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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. REFERENCES 1. A b i e l j a n e G. S., 1955: A n a to m o -fu n k cjo n a ln y e osobenn osti b e d ro -k o le n n n ogo sustava lośadi v sv ja zi so statikoj i din am ikoj ta zov oj konecnosti. Tr. K ije v . Viet. Inst., 12: 114— 124. K ijev . 2. A b i e l j a n e G. S., 1957: M en isk ov o-sv ja zoén y j apparat kolen a svini, lośadi, k ru pn ogó skota i jeg o r o l’ v fu n k c y ji sustava. Tr. K ijev . Viet. Inst., 13: 271— — 277. K ijev . 3. A b i e l j a n c G. S., 1959: T y p y k olin n oh o suhloba ssavciv. Nauk. P raci Vet Fak., 14: 153— 154. K y jiv . 4. A b i e l j a n c G. S., 1962: P rja m i zv ja zk y k olin n oji casky svyni, v e ly k o ji r o h a to ji h udoby i k on i ta ih rol u fu n k cii k olin n oh o suhloba. Nauk. Praci Vet. Fak., 25, 18: 68— 74. K y jiv . 5. A b i e l j a n c G. S., 1968: K is tk o v o -z v ja z k o v y j pojas k olin n oh o suhloba pa lcehid nyh ssavciv. Z ah od y Min. Silsk. G osp. URSR, 3: 171— 173. K y jiv . 6. A k a j e v s k i j A. I., 1968: A n atom ia dom aśnih zivotnyh. Izd. K olos: 1— 230. M oskva. 7. B o c h e n e k A. & R e i c h e r M., 1968: A n atom ia człow ieka. Państw. Zakł. W yd. Lek., W arszaw a. 8. B r ü h l W. (ed.), 1969: C h oroby narządu ruchu. Państw . Zakł. W yd. Lek., W arszaw a. 9. C h a v e a u A. & A r 1 o i n g S., 1903: T raité d ’an atom ie com p a rée des anim aux dom estiques. B ailliére, 1: 1— 684. Paris. 10. D a v l e t o v a L. V., 1960: The grow th o f digestion organs in the course of ontogenesis in sheep. Biul. M osk. O bść. Isp. P rirod y, Biol., 65, 2: 107— 119. [In R ussian w ith English summ.]. 11. D u e r s t J. U., 1926: V ergleich en d e U n tersuchungsm ethoden am Skelett bei Säugern. U rban und S ch w arzen berg, 7; 125— 530. B erlin — W ien. 12. E l l e n b e r g e r W. & B a u m H., 1943: H an dbuch der v erg leich en den A n a tom ie der H austiere. Sprin ger, 1: 1— 1155. Berlin. 13. E m p e l W., 1962: M orp h ologie des Sch ädels von Bison bonasus ( L i n n a e u s , 1758). A cta theriol., 6, 4: 53— 111. 14. E m p e l W., 1969: Studia nad anatom ią chiru rgiczną i rentgen ow sk ą palców bydła. Z eszyty Nauk. S G G W , 1: 7— 50. 15. E m p e l W. & R o s k o s z T., 1963: Das Skelett der G liedm assen des W isents, Bison bonasus ( L i n n a e u s , 1758). A cta theriol., 7, 13: 259— 300. 16. J a n i c k i S., 1938: B adania nad szkieletem żubra (Bison bonasus L.). Prace R oln .-L eśn e P.A.U., 27: 1— 55. 17. J u ś k o J., 1953: D im orfizm p łciow y szkieletu żubra (Bison bonasus). Folia m orph ol., 4, 1: 1— 30. 18. K a d 1 e t z M., 1932: A n atom isch er A tlas E xtrem itätengelen k e v on P ferd und Hund. U rban und S ch w a rzen b erg : 1— 71. B erlin — W ien. Joints of hind-limbs of the European bison 99 19. K o b r y ń H., 1973: The th orax in E uropean bison and other rum inants. Acta theriol., 18, 17: 313— 341. 20. K o b r y ń H. & K o b r y ń c z u k F., 1973: P seu doarthrosis in the European bison. A cta theriol., 18, 18: 347— 350. 21. K o b r y ń c z u k F. & K o b r y ń H., 1973: P ostem b ry on ic g row th o f bones o f the au topodia in the European bison. A cta theriol., 18, 16: 289— 311. 22. K o c h W., 1932: Ü ber W ach stum s- und A ltersverä n deru n gen am S kelett des W isents. A bh. M at.-naturw . Abt., Bayer. A kad. W iss., S u pl.-B d., 15 A b h .: 555— — 678. 23. K o s t y r a J., 1960: Połączen ia kości w koń czyn a ch św ini d om ow ej. Annls Univ. M. C u rie-S k łod ow sk a, Ser. DD 15, 11: 137— 190. 24. K ö h l e r A. A., 1902: U ntersuchungen ü ber P halangenbänder der H austhiere und da V ork om m en der Sesam bein an Zehen der Fleischfresser. A rch . wiss. prakt. T hierheilkunde, 29: 69— 108. Berlin. 25. K r y s i a k K., 1937/38: W spraw ie u ścięgnienia m ięśnia m iędzyk ostn ego p o środ k ow ego u Equidae. Folia m orphol., 8. 26. K r y s i a k K., 1951: W iązadło kark ow e (Ligam entu m nuchae) żubra — Bison bonasus. Folia m orphol., 2, 10: 271— 283. 27. M a r t i n P. & S c h a u d e r W., 1938: L eh rbu ch der A n atom ie der H austiere — 3: 1— 560. Schickhardt. Ebner, Stuttgart. 28. M o n t a n e L. & B o u r d e l l e E., 1917: A n atom ie region ale des anim aux dom estiqu es (Rum inants):, 2: 1— 368. Paris. 29. N i c k e l R. & L a n g e r P., 1953: Z eh en gelen ke des Rindes. T ierärztl. W schr., 14: 211— 253. B erlin und M ünchen. 30. N i c k e l R., S c h u m m e r A. & S e i f e r l e E., 1960: L eh rbu ch der A n atom ie der H austiere. Parey, 1: 1— 206. B erlin— H am burg. 31. Nomina A natom ica V eterinaria, 1973: Vienna. 32. P i l a r s k i W. & R o s k o s z T., 1957: Z ja w is k o uk rzyżow ien ia (sacralisatio) ostatniego kręgu lęd źw iow eg o u sam ic żubra — Bison bonasus (L.). Folia m orfol., 2: 109— 119. ' 33. P i l a r s k i W. , S e r w a t k a S., S w i e ż y ń s k i K. & W ę g r z y n M., 1967: N ew attem pts a f fix in g anatom ical m aterial o f large m am m als. A cta theriol., 12, 31: 353— 358. 34. P o l e i n e r R., 1932: Der anatom ische A u fb a u der E xtrem itäten beim E u ro päischen W isent (Bison bonasus L.) im V ergleich zum H ausrind. Diss. W ien. 35. P o p l e w s k i R., 1927: C elow ość bu d ow y i rozm ieszczenie staw ów . W ych. Fiz. W arszaw a. 36. P o p l e w s k i R., 1934: Rozw ażania teoretyczn e nad budow7ą dłu gich kości saków. Spr. z Pos. Tow . Nauk. W arsz., 26 (1933): 1— 14. 37. P o p l e w s k i R., 1948: A natom ia ssaków . 2: 1— 690. Spółdz. W yd. C zytelnik, Stockh olm . 38. P y t e l S. M., 1969: M orph ology o f digestive tract o f the E uropean bison. A cta theriol., 14, 27: 349— 402. 29. P y t e l S. & K r a s i ń s k a M., 1971: M orp h olog y o f the stom ach and intestines in h yb rid s o f European bison and dom estic cattle. A cta theriol., 16, 31: 471— -4 8 1 . 40. R a d o m s k i R., 1972: Staw y kończyn p iersiow y ch żubra, Bison bonasus (L.), w ro zw o ju postnatalnym . Diss. SG G W , W arszaw a. 41. R a d o m s k i R. & K o b r y ń H., 1969: P ow ik ła n e skręcenie (distorsio) stawu b a rk ow eg o żubra (Bison bonasus L.). M ed. wet., 25, 3: 144— 145. 100 F. Kobryńczuk 42. R e i s e r E., 1903: V ergleich en de U ntersuchungen über die Skelettm uskulatur von H irsch, Reh, S ch a f und Z ieg e: 1— 42. Diss. B ern — W ien. 43. R o s k o s z T., 1962: M orp h olog ie der W irbelsäu le des W isents, Bison bonasus ( L i n n a e u s , 1758). A cta theriol., 6, 5: 113— 164. 44. S i s s o n S. & G r o s s m a n J., 1960: The anatom y o f the dom estic anim als.: 20— 203. Saunders, Ph iladelph ia— London. 45. S o k o l o v I. I., 1972: P ostk ra n ja ln y j skelet p redstav itelej roda Bison. Tr. Zool. Inst., A N SSSR, 48: 198— 219. 46. S z t o l c m a n J., 1926: Żubr, jeg o przeszłość i p rzyszłość: 1— 104. Nakł. Centr. Zw . Pol. Stow . L ow ., W arszaw a. 47. S z u l c M., T r o p i ł o J. & K r a s i ń s k a M., 1971: D ressing percen tage and u tility valu e o f the m eat o f E uropean bison and dom estic cattle hybrids. A cta theriol., 16, 32: 484— 504. 48. Ś w i e ż y ń s k i K., 1962: T he skeletal m u scu latural system o f the E uropean bison, Bison bonasus ( L i n n a e u s , 1758). A cta theriol., 6, 6: 165— 218. 49. W ę g r z y n M. & S e r w a t k a S., 1961: Ligam entu m sacrotuberale latum bei Bison bonasus ( L i n n a e u s , 1758) und Bos taurus ( L i n n a e u s , 1758). A cta theriol., 5, 7: 73— 97. 50. V o k k e n G., G l a g o l i e v P. & B o g o l j u b s k i j C., 1961: A n atom ia d o m aśnih żivotnyh. 1: 39— 236. G os. Izdat. V yssaja Skola, M oskva. 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