in front of the
attachment of the previous ligam ent in the lateral intercondylar area.
The anterior ligaments o f the menisci form an obtuse angle, pointing
to the front. This angle is 180° in the phase o f extension and 130° in
the phase of flexion.
11
Fig. 8. Sagittal section o f right stifle join t through lateral fem ora l condyle. M edial
v iew (8 -y ea r-old male).
Fig. 9. Sagittal section through the m iddle o f the left stifle joint. L ateral v iew
(8 -y ea r-old male).
Fig. 10. Sam e as Fig. 9 (m ale foetus).
Fig. 11. M enisci o f the right fem orotib ia l join t, seen from above (9 -y e a r-o ld fem ale).
Explanations to figures 8— 16 see page 60.
14
Fig. 12. L e ft stifle jo in t after the rem oval o f the lateral sac
joint. A n terior v iew (10 -yea r-old male).
Fig. 13. Right stifle joint. Lateral v iew (8 -y ea r-old
Fig. 14. L e ft stifle joint. M edial v iew (10 -yea r-old
Fig. 15. Right stifle join t after the rem oval o f the m edial sac
joint. Posterior v iew (8 -y ea r-old male).
o f the fem orotibia l
male).
male).
o f the fem orotibia l
60
Fig.
F. Kobrynczuk
16. L eft stifle join t after the rem ov a l o f the jo in t capsule. Posterior
(1 0 -y ea r-old male).
v iew
(In Figs. 8, 9, 10, 12, 13, 14 and 15 the join t capsules are filled w ith gelatin; in
Figs. 8 and 9 — black in colour).
1, 2 — articular cartilage o f fem ora l and tibial condyles. 3 — epiphyseal cartilage
o f tibia, 4, 5 — lateral and m ed ial m enisci, 6 — p rox im a l bursa o f lateral collateral
ligam ent, 7, 8 — anterior tibial ligam ents o f lateral and m edial m enisci, 9, 10 —
posterior tibial ligam ents o f lateral and m edial m enisci, 11 — fem ora l ligam ent
o f lateral m eniscus, 12 — capsule o f fem orotib ia l joint, 13 — jo in t partition, 14 —
su bexten sor pouch, 15 — tendons o f lon g digital ex ten sor and third peroneal m uscle,
16 — su bpopliteal pouch, 17 — popliteal m uscle, 18 — lateral collateral ligam ent,
19 — distal bursa o f lateral collateral ligam ent, 20 — m edial collatera l ligam ent,
21 — distal bursa o f m edial collateral ligam ent, 22, 23 — anterior and posterior
cruciate ligam ents, 24 — lon gitu din al ligam ent o f stifle, 25, 26 — articular cartilage
o f fem ora l trochlea and patella, 27 — capsule o f fem orotib ia l join t, 28 — m iddle
patellar ligam ent, 29 — lateral patellar ligam ent, 30, 31 — its lateral and m edial
portions, 32 — bursa o f biceps m u scle o f thigh, 33 — m edial patellar ligam ent,
34 — fem oropatellotibial ligam ent, 35 — in frapatellar fat b od y, 36 — fem u r, 37 —
tibia, 38 — patella, 39, popliteal artery, 40 — fatty tissue, 41 — third peroneal
m uscle.
Joints of hind-limbs of the European bison
61
The posterior tibial ligam ent of the lateral m eniscus (Figs. 11, 15,
16— 9), the broadest and at the same time thinnest of the ligaments
under discussion, arises from the posteroinferior edge of the meniscus
and ends it the edge of the popliteal notch of the tibia, laterally to the
terminal attachm ent of the posterior cruciate ligament.
The posterior tibial ligam ent of the medial meniscus (Fig. 11— 10),
covered by the posterior cruciate ligament from behind and by the
medial portion of the longitudinal ligament of stifle from above (Fig.
11— 24), arises from the posterior end of the meniscus, runs forw ard and
inward and ends in the posterior intercondylar area.
The fem oral ligam ent of the lateral meniscus (Figs. 10, 11, 15, 16,— 11)
has it origin on the fem ur, at the height of the upper edge of its medial
condyle. H ence it extends downw ards and laterad towards the peripheral
border of the lateral meniscus. It is the only ligam ent of those discussed
that is connected with the fibrous m em brane of the joint capsule.
A ll the ligaments of the menisci are characterized b y the parallel and
relatively loose arrangem ent of their fibres.
The capsule of the fem orotibial joint (Figs. 8, 9, 10, 14, 15— 12) is loose
and com paratively thin. Its fibrous m em brane is best developed on the
posterior side of the joint, w here it blends w ith the adventitia of the
and virtually lacking in the front. In the youngest European bisons the
fibrous m em brane is attached along the line o f the epiphyseal cartilages
of the fem u r and tibia and to the peripheral edges of the menisci except
the groove for the tendon of the popliteal muscle, impressed in the
peripheral border of the lateral meniscus. In adults the attachments of
this m em brane are m oved away from the m argin o f the joint by about
2— 3 cm on the fem ur and coincide w ith these m argins on the tibia.
The synovial m em brane form s tw o sacs, a lateral and a medial, which,
clinging to each other in a sagittal plane of the joint, constitute a par­
tition of the joint (Fig. 10— 13). The cavities of these sacs com m unicate
with each other in 7 per cent of cases, and the ch iefly ini old bisons,
by an aperture in the partition. The medial sac com m unicates with the
fem oropatellar joint cavity in all European bisons. The lateral sac form s
two pouches and a synovial bursa. Its subextensor pouch (Figs. 8, 13— 14)
has the shape of an elongate drop and underlies the com m on tendon
of the long digital extensor and third peroneal m uscles in the groove
for m uscles in the tibia. Measured from the m argin of the articular
surface of the tibia, this pouch hangs for a distance of 4 cm in the
youngest European bisons and 7— 8 cm in the oldest specimens. The other
pouch o f the lateral sac of the joint under description is the subpopliteal pouch (Figs. 8, 13, 15— 16), w hich extends under the tendon of origin
of the popliteal m uscle (Figs. 13, 15, 16— 17). The third structure of the
T able 8
D im ensions
of
collateral ligam ents
of
fem orotibia l join t and cru ciate ligam ents o f stifle join t (in mm). L — length.
W — w idth, T — thickness.
C ollateral ligam ents___________________
Uroup
n
C ruciate ligam ents
M edial
L ateral
L
W
T
■
'
L
W
'
A n terior
T
L
W
P osterior
T
L
W
T
M aies
0
I
II
III
IV
V
G.C.
8
5
11
8
8
7
4*1.0
54 0
78.0
82.0
86.0
88.0
1.91
6.9
7.3
12.7
17.5
17.6
18.1
2.62
1.2
1.8
3.4
3.7
4.2
4.4
3.67
64.0
80.0
135.0
142.0
140.0
142.0
2.22
7.3
7.7
14.0
17.2
17.1
17.0
2.33
0
I
II
III
IV
V
G.C.
4
1
6
1
3
9
43.0
63.0
65.0
84.0
74.0
79.0
1.82
6.3
6.5
9.5
13.5
15.3
15.9
2.52
1.2
1.5
3.3
3.5
3.8
4.0
3.33
60.0
86.0
90.0
129.0
120.0
123.0
2.06
7.2
9.0
13.2
12.8
15.0
15.1
2.10
1.4
1.7
3.4
3.9
4.4
4.7
3.36
33.0
41.5
55.0
57.0
55.0
50.0
1.52
4.0
6.0
10.1
9.7
9.9
9.8
2.45
3.4
3.8
6.2
6.3
6.3
6.1
1.79
44.0
52.0
67.0
70.0
68.0
65.0
1.48
7.5
8.0
8.4
8.3
8.4
8.2
1.09
2.2
2.6
5.3
5.4
5.5
5.0
2.27
1.2
0.8
2.5
2.8
3.5
4.0
3.33
28.0
39.0
42.0
44.0
’ 42.0
42.0
1.53
3.8
6.0
7.2
7.0
7.3
8.0
2.11
3.2
4.0
4.6
3.5
4.7
5.2
1.66
42.2
46.0
56.0
55.0
57.0
58.0
1.37
5.8
6.1
5.8
6.4
6.0
6.1
1.05
2.2
3.0
4.7
5.0
4.7
4.8
2.18
Fem ales
Joints of hind-limbs of the European bison
63
lateral sac is the proxim al synovial bursa of the lateral collateral liga­
ment of the fem oropatellar joint (Figs. 13, 15— 6). This bursa separates
the ligam ent from the substratum, i.e. the tendon o f origin of the popli­
teal muscle.
Ligam ents of the Femorotibial Joint
The lateral collateral ligament (Figs. 11, 12, 13, 15, 16— 18) arises from
the fossa for its attachment on the femur. The deep portion of this
ligam ent ends on the lateral surface of the tibial condyle and its super­
ficial portion passes finally into the fibrous band of the leg, constituting
its superficial layer (Fig. 17— 2). The distal synovial bursa of this liga­
ment (Fig. 13— 19) is autonomous.
The medial collateral ligament (Figs. 11, 12, 14, 15, 16— 20) runs from
the fossa of its attachment on the medial side of the fem ur to the tibia,
where its attachm ent form s a line parallel to the long axis of this bone.
This ligam ent has only a distal synovial bursa (Fig. 14— 21).
The collateral ligam ents of the fem orotibial joint are loosely connected
with the articular capsule and menisci. The medial ligament can be par­
ticularly easily displaced in a sagittal plane. The measurements of these
ligaments are given in Table 8.
The anterior cruciate ligament (Figs. 9, 10, 11— 22) arises from the
intercondylar fossa of the fem ur and extends forw ard and downward.
A bove the posterior intercondylar area of the tibia it reveals a tendency
to split into a lateral and a medial part. Crossing the posterior cruciate
ligament on the lateral side it turns right by 90° round its long axis
and thus its m edial portion becom es a superficial layer and the lateral
portion a deep one. The superficial layer runs to the front above the
attachm ent of the anterior tibial ligament of the lateral meniscus on
the tibia and ends in the anterior intercondylar area. The deep layer
reaches the anterior part of the central intercondylar area and term i­
nates posteriorly to the attachment of the above-m entioned ligam ent of
the lateral meniscus.
The posterior cruciate ligament (Figs. 9, 10, 11, 15, 16— 23) goes away
obliquely from the anteromedial part of the intercondylar fossa of the
fem ur. It extends backwards, clinging closely to the floor o f this fossa,
which, being a sem icylinder, constitutes a sort of pulley for it. The liga­
ment ends flattened at the edge of the popliteal notch of the tibia, partly
overlapped by the fem oral ligament of the lateral meniscus. The measu­
rements o f the cruciate ligaments are given in Table 8.
The longitudinal ligament of the stifle (Fig. 11— 24), w hich is a thin
structure (7 mm w ide and 1 mm thick in adults), extends antero-poste-
F. Kobrynczuk
64
T ab le 9
D im en sions o f trochlea o f fem u r (in m m and degrees).
G.C. — g row th coefficien t, W . — w idth.
G roup,
name
W . p rox im a l
n
x
s
4.4
5.8
4.6
3.2
3.3
2.8
A n g le o f trochlear
ridges
W. distal
V
x
s
European bison, m ales
16.2
24.3
3.6
16.3
30.8
4.7
8.4
41.1
3.5
5.5
44.4
3.5
5.4
45.1
2.2
4.6
44.8
1.4
1.84
V
X
s
14.8
15.2
8.5
7.9
4.9
3.1
4.0
4.4
9.8
9.3
10.2
9.5
2.37
2.3
1.9
1.4
1.0
1.6
1.7
57.5
43.5
14.2
10.7
15.7
17.9
1.9
0.5
1.7
1.5
22.3
5.5
20.4
15.7
1.4
18.1
0
I
II
III
IV
V
G. C.
8
5
16
17
15
13
27.2
35.6
55.0
58.6
61.1
60.5
2.22
0
I
II
III
IV
V
G. C.
4
1
9
6
7
18
24.0
35.5
46.1
51.0
51.0
55.2
2.80
4
49.7
1
E u ropean bison — A m erica n bison hybrids, male
9.4
46.8
62.0
4.1
1.0
1.4
4.7
E uropean bison, fem ales
21.7
31.5
8.9
35.8
4.2
0.2
39.0
1.0
2.7
39.2
2.0
8.5
41.2
4.2
1.90
11.7
2.6
5.1
10.2
A m erican bison, fem ales
40.4
2.8
1.7
8.5
9.1
8.3
9.5
3.42
9.2
Fest
Felon
Fellach
Festyn
Fey
E uropean bison — dom estic cattle hybrids, fem ales
11.7
55.0
48.0
8.3
49.0
62.0
8.8
54.0
68.0
8.8
68.0
54.0
10.3
63.0
47.0
'
Felly
Fela
Fama
E uropean bison — dom estic cattle hybrids, fem ales
11.7
45.0
60.0
9.3
49.0
62.0
9.3
43.0
56.0
2
70.0
10
53.9
1
55.0
D om estic cattle, m ales
59.0
4.5
D om estic cattle, fem ales
8.3
43.6
4.5
6.6
10.3
7.8
W atussi cattle, m ale
43.8
8.1
50.8
W atussi cattle, fem a le
43.8
5.5
1
56.0
Bubalus bubalus, m ale
39.4
11.8
1
46.5
Z ebu , fem a le
38.0
6.5
1
37.5
Y ak, fem a le
34.8
3.5
1
v
Joints of hind-limbs of the European bison
65
riorly. It is situated between the lamellae of the joint partition. It arises,
together w ith the superficial layer of the anterior cruciate ligam ent, from
the tibia. Passing to the rear, it divides into tw o unequal bands, w hich
end at the edge of the popliteal notch of the tibia.
3.2. F em oropatellar Joint
In its structure and function the trochlea of the fem ur of the European
bison com bines elements of a technical pulley, w edge and screw . Its
proxim al w idth exceeds the distal one, and the same is true of the grow th
coefficients o f these widths (Table 9). In consequence, the planes in w hich
the trochlear ridges lie cross below the stifle, form ing a dihedral angle,
the size o f w hich is slight in foetuses and new borns and discernible
without the necessity of measuring in adults (Table 9). The chord or
length of the lateral ridge is smaller than that of the medial. The grow th
coefficien t o f the chord is greater for the medial ridge (Table 10). The
radius o f the medial ridge is greater than that of the lateral and so is
its grow th coefficient. The angle of the curvature is always larger in the
lateral ridge. The differen ce between these param eters increases with
age (Table 10). The fem oral trochlea is deflected laterally b y an angle
of 75° in group 0, this angle being 69° in adults (Fig. 7). O ver its extent
from the top downwards, the articular surface of the trochlea undergoes
a 38° torsion towards the medial side.
The articular cartilage of the fem oral trochlea (Figs. 8, 9, 10— 25)
blends w ith the epiphyseal cartilage in young specimens. It covers the
articular surface of the trochlea and also a small area of its m edial sur­
face. Beyond the medial ridge it passes into the articular cartilage to
the medial condyle of the fem ur. Its thickness on the ridges is consi­
derable, ranging from 5.0 to 9.0 mm in groups 0 and I in males and
from 2.8 to 7.3 mm in these groups in fem ales. In adult bisons of both
sexes it is 1.3— 1.9 mm.
The articular surface o f the patella, together w ith the fibrocartilage
of the patella arising from it, constitutes a partly elastic exact cast of
the fem oral trochlea. The thickness of the articular cartilage (Figs. 8, 9,
10— 26) ranges from 2.9 to 4.3 mm in males of groups 0 and I and from
2.0 to 2.6 mm in fem ales of these groups. In adults of both sexes it is
1.3— 2.1 mm. It is characterized by great individual variation.
The fibrocartilage of the patella is in the form of a lamella inclined
towards the trochlea. In adult European bisons its thickness is up to
3 cm. Interiorly it passes into the medial patellar ligam ent (Figs. 11, 12,
14— 33).
The capsule of the fem oropatellar joint (Figs. 8, 9, 10, 12, 13, 14,
15— 27) is virtually represented by the synovial mem brane. This capsule
66
F. Kobrynczuk
is attached to the articular edges of both bone components of this joint.
On the m edial side it passes into the capsule of the fem orotibial joint.
As a whole, it is a room y, loose, excessively developed sac. A b ov e the
patella this excess manifests itself as a blunt pouch directed upwards,
the length of w hich approxim ates that of the patella, whereas below the
patella the capsule develops two swells on the lateral side and one on
the medial (Figs. 12, 14— 27). The synovial membrane of the fem oropatellar joint is characterized by very great accumulations of villi and
folds. Their largest num bers occur in the region of the apex of the patella
and at its base. The villi are hair-shaped and 1— 6 cm long in adults.
The height of folds reaches 3 cm.
The total capacity of the cavity of the fem oropatellar joint and the
medial sac of the fem orotibial joint averages 80 ml in foetuses and new ­
borns, 125 ml in calves of group I, 225 ml in adult males and 180 ml in
adult fem ales. The amount of synovial fluid obtained from this com m on
cavity is up to 14 ml in adult males and 9 ml in adult females. The
capacity o f the lateral sac of the fem orotibial joint is 16 ml in group 0
and 70 ml in group I of both sexes and, respectively, 100 and 80 ml in
adult males and females.
Ligaments of the Fem oropatellar Joint
The m iddle patellar ligament (Figs. 8, 9, 10, 11, 12, 14— 28) is a band,
uniform in w idth, directly underlying the fascia of the stifle. It is a dow n­
ward continuation of both the tendon of the rectus muscle of the thigh
and the medial portion o f the lateral vastus muscle. Out of the ligaments
o f the patella the m iddle patellar is the only one that has no connections
w ith the joint capsule, since it is separated from it by the infrapatellar
fat body, in w hich it carves a suitable groove. On the tibia it crosses
the attachment of the lateral patellar ligament. A spacious infrapatellar
bursa, 36 by 45 mm in adults, occurs under the insertion of the middle
patellar ligament.
The lateral patellar ligam ent (Figs. 8, 9, 11, 12, 13, 14— 29) extends
at an angle o f 30°, pointing downwards, to the previous ligament. It is
in the form of a trapezium oriented with its shorter base dow n and
consists of tw o portions, a lateral (Fig. 12— 30) and a medial (Fig. 12— 31).
The lateral portion is the extension of the tendon of the anterior part
of the fem oral biceps m uscle. The medial portion, arising from the side
edge of the patella, is som ewhat wider than a half of the w idth of the
lateral portion. Under the unpaired part of this ligament is the spacious
subtendinous bursa o f the biceps muscle (Fig. 13— 32), its capacity being
3— 4 ml in the youngest specimens and 40— 50 ml in adults. The lateral
patellar ligam ent blends w ith the subextensor pouch of the capsule of
the fem orotibial joint.
Joints of hind-limbs of the European bison
67
The m edial patellar ligam ent (Figs. 11, 12, 14— 33) is a thin tape bent
to the rear like a sabre. The further tw o-thirds of this ligament is over­
lapped by the fem oropatellotibial ligament (Figs. 12, 14— 34). The m edial
patellar ligam ent arises from the fibrocartilage of the patella and is
inserted into the tibia. O nly in the m iddle section it loses contact with
the articular capsule, being separated from it by the infrapatellar fat
body.
The dimensions of the patellar ligam ents are given in Table 11. In
analysing these measurements, w e find that the lateral ligam ent is the
strongest of them and the m edial one the weakest in adults and that
Table 11
D im ensions o f patellar ligam ents (in mm).
^
G rou p
n
Lateral
M iddle
M edial
----------------------------------------------------------------------------------------------------------------------L en gth W idth T h ick n . Length W idth T hickn. Length W idth T h ick n .
M ales
0
I
II
III
IV
V
G.C.
8
5
11
8
8
7
70.4
82.0
106.0
114.0
121.0
130.0
1.75
19.2
17.5
25.4
31.0
42.0
42.0
2.19
2.4
3.0
7.3
7.4
9.1
9.3
3.88
0
I
II
III
IV
V
G.C.
4
1
6
1
3
9
58.0
90.0
93.0
108.0
110.0
99.0
1.71
14.0
16.0
23.0
23.0
28.0
32.0
2.31
2.1
3.0
5.9
6.0
7.7
8.1
3.86
88.0
126.0
150.0
159.0
159.0
158.0
1.80
19.3
18.1
25.7
27.2
29.1
28.3
1.47
2.2
3.5
5.1
5.4
6.2
6.1
2.77
74.0
63.4
139.0
161.0
143.0
141.0
1.91
8.8
11.7
22.6
28.4
28.5
29.0
3.30
1.3
1.7
2.9
3.8
4.3
4.5
3.46
16.8
18.5
22.0
22.0
24.0
25.0
1.49
2.1
4.0
4.2
4.1
5.7
6.1
2.90
70.0
106.0
124.0
111.0
135.0
127.0
1.81
8.3
11.2
19.5
18.0
20.0
23.8
2.87
0.8
1.0
1.6
2.7
3.5
3.8
4.75
Fem ales
97.0
112.0
121.0
130.0
130.0
124.0
1.28
the medial ligam ent undergoes the greatest postnatal developm ent and
the m iddle one the smallest.
The fem oropatellotibial ligam ent (Figs. 12, 14— 34) is Y -shaped. Its
arms connect the patella to the fem ur and correspond to the m edial
fem oropatellar ligament. The body o f this structure arises from the
aponeurosis of the m edial vastus m uscle and the fascia of the stifle.
It partly overlaps the medial patellar ligament, with which it terminates
on the tibia.
T he volum e of the infrapatellar fat body (Figs. 9, 10, 14— 35) is 15 cu.
cm in the youngest European bisons. In adult males it ranges betw en 50
and 120 cu. cm and in adult fem ales betw een 35 and 100 cu. cm. It
depends to a great extent on the general condition of the animal.
68
F. Kobrynczuk
4. Fibrous Band of the Leg
The fibrous band o f the leg (Fig. 17) is considered to be a remnant
of the shaft of the fibula. It is set among appropriate muscles of this
region. P roxim ally it is divided into tw o layers, a superficial (Fig. 17— 2),
w hich is a dow nw ard prolongation of the lateral collateral ligament of
17
18
Fig. 17. Fibrous band o f leg.
1 — fib rou s band o f leg, 2 — its su perficial layer (lateral collateral ligam ent o f
fem orotibia l joint), 3 — deep layer, 4 — interosseous m em brane o f leg.
Fig. 18. V estigial fibulae. F rom left to right, three fibulae o f 8-, 10- and 15-year-old
m ale E uropean bisons, the fou rth one o f an adult m ale bu ffalo.
the fem orotibial joint, and’ a deep (Fig. 17— 3), w hich arises from the
tibia, in the place w h ere its fibular process is present in dom estic cattle.
The uniform band ends on the lateral surface of the tibia, in the distal
third of this bone. The interosseous space of the leg, occurring between
Joints of hind-limbs of the European bison
69
this band and the tibia, is filled with the thin interosseous membrane
of the leg (Fig. 17— 4), in w hich there are tw o interosseous foramina.
The presence of the vestigial shaft of the fibula in the band (Fig. 18)
in 15 per cent o f the European bisons belonging to groups III— V is
a notably individual character.
5. Hock Joint
Intertarsal Joints
Tarsocrural joint. The concave articular surface of the cochlea of the
tibia is described w ith the same radius at the corresponding ridges of the
proxim al trochlea of the tibial tarsal articulating with it (Table 12). The
angle of the curvature of the articular grooves of the cochlea is smaller
than that of the ridges of the proxim al trochlea of the tibial tarsal
corresponding w ith them b y about 65°, w hich constitutes the flexion ­
extension range o f the m ovem ents in this joint. The thickness o f the
articular cartilage of the cochlea is strikingly small com nared w ith the
Fig. 19. S yn ovia l fossae (x) on the articular surfaces o f the h ock joint. A — articular
su rface o f tibial coch lea, B — articular su rface o f tibial tarsal bone, C — articular
su rface o f T c + T I V .
thickness of this cartilage in other joints. It is 1.1 mm in groups 0 and I
and only 0.1— 0.2 m m in older groups.
A characteristic m orphological detail of this cartilage of the tibial
cochlea in all European bisons is the presence of a synovial fossa in it
(Fig. 19 A — x). In adult bisons it is visible also on digested bones, w hich
indicates that the defect affects not only the cartilaginous tissue but the
bone as w ell. In foetuses and new borns the future area of this fossa
is m anifested as a place coated with a very thin light-brow n connectivetissue mem brane, w hich peripherally passes into articular cartilage. As
regards several-m onth-old calves, the m em brane is already missing and
the bottom of the fossa is of rough osseous tissue with numerous nutrient
70
F . K o b ry n cz u k
foram ina scattered all over it. In the youngest and oldest males the
length of the fossa is 15 and 25 mm, respectively, and the w idth 7 and
10 mm. In the youngest and oldest fem ales this length is correspondingly
12 and 22 m m and the w idth 6 and 10 mm. The depth o f the fossa is
approxim ately tw ice as great as the thickness of the surrounding articular
cartilage.
The proxim al trochlea of the tibial tarsal has tw o sagittal ridges
unequal in breadth. The lateral ridge is broader and therefore blunter
than the m edial one. The angles of the curvatures of both trochlear
ridges are given in Table 12. The angle is sm aller in the lateral ridge.
In respect of other param eters the ridges resem ble each other (Table 12).
The articular cartilage o f the proxim al trochlea of the tibial tarsal is
1.6 mm thick in the youngest groups and 0.3 mm in the oldest ones.
The tarsocrural joint co-form s the calcaneom alleolar joint, situated
laterallly, w hich in its general outlines is similar to this joint in dom estic
cattle.
P roxim al intertarsal joint. The ridges o f the distal trochlea of the
tibial tarsal bone have smaller chords and radii but larger angles of
curvature than have those of the proxim al trochlea of this bone (Table
12).
The articular cartilage of the distal trochlea passes on to the posterior
surface o f the tibial tarsal, w here it participates in the form ation of the
proxim al talocalcaneal joint. It also passes from the cartilage of the
distal talocalcaneal joint. In the place w here this cartilage passes on to
the posterior surface o f the bone there is a noticeable synovial fossa
(Fig. 19 B— x), the developm ent and m orphological characters of w hich
are sim ilar to those described for the synovial fossa of the cochlea of
the tibia. Its length is 4 mm and the w idth 3 mm in the youngest bisons
and, respectively, 32 and 8 mm in the oldest specimens.
M edially, the proxim al articular surface of the fused central and fourth
tarsal bones (Tc + T IV) is the negative im pression of the distal trochlea
o f the tibial tarsal only that the angle o f its curvature is smaller than
the angle of the curvature o f the trochlear ridges by about 95°. Thus,
theoretically this differen ce is the flexion -exten sion range of m ovem ents
in the proxim al intertarsal joint. The synovial fossa on the proxim al
articular surface of the Tc + T I V (Fig. 19 C— x) is a remnant of the
syndesmosis betw een the central and fourth tarsal bones, w hich occurs
in foetuses and newborns. The cartilage in the proxim al intertarsal joint
is 1.6 m m thick in the youngest specim ens and 0.3 mm in the oldest
ones.
The distal intertarsal and tarsometatarsal joints are plane joints, re­
sembling those of dom estic cattle in structure. Their articular cartilage
Joints of hind-limbs of the European bison
71
is about 1.3 mm thick in the youngest European bisons and 0.2— 0.3 mm
in the oldest.
Intratarsal joints. The proxim al and distal talocalcaneal joints are of
the greatest functional-m otor im portance. The cavity o f the proxim al
talocalcaneal joint between the posterior surface of the tibial tarsal and
the adjoining surface of the fibular tarsal form s a perfect arc, the angular
value o f w hich is 75° in all European bisons. The gliding fibular tarsal
bone m oves over a distance having an angular value of 30°. The distal
talocalcaneal joint is an arthrodia.
The sides and the plantar surface of the capsule of the hock joint
is made of a fibrous membrane w hich here develops strong colateral and
plantar ligaments. O nly on the anterior side and in the angle form ed
by the tibia and the body of the fibular tarsal bone the fibrous m embrane
is thin and relatively loose.
This fibrous m em brane is com m on to all the tarsal joints mentioned.
The synovial m em brane is attached to the margins of the articular sur­
faces of the respective bone com ponents. M oreover, it wraps the inter­
osseous ligaments and the long peroneal m uscle tendon, w hich runs
transversely in the cavity of the tarsometatarsal joint towards its inser­
tion. The synovial membrane, in addition, encloses the artery supplying
the tibial tarsal bone and extending in the cavity of the proxim al talo­
calcaneal joint.
On the anterior side the capsule of the hock joint form s tw o swellings.
The lateral swelling (Fig. 24— 1) is visible between the lateral digital
extensor tendon (Fig. 24— 51) and the long digital extensor tendon
(Fig. 24— 50). The smaller medial sw elling (Figs. 22, 24— 2) is situated
betw een the third peroneal m uscle tendon (Figs. 22, 24— 47) and the
anterior edge o f the medial collateral ligam ent (Figs. 22, 24— 12— 13).
In the angle form ed by the tibia and the b od y of the fibular tarsal
the capsule of the hock joint pouches halfw ay up the body of the fibular
tarsal (Fig. 22— 3). The synovial m em brane of the pouch bears villi.
The synovial m em brane of the hock joint com prises 5 articular cavities.
The first and roomiest o f them is com m on to the tarsocrural, proxim al
intertarsal, and proxim al and distal talocalcaneal joints. The second cavity
is that o f the distal intertarsal joint. The tarsometatarsal joint has the
next three cavities. One of them occurs betw een the first tarsal and
the fused third and fourth metatarsal bones and the other two, separated
from each other by the canal of the long peroneal muscle, belong to the
remaining part of the tarsometatarsal joint.
The cavity of the joint between TI and MtlII + I V has not a com m un­
ication w ith the joint form ed by the vestigial M tl l and MtHI + IV,
although their close vicinity w ould suggest its presence.
72
F . K o b ry n cz u k
The greatest capacity, and so measurable, is that of the first joint cavity.
In foetuses and new borns it is 24— 36 ml, in calves up to one year of age
35— 45 ml, in adult males 50— 60 m l and in adult fem ales about 50 ml.
The other fou r joint cavities are fisures, the capacity of w hich does not
exceed 1 ml each in adults.
Ligam ents of the H ock Joint
Lateral side. In European bisons the lateral collateral ligam ent (Figs.
20, 24, 25— 4— 8) is divided into 5 parts:
1. The tibiom etatarsal part (Fig. 20— 4) arises from the posterolateral
surface of the distal end of the tibia and arches downwards. On its way
it is attached to the m alleolus of the fibular tarsal and Tc + TI V and
terminates on MtIII + IV. The insertion tendon of the long peroneal muscle
(Figs. 20, 24— 48) penetrates under the anterior border of the tibiom e­
tatarsal part at the height of the tarsometatarsal joint. The anterior border
of this part overlaps the calcaneom etatarsal part of the ligament under
discussion (Fig. 20— 6) in adulth bisons, w ith w hich it m ay be coalesced.
2. The calcaneom alleolar part (Fig. 20— 5) is another, deeper lying,
portion o f this ligament. It arises anteriorly to the attachm ent of the
previous part on the m alleolus and above it on the tibia. It ends attached
to a special rough area on the lateral side of the fibular tarsal bone.
3. The calcaneom etatarsal part (Fig. 20— 6) begins on the lateral
surface of the fibular tarsal bone, below the insertion of the calcaneo­
m alleolar part, and ends on MtIII + IV, posteriorly to the tibiometatarsal
part.
4. The tibiocalcaneal part (Figs. 20, 24, 25— 7) is the third, counting
from the outer side, layer o f the ligam ent. It arises anteriorly on the
tibia, below the proxim al annular ligam ent (Figs. 20, 22, 24— 35) and
has a multiplanar spiral course. It runs inferolateroposteriorly and is
attached to the fibular tarsal bone, above the calcaneom alleolar part.
5. The talom alleolar part (Fig. 20— 8) originates at the posterior edge
of the m alleolus and runs the shortest w ay to the fossa for attachment
of the ligam ent on this side of the proxim al trochlea of the tibial tarsal
bone. The dimensions of individual parts of the lateral collateral liga­
ment are given in Table 13.
The lateral talometatarsal ligam ent (Fig. 21, 25— 9) begins in its fossa
for attachm ent on the lateral side of the distal trochlea of the tibial
tarsal and ends on MtIII + IV. In old European bisons it grow s together
with the calcaneom etatarsal ligament, w hich is posterior to it. This last
ligament (Fig. 21, 25— 10) arises on the coracoid process of the fibular
tarsal bone and term inates on the anterolateral aspect of MtIII + IV.
The lateral ligam ent connecting Tc + TI V w ith M t I I I + I V (Fig. 21— 11)
Table 13
D im ensions o f lateral collatera l ligam ent o f h ock jo in t (in mm ).
L — lenght, W — w idth, T — thickness.
n
T ibiom etatarsal
pa rt
L
W
C alcan eom alleolar
part
T
L
W
T
C alcaneom etatarsal
pa rt
L
T ibiocalcan ean
part
W
T
L
W
T
8.8
8.7
12.4
13.3
13.9
13.6
1.54
1.6
1.7
3.2
3.5
3.8
4.0
2.50
35.0
44.3
55.0
61.3
61.1
65.1
1.86
10.5
10.1
14.6
15.2
15.1
15.4
1.47
1.3
2.3
3.0
3.6
4.6
4.5
3.46
8.1
8.8
10.0
13.5
12.8
12.4
1.54
1.4
1.8
2.7
2.5
3.0
3.3
2.36
33.7
45.0
49.2
55.0
58.0
57.0
1.69
9.9
11.0
12.8
12.0
13.7
14.2
1.43
1.2
2.5
2.7
3.5
3.8
3.9
3.25
M ales
8
5
11
8
8
7
67.3
94.0
105.1
113.0
116.6
117.4
1.74
7.1
6.6
9.6
12.4
12.5
12.3
1.74
1.1
1.6
2.6
3.6
4.3
4.8
4.36
38.8
40.7
55.9
54.1
55.8
53.4
1.40
5.7
9.5
10.6
12.3
12.3
12.3
2.16
1.1
1.3
2.0
2.9
3.4
4.0
3.64
66.2
72.0
98.5
113.0
107.7
112.8
1.70
5.8
6.2
8.0
9.6
10.5
10.8
1.86
1.1
1.4
2.0
2.8
3.4
3.7
3.37
36.8
38.0
42.0
51.0
51.0
50.8
1.38
52
6.6
8.7
9.0
10.7
10.8
2.10
1.0
1.7
1.4
2.0
2.9
3.6
3.60
42.7
57.0
74.0
76.8
77.4
77.3
1.82
Fem ales
4
1
6
1
3
9
37.5
48.0
54.5
60.0
63.0
72.6
1.95
o
CM
CD
fO
OJ
ID
LOCO
00
CO
CD
lO
Fig. 20. L eft hock join t w ith its capsule partly rem oved. Lateral view . Superficial
la yer (14 -yea r-old male).
Fig. 21. Right
hock
jo in t w ith
its
capsule rem oved.
(8 -y e a r-o ld male).
Lateral
view .
D eep
layer
rO
Fig.
Fig. 22. Right h ock joint. M edial view . S u perficial layer (1 4 -yea r-old male).
23. Right h ock join t w ith its capsule rem oved. M edial v iew . D eep la yer
(8 -y e a r-o ld m ale).
ю
CM
Joints of hind-limbs of the European bison
77
is divided into tw o parts, an anterior and a posterior, separated b y the
tendon of the long peroneal m uscle (Fig. 21— 48), w hich squeezes in
betw een them.
Medial side. The m edial collateral ligam ent (Figs. 22, 23, 24— 12— 14)
splits into three parts in European bisons:
1. The part connecting the tibia w ith the fibular tarsal and Tc + TI V
(pars tibiocalcaneocentroquartalis — Figs. 22, 24— 12) has the shape of
a trapezium w ith its longer (distal) base against the long plantar ligam ent
(Fig. 22— 29). This part begins on the anterom edial surface of the distal
end of the tibia and terminates on the sustentaculum of the fibular tarsal
bone and Tc + TZV, exchanging fibres w ith the long plantar ligam ent.
2. The tibiometatarsal part (Figs. 22, 24— 13) extends obliqu ely under
the previous part. It arises from the distal end of the tibia and on its
w ay is attached in turn to Tc + TIV, TII + TIII and TI to end on
Its anterior border and the medial talometatarsal ligam ent (Figs. 22, 23,
24, 25— 15) coalesce by the m edium of loose fibres, and the posterior
border blends w ith the long plantar ligam ent (Fig. 22— 29).
3. The tibiotalar part, (Figs. 22, 23— 14) is the third and deepest portion
o f the ligam ent concerned. It begins on the posterior edge of a special
Fig. 24. L eft
Fig. 25. Right
hock joint. A n terior
view . S u p erficia l layer (1 4 -y ea r-old m ale).
h ock join t w ith its capsule rem oved. A n terior v iew . D eep layer
(14 -yea r-old male).
Fig. 26. Right hock join t w ith its capsule rem oved. P osterior (plantar) view . D eep
la yer (8 -y ea r-old male).
(In Figs. 22 and 24 the join t capsules are filled w ith gelatin). 1, 2 — lateral and
m edial sw ellings o f join t capsule, 3 — pou ch o f jo in t capsule, 4— 8 — lateral
collatera l ligam ent,
4 — tibiom etatarsal part, 5 calcan eom a lleola r part, 6 — ca lca neom etatarsal part, 7 — tibiocalcanean part, 8
— ta lom a lleolar part, 9 — lateral
talom etatarsal ligam ent, 10 — calcaneom etatarsal ligam ent, 11 — lateral cen troquartom etatarsal ligam ent, 12— 14 — m edial collatera l ligam ent, 12 — tib ioca lca n eocen troqu artal part, 13 — tibiom etatarsal part, 14 — tibiotalar part, 15 — m edial
talom etatarsal ligam ent, 16— 21 — ligam ents o f the m ed ial side con n ectin g: 16 —
tibia w ith Tc + IV ,
17 — Tc + T IV w ith TI, 18— T c + T IV w ith T II + TIII, 19 —
T I w ith TII + TIII, 20— TI w ith M t I I I + I V , 21 — T I I + T I I I , 22— 27 — ligam ents
o f the an terior side con n ectin g: 22 — Tt w ith Tc + TIV , 23 — Tt w ith M tIII + IV, 24 —
T c + T IV w ith M tIII + IV, 25 — TII + TIII w ith T c + IV, (intratarsal), 26 — T II + TIII
w ith Tc + T IV (intertarsal), 27 — TII + TIII w ith M tIII + IV, 28, 29 — lateral and
m edial parts o f long plantar ligam ent, 30— 33 — ligam ents o f the p lantar side
con n ectin g: 30 — T f w ith Tc + TIV, 31 — Tc + T IV w ith TI, 32 — T c + T IV w ith
M tIII+ IV , 34 — interosseous ta localcan eal ligam ent, 35, 36 — p ro x im a l and distal
annular ligam ents, 37 — tibia, 38 — m alleolus, 39 — fib u la r tarsal bone, 40 — tibial
tarsal bone, 41— Tc + TIV, 42 — TI, 43 — TII + TIII, 44 — M tIII + IV , 45 — su rface
fo r articulation
w ith M ill, 46 — tarsal canal, 47 — ten don o f th ird p eron ea l m uscle,
48 — ten don o f lon g peroneal m uscle, 49 — ten don o f ex ten sor m u scle o f digit III,
50 — ten don o f long exten sor m uscle o f digits, 51 — tendon o f ex ten sor m u scle o f
digit IV, 52 — tendon o f su perficial fle x o r m uscle o f digits, 53 — ten don o f deep
fle x o r m u scle o f digits, 54 — ten don o f lon g fle x o r m u scle o f great toe and posterior
tibial m uscle, 55 — tendon o f lon g fle x o r m u scle o f digits, 56 — short ex ten sor
m u scle o f digits, 57 — m iddle interosseous m uscle, 58 — an terior tibial m uscle,
59 — lateral plantar vein.
F. Kobryñczuk
78
process of the tibia, and ends in the fossa for attachment on this side
of the proxim al trochlea of the tibial tarsal bone. The dim ensions of the
medial collateral ligam ent are included in Table 14.
The m edial talometatarsal ligam ent (Figs. 22, 23, 24, 25— 15) is an
autonom ous tape-like structure. It arises from its fossa for attachm ent
on this side of the distal trochlea of the tibial tarsal bone and ends on
MtIII + IV.
A fter the rem oval o f the ligaments described above, the ligaments that
connect the follow in g bones appear on the m edial side: the tibia with
Tc + T /V (Fig. 23— 16), Tc + T iV with TI (Figs. 23, 26— 17), T c + T I V with
T ab le 14
D im ensions o f m ed ial collatera l ligam ent o f h ock join t (in m m ).
L — length, W — w idth, T — thickness
G roup
n
Part con n ectin g tibia
w ith T c + T I V and T f
W idth
p rox.
dist.
T
L
T ibiom etatarsal
part
T ib iota lar part
L
W
T
L
W
T
52.8
75.3
86.3
82.2
83.7
85.6
1.62
5.4
5.5
7.5
8.0
9.0
9.3
1.72
1.9
1.9
2.7
3.6
3.7
3.9
2.05
18.8
17.7
29.2
33.3
32.0
31.3
1.67
11.6
11.0
13.1
14.3
13.9
14.0
1.21
1.1
2.2
2.7
3.1
4.1
4.1
3.73
5.3
6.0
7.2
7.5
7.2
8.0
1.50
1.8
2.3
2.2
2.4
3.1
3.0
1.67
16.0
16.0
25.4
26.0
27.3
27.8
1.74
9.7
11.0
13.8
12.0
12.0
12.6
1.24
1.0
2.0
2.2
2.0
2.5
3.0
3.00
M ales
0
I
II
III
IV
V
G. C.
8
5
11
8
8
7
51.0
62.0
70.0
74.5
79.7
83.6
1.64
9.0
11.3
14.3
18.4
19.0
18.7
2.08
26.5
40.0
47.3
48.1
44.7
47.8
1.84
0
I
II
III
IV
V
G. C.
4
1
6
1
3
9
46.8
56.0
66.0
80.0
70.3
76.0
1.62
8.7
9.0
12.1
14.0
14.7
15.8
1.82
23.8
35.0
42.5
40.0
44.7
41.9
1.76
2.1
2.7
3.8
5.6
8.0
8.0
3.81
Fem ales
1.7
2.0
3.6
4.8
6.8
6.1
3.59
47.7
60.0
67.5
65.0
65.3
73.9
1.55
TII + TIII (Fig. 23— 18), TI with TII + TIII (Fig. 23— 19), TI w ith
MtIII + I V (Figs. 23, 26— 20), TII + TIII w ith MtIII + IV (Fig. 23— 21).
A nterior side. The rem oval of the short extensor m uscle of the digits
in European bisons (Figs. 20, 22, 24— 56) uncovers the poorly developed
dorsal tarsal ligaments connecting the follow in g bones: Tt w ith Tc + TIV
(Fig. 25— 22), Tt w ith MtIII + I V (Fig. 25— 23), Tc + TIV w ith M t I I I + I V
(Fig. 25— 24), TII + TIII w ith Tc + IV (intratarsal — Fig. 25— 25), TII + TIII
with Tc + TI V (intertarsal — Fig. 25— 26), TII + TIII w ith MtIII + IV (Fig.
25— 27).
Joints of hind-limbs of the European bison
79
Posterior (plantar) side. The long plantar ligam ent consists of a lateral
and a m edial part joined b y a m iddle sheet. These three elements rem ain
distinct only in foetuses and newborns, w hereas in adult European bisons
they exchange fibres w ith each other.
The lateral part (Fig. 20— 28) begins over the w hole length of the
posterolateral border of the fibular tarsal bone, being initially united
with the m iddle sheet. Further towards the digits it separates from this
last and at the height of the sustentaculum tali is divided into an anterior
and a posterior portion. The anterior portion is attached to the postero­
lateral surface o f T c + T /V , its anterior m argin being grow n together w ith
the calcaneom etatarsal part of the lateral collateral ligam ent and term i­
nates on MtIII + I V between the attachm ent of this ligament and the
middle interosseous muscle, running in this place. The posterior portion
runs perpendicularly, is attached to Tc + T /V and a special tubercle on
this side of the proxim al end of MtIII + I V and next merges w ith the
superficial part of the m iddle interosseous muscle.
The medial part of the long plantar ligam ent (Fig. 22— 29), m uch
stronger than the lateral part, passes from the tuberosity and body of
the fibular tarsal bone on to the posterom edial border of the sustentacu­
lum tali and next on to Tc + TIV, thus form ing two bridges over the
passages for appropriate tendons of the deep flex or m uscle of the digits.
In its further course the medial part splits into an anterior and a poster­
ior portion. The anterior portion runs on to the posterom edial surface
of Tc + T /V , w here it is attached, next reaches TII + TIII and ends on
this side on MtIII + IV. At the height o f Tc + TZV the posterior portion
is divided into an anterior and a posterior branch, the first of w hich ends
on M t /// + /V . The posterior branch behaves analogously to the posterior
portion of the lateral part of the ligam ent being described.
The m iddle sheet becom es thinner in feriorly and ends in a sharp sem i­
lunar m argin, from under w hich the tendon o f the deep flexor m uscle
of the digits em erges. The tendon of the superficial flex or muscle of the
digits is contiguous to the posterior surface of this sheet.
The rem oval o f the long plantar ligam ent and m iddle interosseous
muscle detects the ligam ents connecting the follow ing bones on the plantar
side of the hock joint: Tf with Tc + T /V (Fig. 26— 30), Tc + T /V w ith Tl
(Fig. 26— 31), T c + T I V w ith MtIII + IV (Fig. 26— 32), Tl with MtIII + IV
(Fig. 26— 33).
Interosseous Ligaments of the Hock Joint
The interosseous talocalcaneal ligam ent (Fig. 25— 34) of European bi­
sons is a strong structure extending transversely in the tarsal sinus.
Table 15
D im ensions o f articular surfaces o f m edial m etatarsophalangeal join t
Curvature
Length
(chord)
n
r
s
V
*1
x
2.8
3.4
1.9
2.0
1.8
1.7
12.2
10.8
4.7
5.0
4.3
4.1
11.9
16.3
20.6
20.7
21.9
21.5
1.81
211
212
204
209
213
207
0.98
11.0
17.3
18.0
20.0
19.6
19.4
1.76
212
215
200
211
205
208
0.98
0
I
II
III
IV
V
G.C.
8
5
16
17
15
13
23.0
31.4
40.3
40.1
41.9
41.9
1.82
0
1
II
III
IV
V
G.C.
4
1
9
6
7
18
21.2
33.0
37.5
38.5
38.3
37.6
1.77
Length (chord)
A n gle
x
v
R ange
o f m ove
m ents
P roxim a l articular su rface
of p rox im a l ph alanx
M edial trochlea o f M t I I I + I V
G roup
(in m m and degrees).
s
M ales
4.5
2.1
1.9
4.1
2.0
4.0
2.7
5.7
2.8
5.9
3.6
1.7
x
s
21.0
21.6
29.7
30.5
31.5
31.7
1.51
1.2
1.7
1.7
1.2
0.0
1.0
V
5.7
7.9
5.7
3.9
0.0
3.2
A n g le
x
87
83
93
95
92
94
1.08
v
s
4.3
4.9
5.8
4.0
5.6
5.2
4.9
5.9
6.2
4.2
6.1
5.5
Xi— X2
X
124
129
111
114
121
114
0.91
Fem ales
2.1
2.3
1.4
2.6
5.6
6.0
3.6
6.9
4.2
4.8
4.2
5.1
2.1
2.3
2.0
2.4
19.4
21.0
25.8
28.3
27.9
27.4
1.41
1.6
1.4
1.0
2.4
6.2
4.9
3.6
8.8
82
£6
86
92
91
89
1.08
3.7
4.3
1.5
3.7
4.3
4.7
1.6
4.2
130
129
114
118
114
119
0.90
Joints of hind-limbs of the European bison
81
The other interosseous ligaments are small structures connecting the
follow ing bones: Tc + TIV w ith MtHI + IV, Tc + TIV with TII + TIII,
TII-{-Till w ith TI, TII + TIII w ith MtHI + IV and TI with MtHI + IV.
6. Metatarsophalangeal Joints
The structure of tw o trochleae o f the metatarsal bones is similar. The
dimensions of the ridges of the medial trochlea are given in Table 15.
Fig. 27. System o f exten din g forces o f h ock joint.
F — fo r ce o f exten sors o f h ock join t, O — axis o f rotation o f tarsocru ral joint,
O t — axis o f rotation o f p rox im a l intertarsal joint, p(A O ) — arm o f extending
fo r ce traced fro m centre O, q(C O t) — arm o f extending fo rce traced fro m centre
O u a — angle o f in clination o f lon g axis o f tibial tarsal bone to arm s p and q.
The articular cartilage w hich covers the articular surfaces of the tro­
chleae is thin, 1.6— 1.9 mm in the youngest specimens of both sexes and
0.2— 0.5 m m in adults. Covering the working articular surfaces of the
trochleae, it also passes on to their axial and abaxial surfaces, occupying
small crescent areas.
82
F. Kobrynczuk
The proxim al articular surfaces of the proxim al phalanges are negative
copies of the trochleae of the metatarsal bones, but the angles of their
curvatures are smaller than those in the trochleae (Table 15). This d iffe ­
rence (Ti— oc2) is at the same time the flexion -exten sion range of m ov e­
ments in the two metatarsophalangeal joints (Table 15).
The thickness of the articular cartilage on the articular surfaces of the
proxim al phalanges is similar to that of the cartilage of the trochleae.
In structure and arrangem ent the sesamoid bones of the proxim al
phalanges resem ble those of dom estic cattle.
In European bisons either metatarsophalangeal joint has its ow n joint
capsule, but both capsules have their fibrous m em brane in com m on. The
attachment of this m em brane coincides w ith the margins of the articular
surfaces o f particular bone com ponents of the joints, except fo r the dorsal
surface, w here it is m oved aw ay to the outside from these edges by about
1 cm. The attachm ent of the synovial m em brane presents itself sim ilarly.
The capsule of either of these joints form s tw o pouches. The dorsal
pouch (Fig. 28— 1) extends above the appropriate trochlea of the m eta­
tarsal bones and is 4 cm long in adult bisons. The plantar pouch (Fig.
28— 2) squeezes in upwards betw een the portions of the deep part of the
m iddle interosseous m uscle and the metatarsal bones, being 3 cm long
in the youngest specimens and 7— 8 cm in adults of both sexes.
The synovial membranes o f both joints cling to each other in a sagittal
plane and form a joint partition. In all European bisons this partition
has an oval aperture, by w hich the cavities of these joints com m unicate
with each other. The total capacity of both these cavities is 17 mi in
males of group 0, 15 m l in fem ales of this group and up to 20 ml in
calves of both sexes up to 1 year of age. In adult males it ranges from
35 to 45 ml and in adult fem ales from 30 to 35 ml. The amount of sy ­
novial fluid obtained from both these cavities reaches 3 ml in adults.
Ligaments of the Metatarsophalangeal Joints
The abaxial collateral ligam ent (Fig. 30— 3) of European bisons consists
of two portions, a superficial and a deep, w hich cross at an acute angle.
The axial collateral ligam ent (Fig. 31— 4) is situated in the intertrochlear
incisure of the metatarsal bones. It is in the form of a distally broadening
lamina. Betw een the axial collateral ligam ents of the tw o metatarsopha­
langeal joints there is a fissural space, through w hich the interdigital
branch of the m iddle interosseous m uscle passes (Figs. 31, 32— 13).
Sesam oidean Ligaments
Proxim al seamoidean ligaments. The proxim al sesamoidean ligaments
Joints of hind-limbs of the European bison
83
are appropriate portions of the m iddle interosseous muscle. In European
bisons this m uscle divides halfw ay dow n the metatarsus into a superficial
and a deep part. The superficial part (Figs. 28, 30— 5) is the dow nw ard
extension of the fibres of the long plantar ligam ent and unlike the deep
part has no m uscle fibres. It splits into two portions, a lateral and a
medial, w hich w ith the corresponding branches of the superficial flexor
muscle of the digits form tw o tendinous sheaths (Figs. 30, 31, 32— 6),
w hich em brace the appropriate tendons of the deep digital flexor. The
deep of the m iddle interosseous m uscle (Figs. 29, 30, 31, 32— 7— 23) un­
dergoes num erous divisions on its w ay down. Thus, at the distal third
of the metatarsus it divides into lateral and medial abaxial portions
(Figs. 30, 32— 8, 9) and a m iddle sheet (Fig. 32— 10). The sheet, in turn,
splits into three branches, tw o of w hich are axial, a lateral (Fig. 32— 11)
and a m edial (Figs. 31, 32— 12), and the third — the interdigital branch
(Figs. 31, 32— 13) — is unpaired and lies between the axial branches.
The abaxial portions of the deep part o f the m iddle interosseous m uscle
reach the abaxial and, partly, axial sesamoid bones. A bove the sesamoid
bones they send o ff the lateral and medial abaxial branches (Figs. 29,
30, 32— 14, 15) to the external surfaces of the joints. These branches
reach the dorsal surfaces of the m iddle phalanges, where most of their
fibres term inate together w ith the medial and lateral digital extensor
tendons. The other fibres of these branches, as the lateral and medial
bands of the distal phalanges (Figs. 29, 30— 16, 17), disappear in their
periosteum. A t the height o f the metatarsophalangeal joints the abaxial
portions give o ff the lateral and medial sesamoid bands (Figs. 30, 32— 18,
19) to the external edges of the abaxial sesamoid bones.
The axial portions of the deep part of the muscle under study reach
the axial and, externally, also abaxial sesamoid bones.
The interdigital branch of the muscle sheet goes under the interdigital
intersesamoidean ligament (Figs. 31, 32— 26) in the intertrochlear incisure,
w here it undergoes a strong lateral flattening. A t this height it sends
o ff some of its fibres (Fig. 31— 20) to the interdigital intersesamoidean
ligam ent (Figs. 31, 32— 26) and another bundle (Fig. 31— 21) to the pro­
xim al interdigital ligam ent (Fig. 31— 39). A fter sending o ff this last bun­
dle of fibres, the interdigital branch divides into the lateral and medial
axial twigs (Figs. 29, 31, 32— 22, 23). These twigs pass on to the dorsal
surfaces of the toes and end on the m iddle phalanges together w ith the
corresponding medial and lateral digital extensors.
On the surfaces o f the non-divided deep part of the m iddle interosseous
m uscle and on its longitudinal and transverse sections one can see muscle
and on its longitudinal and transverse sections one can see m uscle fibres
em bedded in the fibrous structure of this muscle. The num ber of muscle
Fig. 28. Sagittal section through digit IV o f right h in d -lim b . Joint capsules o f digits
filled w ith gelatin (black) (8 -y e a r -o ld male).
Fig. 29. D igits o f right h in d -lim b . A n terior v iew . (1 6 -m on th -old male).
E xplanation o f Figs. 28— 32.
1, 2 — dorsal and plantar pou ch es o f capsule o f m etatarsophalangeal join t, 3, 4 —
abaxial and axial collateral ligam ents o f m etatarsophalangeal join t, 5— 23 — m iddle
interosseous m uscle, 5 — su perficial part, 6 — tendinous sheath o f deep fle x o r
m uscle o f digits, 7 — deep part, 8, 9 — lateral and m edial ab axial portions, 10 —
m iddle sheet, 11, 12, 13 — lateral, m ed ial and interdigital a xia l branches, 14, 15 —
lateral and m edial abaxial branches, 16, 17 — lateral and m edial bands fo r distal
phalanx, 18, 19 — lateral and m edial sesam oidean bands, 20, 21 — fib res o f in ter­
digital bran ch to interdigital intersesam oidean and p roxim a l interdigital ligam ents,
22, 23 — lateral and m edial a xia l tw igs, 24— 26 — lateral, m ed ial and interdigital
intersesam oidean ligam ents, 27, 28 — lateral and m edial collatera l sesam oidean
ligam ents, 29, 30 — lateral and m edial abaxial obliqu e sesam oidean ligam ents,
31, 32 — lateral and m edial axial obliqu e sesam oidean ligam ents, 33, 34 — lateral
and m edial abaxial cru ciate sesam oidean ligam ents, 35, 36 — lateral and m edial
axial cru ciate sesam oidean ligam ents, 37, 38 — lateral and m edial phalangosesa-
Joints of hind-limbs of the European bison
85
Fig. 30. Joints o f digit IV in left hind-lim b. A x ia l v iew (1 0 -y ea r-old male).
m oidean ligam ents, 39 — p rox im a l interdigital ligam ent, 40, 41 — dorsal and plantar
pouches o f capsule o f p rox im a l interphalangeal joint, 42, 43 — ab ax ial and axial
collateral ligam ents o f p roxim a l interphalangeal joint, 44 — ax ia l ligam ent o f
p roxim a l and distal interphalangeal joints, 45— 47— axial, ab axial and central
plantar ligam ents, 48, 49 — dorsal and plantar pouches o f capsule o f distal in ter­
phalangeal join t, 50, 51 — abaxial and axial collateral ligam ents o f distal in ter­
phalangeal join t, 52— 55 — abaxial and axial ligam ents attaching sesam oid bone
o f distal ph ala n x to m iddle and distal phalanges, 56— 61 — distal interdigital
ligam ent, 56, 57 — lateral and m edial p rox im a l branches, 58, 59 — lateral and
m edial m iddle branches, 60, 61 — lateral and m edial distal branches, 62, 63 —
p rox im a l and distal annular ligam ent, 64 — m etatarsal bones I I I + I V ( M t I I I + I V ) ,
65 — p rox im a l phalanx, 66 — sesam oidean bon e o f p ro x im a l p h ala n x, 67, 68 —
m iddle and distal phalanges, 69 — sesam oidean bone o f distal ph alanx, 70 — tendon
o f lon g exten sor m uscle o f digits, 71 — ten don o f exten sor m u scle o f digit IV,
72 — tendon o f exten sor m u scle o f digit III, 73 — tendon o f su p erficial fle x o r
m uscle o f digits, 74 — its portion fo r digit III, 75 — tendon o f deep fle x o r m uscle
o f digits, 76, 77 — its portion digits IV and III.
86
F . K o b ry n cz u k
32
Fig. 31. Joints o f digit IV in right h in d -lim b . A b a x ia l v iew (1 0 -yea r-old male).
Fig. 32. Joints o f digits in left h in d -lim b w ith join t capsules and p roxim a l digital
ligam ent rem oved (1 0 -yea r-old male).
E xplanations see pages 84 and 85.
fibres decreases considerably with age. In young specimens they are
particularly distinct in the distal part of the muscle.
M iddle sesamoidean ligaments. The m edial and lateral intersesamoidean
ligaments (Fig. 32— 24, 25) connect the abaxial sesamoid bones to the
axial ones in the corresponding digit. Im pregnated w ith cartilaginous
tissue, these ligaments constitute the bottom s of the gliding grooves for
two sheaths of the deep digital fle x o r tendons.
The interdigital intersesamoidean ligam ent (Figs. 31, 32— 26) connects
the tw o axial sesamoid bones. It has, in addition, a connection with the
interdigital branch of the m iddle interosseous m uscle (Fig. 31— 20).
Joints of hind-limbs of the European bison
87
The lateral and medial collateral sesamoidean ligaments (Figs. 30, 32—
— 27, 28) connect the external edges of the abaxial sesamoid bones with
the tubercles for ligaments on the proxim al phalanges and also by means
of thin strips (not shown in the figures) w ith the abaxial surfaces of the
tw o trochleae of the metatarsal bones. In some specimens synovial bursae
w ere found under the distal attachments of these ligaments.
Distal sesamoidean ligaments. The m edial and lateral abaxial oblique
sesamoidean ligam ents (Figs. 30, 32— 29, 30) and the medial and lateral
axial oblique sesamoidean ligaments (Fig. 32— 31, 32) arise one from
each sesamoid bone and run dow nw ard to the proxim al phalanges. In
European bisons they may be regarded as extensions of som e fibres of
the axial and abaxial portions of the m iddle interosseous muscle.
The lateral and medial abaxial cruciate sesamoidean ligaments (Fig.
32— 33, 34) and the lateral and medial axial cruciate sesamoidean liga­
ments (Fig. 32— 35, 36) begin on the distal edges of the sesamoid bones
corresponding to their name, below w hich bones they cross in pairs to
end on the plantar surfaces of the proxim al phalanges.
The lateral and medial phalangosesamoidean ligaments (Fig. 32— 37,
38) arise from the distal edges of the axial sesamoid bones, below which
they cross. The lateral ligament terminates on the proxim al phalanx of
digit III and the m edial one on the proxim al phalanx of digit TV.
The proxim al interdigital ligament (Fig. 31— 39) is a conspicuous bundle
of fibres w hich run transversely betw een the surfaces of the proxim al
phalanges facing each other. Some fibres of this ligam ent reach the axial
plantar ligam ents of the proxim al interphalangeal joints as w ell as the
distal interdigital ligament. On the other hand, the proxim al interdigital
ligament receives fibres from the interdigital branch of the m iddle in­
terosseous m uscle (Fig. 31— 21).
7. Proxim al Interphalangeal Joints
Structurally these two joints are autonom ous. They are separated from
each other by the fasciae and loose tissue that penetrate into the inter­
digital space.
The articular surface of the trochlea o f the proxim al phalanx has the
shape of a heart whose tip is bent forw ard and slightly upward.
The articular fovea of the m iddle phalanx is an im perfect negative
copy of the trochlea of the proxim al phalanx.
The articular cartilage covering both articular surfaces is 1.4— 1.6 mm
thick in foetuses and newborns, 0.5— 0.8 mm in calves up to 1 year of
age, and 0.2— 0.3 mm in adults.
The attachm ents of the joint capsule coincide with the margins of the
articular surfaces except the dorsal surface, w here the line of attachment
88
F. Kobrynczuk
is m oved somewhat upwards. The joint capsule pouches upwards (dorsal
pouch — Fig. 28— 40), squeezing in under the tendon of the extensor
muscle of digit III or IV. The axial pouch sinks as far as the distal interphalangeal joint on the axial side of the digit. The room iest plantar
pouch (Fig. 28— 41) extends halw ay up the proxim al phalanx. In spite
of these pouches the cavity of the joint discussed is small. Its capacity
is 1.5— 2.0 m l in group 0 of both sexes, 3.0— 4.5 ml in calves up to 1 year
of age, 6.6 ml in adult males and 4.6 m l in adult fem ales.
Ligaments o f the Proxim al Interphalangeal Joints
The abaxial collateral ligam ent (Fig. 30— 42) arises w ith the proxim al
portion o f the distal interdigital ligam ent from a fossa for attachment
on the proxim al phalanx (Figs. 30, 32— 56, 57) and ends at a tubercle
for attachm ent on the m iddle phalanx.
«
The axial collateral ligam ent (Fig. 31— 43) much stronger than the
previous one, begins in a fossa for attachm ent on the proxim al phalanx
and ends on a tubercle for attachm ent on the m iddle phalanx. In its distal
section the posterior portion of fibres intertw ine w ith the fibres of the
m iddle portion of the distal interdigital ligament (Figs. 31, 32— 58, 59).
Small synovial bursae lie under the proxim al and distal attachments of
this ligament.
The axial collateral ligam ent of the proxim al and distal interphalangeal
joints (Fig. 31— 44) extends over tw o joint. It begins together w ith the
previous ligam ent and terminates on the axial surface of the distal pha­
lanx. The axial pouch of the capsule of the proxim al interphalageal joint
is squeezed in under this ligament.
The plantar ligaments form a Y-shaped structure. Its paired parts are
the axial and abaxial plantar ligam ents (Figs. 30, 31, 32— 45, 46) and the
central plantar ligam ent constitutes the unpaired part (Figs. 28, 32— 47).
As a w hole, the plantar ligam ents are strong structures. Situated on the
plantar side of the joint, they inhibit its hyperextension.
8. Distal Interphalangeal Joints
Apart from the com m on structure, w hich is the distal interdigital li­
gament, these joints are autonomous.
The articular surface of the trochlea of the m iddle phalanx is an irre­
gular sem i-cylinder. The gentle groove o f the trochlea runs obliquely
and consequently the trochlea resem bles a screw. The angle of the cur­
vature of the articular surface of the distal phalanx is greater than that
of the curvature of the trochlea on the m iddle phalanx.
The articular cartilage is 1.8 mm thick in foetuses and newborns and
ranges between 0.3 and 0.4 mm in adults.
Joints of hind-limbs of the European bison
89
The dorsal pouch of the capsule of the distal interphalangeal joint
(Fig. 28— 48) extends upwards above the coronary border of the hoof for
about 0.5 cm in the youngest specimens and up to 1.5 cm in adults. It
reaches to the axial pouch of the proxim al interphalangeal joint. The
plantar pouch (Fig. 28— 49), m uch m ore spacious than the dorsal one,
squeezes in upwards under the insertion tendon of the*,deep digital flex or
and runs nearly halfw ay up the m iddle phalanx.
Betw een the distal phalanx and its sesamoid bone is the fissural joint
cavity connected w ith the cavity of the distal interphalangeal joint.
Ligaments of the Distal Interphalangeal Joint
The abaxial collateral ligam ent (Fig. 30— 50) is an isosceles triangle
in shape, its apex being directed upwards. It begins on the peripheral
surface of the m iddle phalanx, from w here it tends, assuming the shape
of a broad fan, towards the distal phalanx. In some European bisons the
anterior m argin o f this ligam ent is accom panied by vertically running
fibres w hich arise on the proxim al phalanx. These fibres correspond to
the abaxial collateral ligam ent of the proxim al and distal interphalangeal
joints.
The axial collateral ligam ent (Fig. 31— 51) is a weak fibrous structure,
united w ith the joint capsule.
The sesamoid bone of the distal phalanx is attached to this phalanx
by tw o small ligaments, an abaxial and an axial (Fig. 32— 54, 55), and
to the m iddle phalanx also by two, axial and abaxial, ligaments (Fig.
32— 52, 53).
The distal interdigital ligam ent is an unpaired sym m etrical six-branched
structure, having attachm ents on all phalanges. The lateral and medial
proxim al branches of this ligament (Figs. 30, 32— 56, 57) take rise on the
abaxial surfaces of the proxim al phalanges. The stronger lateral and
medial m iddle branches (Figs. 31, 32— 58, 59) arise from the surfaces of
the m iddle phalanges facing each other, and the lateral and m edial distal
branches (Fig. 32— 60, 61) from the axial edges and surfaces o f the distal
phalanges and also from their sesamoid bones. The body of the distal
[nterdigital ligament is the place w here fibres of its particular branches
meet and exchange.
9. The Joints of Accessory Digits II and V
The bone skeleton of vestigial digits II and V of European bisons
consists o f tw o rudim entary phalanges connected w ith each other by
means of a small joint. The superior phalanx in both digits resembles
a wheat grain in size and shape and the inferior one a pum pkin seed.
90
F. Kobryńczuk
The accessory digits are joined to each other and also to digits III and
IV only by loose tissue. The so-called tendons of the accessory digits are
distinct structures; they arise from the connective tissue surrounding the
hooves of the rudim entary digits and take the shortest w ay to the abaxial
surfaces of the distal phalanges of the chief digits and their sesamoid
bones.
IV. D ISC U SSIO N
The locom otive organ of the European bison, a typical inhabitant of
the primaeval forest, is efficient, as regards its limbs, for all lifetim e.
The natural environm ent in w hich it dwells, free from human intervent­
ion, permits this animal to keep its body mass in a correct physiological
relation to the biom echanical strength of its limbs and, consequently, is
unpropitious to the form ation of m orphological distortions in them. The
efficiency of the locom otive organ of the European bison is confirm ed
by studies on joints in the pelvic and also the thoracic lim b ( R a d o m ­
s k i, 1972). Neither have pathological changes been found in the osseous
and m uscular system s ( E m p e l & R o s k o s z , 1963; S w i e ż y ń s k i ,
1962). The cases o f deform ation in the lim b joint of European bisons
signalled in literature (J u ś k o, 1953; R a d o m s k i & K o b r y ń, 1969)
and changes in the articular cartilages o f the stifle join t observed in the
present study constitute a v ery small proportion of the large num ber
o f specim ens exam ined.
It is thought that in its manner of locom otion the European bison is
similar rather to sm all ruminants like sheep, goat, roe, etc. than to the
dom estic cattle ( S z t o l c m a n , 1926; W r ó b l e w s k i , 1927). Its m uscu­
lar system also shows m ore similarities to this system of, e.g., the deer
or sheep than to that of the cow ( S w i e ż y ń s k i , 1962). On the other
hand, in skeleton m orphology it comes nearer to dom estic cattle than to
small ruminants (E m p e 1, 1962; R o s k o s z , 1962: E m p e l & R o s k o s z ,
1 9 6 3 ; K o b r y ń , 1973).
In studies carried out so far ( E m p e l & R o s k o s z , 1963) the age
at w hich the pelvic sym physis undergoes ossification is given in appro­
xim ation. The author’s own observations have perm itted the specification
of the time and duration of ossification of this joint, allow ing for di­
m orphic differences.
The newborns and calves o f European bisons, like other mammals (K o ­
s t y r a, 1962; B o c h e n e k & R e i c h e r , 1968), have their acetabula
shallow and, com pared w ith dom estic cattle (V o k k e n et al., 1961),
bisons have the acetabular lip low er.
The lack of the ligam ent of the fem oral head isobserved in many
mammals ( B o c h e n e k & R e i c h e r , 1968; K o s t y r a, 1962). Am ong
Joints of hind-limbs of the European bison
91
others it has not been met w ithin one o f the European bison dom estic
cattle hybrids exam ined (ow n observation). The reduction of this structu­
re is accom panied also by the disappearance of the fovea in the fem oral
head. The great individual variation o f the ligam ent of the fem oral head
in European bisons w ith regard to its size and thickness and the com plete
disappearance o f this ligam ent in some m am mals indicate that unlike all
the other ligaments of the joint it does not strengthens the fram e of the
joint (V o k k e n et al., 1961), but in all probability serves as a track
for the passage o f the blood-vessels supplying the fem oral head. The
m ediocre m echanical role o f this ligam ent is suggested b y the fact that
it is not built o f fibroconn ective tissue but o f fibrocartilaginous one
( A k a j e v s k i j , 1968). The m acroscopic appearance of the ligam ent of
the fem oral head in European bisons confirm s its cartilaginous structure,
except for its extraacetabular part built of fibroconnective tissue.
T able 16
A rticu lar angles o f European bison at the tim e o f w alkin g (in degrees).
Angle
V ertebral colum n — h ip1
Hip — fem u r
Fem ur — tibia
Tibia — m etatarsus
In flex ion
w
158
55
121
136
1 A fter W ę g r z y n & S e r w a t k a
In extension
R ange o f m ovem en t
158
112
128
156
0
57
7
20
(1961)
In contradistinction to dom estic cattle but sim ilarly to the horse (K a­
d 1 e t z, 1932) the European bison possesses the ilioischiofem oral ligament
in the hip joint. On the other hand, it lacks the pubofem oral ligament
observed in dom estic cattle (V o k k e n et al., I.e.), in w hich it strengthens
the capsule o f the hip joint on the ventral side. These differences in the
structure of the fibrous m em brane w hich makes up both the ligaments
m entioned persuade us to suppose that unlike the cow but v ery like the
horse the European bison can perform rapid m ovem ents straightening
the hind legs to the rear, colloquially called »kicking«, as reported by
W r ó b l e w s k i (1927).
In European bisons the rotation axes o f the tw o hip joints lie in a ho­
rizontal plane and cross at 120° in the pelvic cavity. As a result, at the
time of flexion in these joints the stifles alternately straddle the belly
and the fem ora describe the curved surface of conic sectors. The deviation
of the stifles to the outside in their straddling the belly has been esta­
blished at 35° in the cow (V o k k e n et al., 1961) and it is undoubtedly
sm aller in European bisons on account of the fact that their abdom inal
tunics are pulled up. The weight ratio of the organs of the alim entary
92
F. Kobryńczuk
system to the body is greater in dom estic cattle and European bison—
— dom estic cattle hybrids than in European bisons ( S z u l c et al., 1971;
P y t e l , 1969). Further, the abdom inal tunic is better developed in E uro­
pean bisons than in dom estic cattle ( Ś w i e ż y ń s k i , 1962). As a result
of all this, the belly o f the European bison does not flag so loosely and,
what follow s, the fem ora m ove in planes approxim ating to sagittal ones
during flexion at the hip joint. Hence, the European bison has a »light
and graceful« step ( W r ó b l e w s k i , 1927), in w hich, perhaps just for
this reason, it resem bles small ruminants ( S z t o l c m a n , 1926).
In Bison bison (ow n observations) the capacity of the hip joint cavity
is 38 ml and in cows 50 ml (V o k k e n et al., I.e.). In adult European
bisons this capacity has interm ediate values.
In dom estic cattle the lateral meniscus is longer and broader than the
medial one ( A b i e l j a n c , 1957) and thus, except for a few cases, the
situation here is opposite to that in European bisons. H ow ever, the lateral
meniscus is thicker than the m edial in all, European bisons, dom estic
cattle, and European bison— dom estic cattle hybrids ( M o n t a n ć &
B o u r d e l l e , 1917).
The menisci of European bison— d om estic cattle hybrids com pared w ith
those of European bisons (Table 7) are characterized by m orphological
heterosis, and so are the internal organs of these hybrids ( P y t e l &
K r a s i ń s k a , 1971; S z u l c et al., 1971).
The sym m etry of the stifle joint depends on the size ratio of the
menisci ( A b i e l j a n c , 1959). The m ore menisci resem ble each other
in shape and size, the m ore sym m etrical the stifle is and, consequently,
the m ore efficacious it is as an elem ent o f the locom otive organ. D igiti­
grades have sym m etrical stifle joints, plantigrades asym m m etrical ones,
and ungulates hold an interm ediate position. In this respect the stifle
joint of European bisons must be regarded as m ore sym m etrical than
that of European bison— dom estic cattle hybrids.
It is thought ( A b i e l j a n c ,
1957; A k a j e v s k i j ,
1968; B r i i h l ,
1969) that as fibrocartilaginous structures the m enisci absorb shocks in
the stifle joint. In the author’s opinion, their presence in the gap of the
fem orotibial joint conditions also the simultaneous occurrence of two sorts
of m ovem ents, gliding and rolling.
The transerverse ligam ent of the stifle, present in other mammals
(Abieljanc,
1957; K o s t y r a ,
1962, V o k k e n
et al., 1961; B o ­
c h e n e k & R e i c h e r, 1968), is missing in European bisons. The lack
of this structure has also been observed in Ft European bison— domestic
cattle hybrids and in Bison bison. On the other hand, in all European
bisons a small ligamental structure, absent from other mammals, has
been found extending from front to rear betw een the intercondylar tu­
Joints of hind-limbs of the European bison
93
bercles of the tibia; the term »longitudinal ligament of stifle (lig. longi­
tudinale genus)« has been proposed for it (Fig. 11— 24).
The insertions of the collateral ligaments of the fem orotibial joint,
situated relatively low on the tibia, allow the condyles of the fem ur to
glide over the articular surface of the tibial condyles, forw ard during
extension and backward during flexion. As in other ungulates (V o k k e n
et al., 1961), these ligaments are tensed very m uch w hen the stifle joint
is extended, preventing any movem ents except those sagittal in direction.
A small range of adduction, abduction and rotation is possible when the
joint is h a lf-flex ed or flexed and the collateral ligaments are m oderately
slackened. The tension and relaxation of these ligaments is caused by
the fact that the fossae fo r their attachment on the fem u r do not lie in
the rotation axis of the fem orotibial joint, but above it and somewhat
to the rear. W riters differ in opinion as to the structure of the lateral
collateral ligam ent of the fem orotibial joint. This ligam ent is believed
to arise from the fibres of the fibrous m em brane of the joint capsule
and those of the tendon of the peroneus longus m uscle ( S w i e z y n s k i ,
1962), or exclu sively from the fibres of the capsule (P e 1 e i n e r, 1932).
The author’s investigation confirm s the first view and at the same time
it shows that some of the fibres of the lateral collateral ligament extend
dow nw ard to form the superficial layer of the fibrous band of the. leg.
The cruciate ligaments of the stifle joint ( E l l e n b e r g e r & B a u m ,
1943; P o p l e w s k i , 1934) are said to inhibit hyperextension in this
joint. Observations made on preparations of the stifle joints of European
bisons show that these ligaments, in addition, fu lfil the task, not less
important, o f gliding the fem oral condyles over the articular surfaces
of the tibial condyles. This function is perform ed as follow s: w hile the
stifle joint is being straightened, the anterior cruciate ligam ent winds
on the sem i-cylinder form ed by the bottom of the intercondylar fossa
of the fem ur. As a result, the ligament is shortened and pulls the distal
end of the fem ur to the front. In flexion the anterior cruciate ligament
unwinds and, in turn, the posterior cruciate ligament becom es wound on
the sem i-cylinder m entioned; thus shortened, it pulls the distal end of the
fem ur backwards. The m enisci which constitute the natural substratum
of the condyles of the fem ur, travel forw ard and backw ard together
w ith them.
There are controversial view s on the com m unication betw eeen the two
cavities of the fem orotibial joint in dom estic cattle. Som e investigators
think that these cavities are isolated from each other ( C h a u v e a u et
al., 1903; S i s s o n & G r o s s m a n , 1960) and others claim that there
is always a com m unication between them (V o k k e n et al., 1961; N i c ­
k e l et al., 1960). M y ow n observations show that hardly 7 European
94
F. Kobrynczuk
bisons in a hundred, and then ch iefly old ones, have the tw o cavities
connected, whereas in most European bisons, European bison— dom estic
cattle hybrids, Watussi cattle and buffaloes they are isolated from each
other by a joint partition. A dm ittedly, how ever, there is a com m unicat­
ion betw een the medial sac of the fem orotibial joint and the cavity of the
fem oropatellar joint in all the Bovinae not excluding European bisons.
A ccordin g to som e authors ( E l l e n b e r g e r & B a u m , 1943; V o k k e n et al., I.e.), all the patellar ligaments are the insertion tendons of
the quadriceps m uscle of the thigh. A b i e l j a n c (1962) writes that only
ihe m iddle patellar ligam ent is derived from that muscle, whereas the
medial patellar ligam ent has origin from the medial fem oropatellar liga­
ment. In his and other author’s opinion ( S i s s o n & G r o s s m a n , 1960),
in respect of derivation the lateral patellar ligament is associated w ith
the lateral fem oropatellar ligam ent and the tendon of the biceps m uscle
of the thigh.
In European bisons the m iddle patellar ligament is the dow nw ard
extension o f the fibres of the rectus m uscle of the thigh and, partly, the
lateral vastus m uscle. The author is of opinion that the m edial patellar
ligam ent of these animals is as a w h ole a product of the fibrous m em ­
brane of the joint capsule, from w hich also the lateral patellar ligament
arises (medial portion), receiving some fibres from the tendon of the
biceps m uscle of the thigh (lateral portion).
European bisons and Fx European bison— dom estic cattle hybrids have
a ligam ental structure w hich does not occur in dom estic cattle but is
observed in a som ewhat m odified form in the pig (K o s t y r a, 1962),
i.e. the fem oropatellotibial ligam ent (Figs. 12, 14— 34). This ligament ari­
ses from the com bination of the medial fem oropatellar ligament proper
w ith the aponeurosis of the fedial vastus m uscle and the fascia genus.
The lateral fem oropatellar ligam ent is not present in European bisons.
The opinion prevails ( A b i e l j a n c , 1955, 1962) that in dom estic cattle,
as in the horse but to a smaller extent, the so-called static loop exists
in the stifle joint, locking it w henever the animal rests in a standing
position. The elements of this loop are, among others, the m iddle and
m edial patellar ligam ents and the medial ridge of the fem oral trochlea.
In European bisons and F t European bison— dom estic cattle hybrids the
loop should be still m ore efficacious than it is in dom estic cattle on
account of its additional element, w hich is the fem oropatellotibial liga­
ment.
In the shanks of some European bisons and one bu ffalo exam ined the
rudim entary fibulas w ere found (Fig. 18), w hich had not even been m en­
tioned before in the osteology of lim bs o f the European bison (E m p e 1
& R o s k o s z , 1963) and dom estic cattle.
Joints of hind-limbs of the European bison
95
A peculiarity of the hock joint is that in com parison with other sy n o­
vial joints its postnatal developm ent is small, as evidenced by the small
increm ents in its chief cavity and the sm all grow th of its osseous com ­
ponents after birth ( K o b r y ñ c z u k & K o b r y ñ , 1973).
The relative thinness of the articular cartilage of the tarsocrural joint
in European bisons is w orthy of notice, and this is true as regards juvenile
specimens. It is generally accepted that articular cartilage thickens during
w ork (V o k k e n et ah, 1961) and that its thickness is proportional to
the pressure exerted on it ( B o c h e n e k & R e i c h e r , 1968). H ow ever,
P o p l e w s k i (1927) gives a different opinion. A ccording to him, pressu­
re makes articular cartilage thinner. This last view w ould seem to be
true, since, fo r instance, the stress in the tarsocrural joint of European
bisons is greater than that in the fem oropatellar joint and yet its articular
cartilage is considerably thinner than that in the latter joint.
Synovial fossae (Fig. 19) in the hock joint are noticeable. The distribu­
tion of these structures and the role they play in mammals have not as
yet been explained. A b i e l j a n c (1968) finds them situated m ost fr e ­
quently in the elbow, hip, hock and m etacarpo- and metatarsophalangeal
joints. In man synovial fossae are believed to be pathological lesions
( B r ü h l , 1969). Some writers (V o k k e n et ah, I.e.) think that their
function is to produce synovial fluid. The accum ulation of synovial fossae
in the tarsal joint of European bisons suggests another hypothesis, nam e­
ly, tnat they appear in joints in w hich a relatively large articular surface
is accom panied by a synovial m em brane small in area, w hich is the case
in the tarsal joint.
In European b'sons, as in other Paraxonia, tw o funcionally equivalent
joints occur in the hock, the tarsocrural and the proxim al intertarsal. The
flexion-exten sion range approxim ates to 65° in the first of these joints
and to 95° in the second, adding to 160°. This cum ulative flex ion -ex ten ­
sion range is great relative to the »dem ands« of the tarsal joint during
normal walk. It should then be supposed that the tarsocrural and pro­
xim al intertarsal joints do not w ork sim ultaneously, the first of them
being active during normal walk, whereas the other joint is activated
only in situations demanding special effort, e.g., in fighting w ith the horns
or clim bing up a hill. In both cases, how ever, the straightening fo rce F
(Fig. 27) is equal, the tuberosity of the tibial tarsal bone being the point
of its application. There are two arms o f this force, p and q. The shorter
arm p arises from the rotation centre o f the tarsocrural joint (0), the
other, longer arm q begins at the rotation centre of the proxim al inter­
tarsal joint. Thus, w e are here concerned w ith tw o mom ents of one and
the same force, for which
F •p < F • q
96
F. Kobrynczuk
The dorsal pouches of the capsules of both metatarsophalangeal joints
have greater absolute dimensions in European bisons than in cattle (E m p e l , 1969); they are 6 cm long in cow s and may reach up to 8 cm in
European bisons.
In cow s two axial collateral ligam ents o f the metatarsophalangeal joints
have been reported ( K o h l e r ,
1902; N i c k e l & L a n g e r, 1953) to
have their anterior edges joined. In European bisons these ligaments are
anteriorly connected only by the synovial membrane. M oreover, the ab­
axial collateral ligam ents o f the m etatarsophalangeal joints are uniform
structures in cows, evidently consisting o f tw o layers in European bisons
(Fig. 30— 3).
M acroscopic observation o f the m iddle interosseous m uscle of the Euro­
pean bison made it possible to com plim ent the results obtained by § w i ez y n s k i (1962). M uscle fibres em bedded in the fibrous structure of this
m uscle belong only to its deep portion, naturally before its division into
branches. Red m uscle fibres w ere observed on n on -fixed preparations
of this muscle, both on its surface and in sections. In longitudinal sections
these fibres concur in fou r vertical trips separated from each other by
strips of fibrous tissue, whereas in cross-sections there appear four fields
lying side by side and consisting of concentric rings or arches of muscu­
lar tissue. In both cases the arrangem ent of the m uscle fibres is remini­
scent o f the fou r interosseous muscles in the tetradactyl limb. In
contradistinction to the situation found in the horse, in w hich the proxi­
mal portion of the m iddle interosseous m uscle contains m ore muscle fi­
bres (K r y s i a k, 1937/38), in European bisons m ore of these fibres are
accum ulated in the distal portion of the m uscle discussed.
The presence of m uscular tissue in the interosseous m uscle is observed
in all European bisons, this tissue being particularly abundant in juveni­
les, whereas in dom estic cattle ( V o k k e n et alt., 1961) its fibres are
met w ith only in calves. N i c k e l et al. (1960) hold the opinion that in
small ruminants this elem ent has a m uscular structure, but R e i s e r
(1903) w rote nothing on this subject in his study on the m uscular system
in small ruminants. If how ever N i c k e l et al. (1960) are right, the
m iddle interosseous muscle of European bisons w ould be m ore similar
in this respect to this muscle in small ruminants than to its counterpart
in dom estic cattle.
In the deer, sheep and goat ( R e i s e r , 1903) the interdigital branch
of the m iddle interosseous muscle, before sinking under the interdigital
intersesamoidal m uscle, sends o ff fibres sym m etrically to the sides, to
the axial sesamoid bones, w hich has also been found in European bisons
but in a small num ber of specimens.
The plantar pouch of the proxim al interphalangeal joint extends one-
Joints of hind-limbs of the European bison
97
third of the w ay up the proxim al phalanx in dom estic cattle (E m p e 1,
1969) and therefore it is shorter here than in European bisons. N i c k e l
&r L a n g e r (1953) have found this pouch situated farther to the outer
side in cattle than it is in European bisons.
No elastic ligam ents have been encountered in the distal interphalangeal joint of the European bison, whereas in dom estic cattle it is situated
on the antero-axial side of the joint ( K ö h l e r , 1902), neither has the
elastic nature of the ligam ent of the sesamoid bone of the distal phalanx
been established, although it has been recorded from dom estic cattle by
N i c k e l & L a n g e r (1953).
Ś w i e ż y ń s k i (1962) w rites that the m iddle interosseous muscle in
the fore-lim b of the European bison gives o ff special branches to accesso­
ry digits II and V. R a d o m s k i (1972) is o f the opinion that these digits
in the fo re -lim b of the European bison have no direct connection w ith
the m iddle interosseous muscle, but are held, above all, by the fascia
metacarpi. The same opinion is expressed by K ö h l e r (1902) w ith regard
to the attachm ent of these digits in dom estic cattle and a similar situation
has also been observed in the hind-lim bs of European bisons.
Inform ation concerning the osseous structure of rudim entary digits
II and V o f dom estic cattle is controversial ( N i c k e l et al., I960;
V o k k e n et al., 1961). In bisons, European bisons and W atussi cattle
each rudim entary digit consists of two phalanges. It w ould be inconside­
rate to claim that in the Bovinae it is the distal phalanx that is absent
from such digits, although this w ould be suggested by the law of reduct­
ion of digits ( P o p l e w s k i , 1948).
The tendons of accessory digits II and V should be considered, in spite
of their name, to be descended from the interdigital ligaments, connecting
the abaxial digits w ith the axial ones in tetractyl animals. The m echa­
nical role o f the accessory digits of the European bison seems to be clear.
Their hoofs bear sings of w ear and therefore, notwithstanding their high
position, they com e into contact with a soft substratum, thus increasing
the bearing surface of the digital organ in these animals. Easy movem ents
of European bisons on swam py and m iry ground is m entioned also by
W r ó b l e w s k i (1927).
It is difficu lt to determ ine the angles form ed by long bones meeting
in joints on carcasses and the flexion -exten sion range of m ovem ents in
particular joints of m oving European bisons. The m ethod o f determ inat­
ion of these values on the basis of photographs is not precise and this
is w hy values obtained in this way, offered in Table 16, are left without
any comm ents.
A t an analysis of the dimensions of structures m aking up particular
join t of bones in the hind-lim bs o f European bisons it should be stated
98
F. Kobryńczuk
in general that the postnatal grow th of these joints in males of groups 0,
I and II is intenser than in fem ales of the same age groups. In adult
European bisons becom e sm aller and sm aller in males of groups III— V,
this drop in increase being m ilder in fem ales.
M oreover, it m ay be stated that the dimensions presented are nearly
alw ays larger in males and so are the grow th coefficients.
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51. W r ó b l e w s k i K., 1927: Ż u b r P uszczy B iałow ieskiej. 1— 232. W yd. Polskie,
Poznań.
52. Ż a b i ń s k i J. (Ed.), 1947— 65: P edigree b o o k o f the E uropean bison. 1— 370.
Państw . W yd. Nauk., W arszaw a.
53. Ż a b i ń s k i J. & R a c z y ń s k i J. (Eds.), 1972: E uropean bison pedigree book
1965— 69. 1— 78. Państw . W yd. Nauk., W arszaw a.
A cc ep te d , January 20, 1975.
Franciszek K O B R Y Ń C Z U K
P O Ł Ą C Z E N IA K O Ś C I K O Ń C Z Y N M IE D N IC Z N Y C H Ż U B R A
W R O Z W O JU P O Z A P Ł O D O W Y M
Streszczenie
Badania p rzeprow a dzon o na zw łok ach 71 żu b rów (47 sam ców i 24 samic). Ponadto
dokonano ob serw a cji p ow ierzch n i staw ow ych na k ośćcach 48 żu b rów (27 sam ców
i 21 sam ic) stan ow iących część zb iorów m u zealnych O środka B adań nad A natom ią
Ż ubra przy A k a dem ii R olniczej w W arszaw ie. P ad łe zw ierzęta p och od ziły z k ra­
jo w y ch ośrod k ów h odow lan ych. M ateriał p od zielon o na 6 grup o b ejm u ją cy ch zw ie­
rzęta począw szy od p łod ów tuż przed u rodzen iem i n ow orod k ów aż do osobnik ów
najstarszych. W celach p orów n a w czych p rzep row a d zon o rów n ież fragm en taryczn e
badania na innych Bouinae a m iędzy in nym i na b y d le d om ow y m i k rzyżów kach
bydła d om ow eg o z żubrem .
Zbadano w szystkie połączenia k ości w k oń czyn a ch m iedniczych . W badaniach tych
określon o, p osłu g u jąc się m etodą w łasną, w ielk ość kątow ą k rzyw izn pow ierzch n i
staw ow ych, w artość prom ieni i cię ciw tych k rzyw izn oraz inne param etry kątow e.
Na zw łokach św ieżych zbadano p ojem n ość i kształt ja m sta w ow ych przez w p ro ­
w adzen ie doń k olorow eg o, ciepłego roztw oru żelatyny. P onadto na zw łok ach u trw a­
lon ych przy p om ocy form alin y zm ierzon o g ru bość chrząstek staw ow ych oraz d o­
kon an o pom iarów w ięzad eł i łąkotek. Poza tekstem u jm u ją cy m m orfolog ię p o łą ­
czeń, badania u dok um entow ano 16 tabelam i i 32 rysunkam i w yk on an ym i przez
autora.
Table 1C
D im ensions o l ridges o f fe m o ra l troch lea (in m m and degrees).
Lateral ridge
G roup, No
nam e
Species, sex
Length
(chord)
Me,
Curvature
A ngle
S
European bison, M
0
I
II
III
IV
v
G.C.
European bison, F
0
I
II
III
IV
V
8
5
16
17
15
13
52.0
61.1
80.6
85.3
89,4
87.9
1.69
4
1
9
6
7
18
48.3
65.0
71.3
75.8
78.8
81.8
1.69
4
69.6
G C.
Bison bison, F
Bison bison
— B oonasus
h ybrid , M
1
European bison
— dom estic cattle
hybrids, M
European bison —
dom estic cattle
hybrids, F
3.9
2.2
4.4
7.9
3.6
7.9
1.8
3.7
5.0
3.3
7.5
3.6
5.4
9.3
4.0
9.0
2.5
4.9
6.3
4.0
91 8
Fes!
Felon
Fellach
Festyn
Fey
Fez
Farad
Felly
Fela
Fam a
31.5
35.9
41.9
43.5
45.5
44.8
1.42
111
117
148
157
159
158
1.42
29.2
37.0
37.7
39.2
40.3
42.5
1.46
111
123
142
150
156
154
1.39
37 3
137
48.2
46.4
46.5
50.8
50.3
42.6
47.5
52.4
40.6
40.5
41.5
139
149
127
138
154
156
145
131
178
155
48.5
40.2
43.6
41.4
158
147
146
148
2
10
1
1
95.2
77.1 5.3 6.9
83.5
79.5
V
10
3.5
9.7
3.9
4.4
6.2
0.'9
3.0
6.6
2.4
2.8
3.9
8.2
2.0
6.0
5.6
5.8
1.3
3.8
3.6
57.0
77.5
107.3
113.1
115.9
114.1
2.00
57,7
81.0
94.7
100.5
103.6
106.7
1.85
9.0
1.4
7.8
5.6
4.5
6.0
5.3
1.8
7.3
5.0
3.9
5.2
9.3
4.5
6.6
5.3
9.8
4.5
6.4
5.0
104 P
144
87.0
90 0
91.0
94.0
83 0
93 0
100.0
74.0
81.0
81.0
D om estic cattle, M
D om estic cattle, F
W atussi cattle, M
W atussi cattle, F
Length
(chord)
2.1
35.2
47.2
62.3
64.2
64.3
62.3
1.78
34.5
46 0
56.4
56.9
58.7
60.1
1 74
59.6
108
110
119
123
128
129
1.19
113
114
124
124
123
1 09
69.0
118
609
59.6
73.5
89.9
60.6
72.0
78.5
52 8
62.6
60.4
127
146
118
114
124
119
123
130
121
129
67 0
62.0
66.1
56.0
120
109
115
129
1.3
.
3.2
9.3
4.7
10,4
8.5
38
8.4
6.3
3.6
3.3
2.6
8.2
124
118.5
1.3
42
1.5
95
76
4.6
1.7
11.3
9.4
3.3
41
122
109.0
114.0
126.0
131 0
107.0
1240
138.0
102.0
109.0
109.0
116.0
100.6
111.5
102.5
1.4
ridge
C urvature
r
Angle
s
x
Bubalus bubalus L., M
1
82.4
44 5
136
100.0
556
128
Zebu, F
1
72.0
37.0
154
90.3
56.6
110
Y ak. F
1
57.2
31.1
134
75 2
56 5
128
T able 12
D im ensions of trochlea o f tibiotarsone (in m m and degrees).
P rox im a l trochlea
D istal trochlea
G rou p
n
Length
(chord)
X
s
8
5
16
17
15
13
38.1
42.8
50.2
50.5
52.6
52.0
1.36
3.1
1.8
3.3
2.4
1.4
3.7
4
1
9
6
7
18
37.5
42.0
44.7
46.8
47.8
48.2
1.28
r
V
Lateral ridge
M edial ridge
L ateral ridge
C urvature
A ngle
X
X
s
21.1
22.7'
26.4
26.5
26.9
27.2
1.29
129
141
143
145
154
149
1.16
7.8
6.6
9.4
9.0
6.8
8.1
21.4
27.8
23.9
24.4
25.0
25.3
1.18
123
134
139
147
143
144
1.17
y
Length
(chord)
s
X
Curvature
A n gle
X
s
V
X
4.4
11.0
6.9
4.0
1.2
3.6
18.6
22.2
24.2
24.9
25.1
25.1
1.35
151
164
167
171
180
180
1.19
19.6
20.5
21.9
22.7
23.4
23.8
1.21
140
155
166
173
169
173
1.24
V
X
Length
(chord)
s
r
V
X
M edial ridge
Curvature
_
A n gle
X
s
Length
(chord)
r
x
C u rvatu re
A n gle
M ales
0
I
II
III
IV
V
G.C.
8.1
4.2
6.6
4.8
2.7
7.1
6.0
4.7
6.6
6.2
4.4
5.4
36.5
44.7
48.0
49.8
50.2
50.2
1.39
1.6
4.9
3.3
2.0
0.6
1.8
10.4
4.4
11.0
11.3
10.1
8.2
6.9
2.7
6.6
6.6
5.6
4.6
25.0
30.0
34.4
35.0
36.5
36.6
1.47
0.7
1.9
2.2
2.1
1.8
2.0
2.8
6.3
6.4
6.0
4.9
5.5
12.8
15.2
17.4
17.7
18.6
17.9
1.40
202
201
198
194
192
190
0.93
12.5
15.1
16.6
15.9
16.8
16.3
1.30
223
200
201
188
195
192
0.86
9.5
6.7
8.2
9.9
7.3
7.1
4.7
3.3
4.1
5.1
3.8
3.7
28.8
34.4
38.7
39.4
41.2
40.8
1.42
1.4
1.9
1.7
2.1
1.6
2.0
4.9
5.5
4.4
5.3
3.9
4.9
14.4
17.6
20.3
20.4
21.4
21.2
1.47
178
199
215
213
205
212
1.19
13.4
18.1
18.0
18.5
19.1
19.3
1.44
206
208
209
213
215
211
1.02
78
7,7
103
75
6.0
6,2
4.4
3.9
4.8
3.5
2.9
2.9
5.3
5.7
5.7
11.1
2.5
2.7
2.6
5.3
Fem ales
0
I
II
III
IV
V
G.C.
1.3
2.3
3.3
2.7
2.9
4.9
6.9
5.6
10.9
5.8
8.4
11.3
7.8
3.9
5.9
7.8
36.8
40.0
43.4
45.4
46.4
47.3
1.28
,
0.8
1.1
2.1
1.5
1.8
2.4
4.5
3.2
5.6
8.1
8.1
8.1
3.4
4.7
4.8
4.7
23.2
29.8
30.7
31.8
33.4
32.2
1.39
1.4
1.8
2.0
1.7
4.7
5.7
6.0
5.3
10.0
6.5
11.3
13.9
5.0
3.5
5.8
7.2
26.2
35.1
34.8
35.4
36.3
37.1
1.42
1.4
2.0
0.3
1.6
4.0
5.6
0.9
4.3
T able 5
D im ensions o f fem ora l con d y le (in m m and degrees).
M edial con d y le
L ateral con dyle
C urvature
G rou p
Sex
Species
n
s
x
0
I
II
III
IV
V
G.C.
M
European bison
0
I
II
III
IV
V
G.C.
5
16
17
15
13
4
1
9
6
7
18
Bison bison
B. b o n a s u s —
B. bison h yb rid
D om estic cattle
M
M
2
F
10
W atussi cattle
M
F
Bubalus bubalus
M
1
1
1
1
F
Z ebu
W idth
L ength (chord)
36.8
53.5
66.0
69.7
70.8
71.8
1.95
35.3
50.0
60.5
64.0
63.7
64.6
1.83
2.2
3.2
4.8
2.6
1.0
3.9
X
6.0
21.0
6.0
7.3
3.7
1.4
5.4
32.2
41.0
46.7
48.8
49.6
2.36
2.0
3.6
3.0
2.4
X
9.5
19.0
27.5
33.2
35.1
35.9
36.2
1.88
151
153
167
167
161
164
1.09
18.7
25.3
30.4
32.7
31.9
32.5
1.74
150
162
168
156
172
162
1.08
7.3
5.1
2.0
7.4
3.7
2.2
2.0
3.0
29.0
38.5
41.2
42.8
43.4
2.09
4.6
3.4
3.1
4.6
2.0
9.4
4.8
4.7
2.6
6.0
3.6
2.0
s
X
11.2
1.0
_
V
S
20.8
2.8
A n g le
r
v
r
w idth
ratio
V
18.1
12.0
21.2
13.8
6.0
'5.7
3.0
4.5
10.1
9.5
4.8
7.4
6.4
9.5
4.0
3.7
2.6
1.6
15.9
6.2
x
1: 0.66
1:0.61
1:0.49
1:0.45
1:0.45
1:0.78
1:0.70
1:0.57
1:0.55
1:0.49
1:0.49
s
V
13.8
5.4
4.3
2.0
2.4
6.6
48.0
64.2
69.9
71.4
72.1
1.92
3.5
3.0
1.4
1.7
C urvature
W idth
Length (chord)
X
s
V
32.0
43.3
47.4
49.4
50.6
2.40
4.4
3.2
2.2
1.7
1.4
13.8
7.4
4.6
3.4
2.8
1.7
1.4
2.4
6.7
3.0
2.2
3.8
10.5
29.0
38.2
41.8
43.0
43.9
2.18
2.8
2.2
2.2
2.2
7.3
5.3
5.1
5.0
40.3
56.9
x
X
20.1
34.3
46.0
56.8
63.0
63.3
64.0
1.87
24.0
32.1
35.0
35.7
36.1
1.90
175
182
179
177
177
1.07
17.3
23.0
28.4
31.5
32.2
31.8
1.86
166
174
181
179
176
181
1.09
28.5
173
0
I
II
III
IV
V
G. C.
0
I
II
III
IV
V
G.C.
M
Fest
F elon
F ellach
F estyn
Fey
Fez
Farad
F elly
F ela
Fam a
8
5
11
8
41.3
43.5
60.3
61.4
65.0
64.0
1.54
37.5
43.0
51.8
51.0
54.7
59.7
1.59
2.6
2.3
1.0
2.6
153
1:0.29
69.2
55.0
34.7
172
1:0.23
1:0.35
1:0.46
76.4
62.2
55.6
40.7
38.3
31.1
186
176
1:0.37
1:0.53
69.0
61.0
48.5
48.3
35.7
30.8
150
163
1:0.42
1:0.26
79.2
63.9
5.9
58.5
43.8
9.2
3.6
1.6
166
166
39.9
32.2
7.9
13.1
6.1
3.8
3.2
7.9
3.2
6.2
1.4
2.3
36.3
29.9
162
184
1 : 0.21
1 : 0.20
66.8
50.5
33.9
161
1:0.32
66.2
43.5
33.1
185
1:0.52
27.9
180
1:0.28
54.5
37.6
27.3
174
1:0.45
43.8
55.9
M edial m eniscus
x
s
s
s
v
V
X
34.0
14.4
25.0
9.9
12.1
7.1
38.7
46.1
63.5
64.9
67.0
70.0
1.81
s
V
X
6.7
6.9
7.9
2.2
3.3
8.0
27.8
32.0
45.3
46.7
51.0
51.0
1.83
s
V olum e
Thickness
W idth
Length
V olum e
Thickness
23.2
28.0
34 6
37 0
37.3
41.5
1.79
0.6
1.5
1.0
2.0
2.2
2.2
2.2
5.4
2.4
4.8
5.1
5.0
1.0
2.9
2.0
4.8
5.0
8.3
1.7
20.0
22.2
5.0
3.0
I.45
24.8
28.1
5.62
11.9
II.2
16.0
16.8
17.8
18.3
1.64
4.0
8.0
11.2
13.0
14.0
18.6
4.65
1.3
13.9
13.9
18.0
19.3
1.4
2.4
20.0
20.2
0.6
0.6
1.6
9.4
11.5
7.8
12.4
3.0
3.0
1.2
7.5
1.0
5.5
V
X
s
2.9
1.6
1.8
0.2
9.0
7.8
15.3
16.2
15.7
15.8
1.76
0.7
0.8
1.8
0.8
1.0
1.2
V
X
s
1.2
2.2
2.0
1.7
15.2
1.2
6.4
36.2
46.0
50.8
55.0
58.0
59.7
1.65
2.6
3.0
5.0
1.4
2.2
5.6
3.2
6.3
1.8
3.0
25.5
34.5
36.2
38.5
41.0
44.3
1.74
0.8
0.5
0.8
0.1
0.5
1.8
1 .0
3.5
1.0
2.3
2.8
6.3
9.9
9.3
11.5
12.3
11.3
13.6
1.37
7.8
10.2
11.8
4.9
6.4
7.6
4.3
7.0
19.7
20.6
23.8
28.1
6.55
0.3
3.0
4.8
4.1
3.9
2.2
7.0
4.3
24.4
19.9
16.4
7.8
0.8
7.0
1.6
16.0
1.0
7.4
3.2
7.0
10.0
11.5
13.3
15.5
4.84
ic cattle hybrid
61.0
76.0
65.0
64.0
42.0
40.0
48.0
44.0
45.0
21.0
20.0
20.3
20.0
18.7
24.0
21.0
30.0
27.0
25.0
75.0
55.0
66.0
53.0
48.0
43.0
43.0
36.0
14.0
19.0
18.0
18.3
30.0
19.0
21.0
17.0
68.0
A c ta t h e r i o l . , 21, 4, w k l e j k a s t r . 64/65.
2.7
59.5
48.0
26.9
28.0
41.3
42.8
43.4
44.4
1.65
1.6
4.8
71.7
59.8
6.3
6.3
1.0
1:0.41
38.6
X
1.4
4.7
4.0
4.4
6.7
58.2
v
1.6
2.6
2.2
2.5
3.7
1:0.63
1:0.59
1:0.49
1:0.51
1:0.47
1:0.46
1.8
75.1
W idth
4.2
2.3
8.1
3.2
8.9
1:0.51
V
European bison
M
5.0
1:0.50
1:0.48
1:0.47
1:0.44
1:0.42
168
Length
s
6.2
5.9
29.3
Lateral m eniscus
n
3.5
3.2
4.5
s
38.6
Linear dim ensions and volu m e o f m enisci (in m m and cu. cm).
G rou p
Length:
w idth
ratio
58.2
T ab le 7
S ex
-------
A n gle
•r
58.0
64.0
70.0
71.0
61.0
78.0
74.0
57.0
65.0
58.0
44.0
43.0
51.0
46.0
47.0
52.0
53.0
40.0
47.0
40.0
14.0
12.7
14.3
12.5
13.0
14.7
14.5
13.0
11.7
13.0
10.0
14.5
23.0
21.0
18.0
—
34.0
12.0
11.0
14.0
1.7
11.0
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