Beruflich Dokumente
Kultur Dokumente
F.: Uher die \Terteilnng der spongiosadiehte im coxalen Femurende und ibre Becloo.
tung fr (lie Lehre vom funktionellen Ban des Knochens. Morph. Jb. 95 (1954) 3554.
Gesammelte Abbandlungen zur ftmktionellen Anatomic des BewegungsaPParates. Berlin: Springer 1965.
Atlas zur Biomechanik der gesunden nnd kranken Hfte. Berlin: Springer 1973.
mk cler Organismen. 2 Bde.,
Roux, W.: Gesammelte AbhandlUngen fiber ieldtmg5me
Leipzig: Engelmanfl 1895.
SCHMITT, H. P.: Uber die Beziehung zwischen Dicbte und Festigkeit des Knoelrnns am Beiepiel
des mensebbeben Femur. Z. Anat. Entwicklgm 127 (1968) 124.
Smuco, K.: Uber die Hftpfanne. I. Mitt. Z. Morph. Anthrop. 17 (1915) 325356.
TmbMANN, B.: Die Beansprueltung des mensebliehea Huftgelenks. Hf. Die Form der Faeies
lunata. Z. Anat. Rntwickl.gesCll. 128 (1969) 329349.
Zur Lokalisation von clegenerativea eranderungen am Femurkopf bei der coxartlirose. Z.
PAUWELS,
WALDEYEE,
Oliver Rieppel
Dr.
I
With 14 figures and 2 tables
HnNz.JUReEN KURRAT
Anatomiscbes Institut
der Univereitt zu Koln,
Lindenburg
D 5000 Kbln 41
A. Introduclion
798
0. Rirppn: A Functionai
Interpretation of the Varanid
Dentition
Varanus is a lizard genus adapted to relatively large prey. This is reflected in the me
chanics of its head. Consequently, no generalisations will be attempted concerning
the amphikinesis among Modern lizards from a mechanical point of view.
The study reported on here is part of a thesis presented to the Faculty of Natural
Sciences at the University of Basel in partial fulfillment of the requirements for the
degree of a Dr. phil.
799
F5]H 17145, 22084, MCZ 1066); T. eec/lotus (FMNH 22354); V. presinus (MBS 10 504*); V. re
dice//is (MBS 18874, FMNH 98947, 131538); J7 so/voter (MBS 3713*, 3736*, 6138*, 10755*,
osteol. 5028, 5878); V. stone (MBS 17891*); V. venus (AMNH 28698, 73361, FMNH 1492).
Abbreviations: AMNH. American Museum of Natural History, New York; BMNH British Muscum of Natural History, London; FMNH, Field Museum of Natural History, Chicago; MBS,
Natural History Museum,Basel, osteol, = osteological collection; MCZ Museum of Comparatie
Zoology, Cambridge, Mass.
The decomposition of the kinetic lizard skull into various functional components
goes back to VER5LUY5 (1912). He subdivided the lizard skull into the oeeipfl
segment and into the maxillary segment which represent the braincase and the
dermatocranium respectively. The braincase is movably suspended within the des
matocranium on its paroccipital processes posterodorsally and on its basipterygotd
processes anteroventrally (Fig. 1). As the maxillary segment moves up and down
around the braincase, the cartilaginous processus ascendens which projects from the
anteromesial tip of the supraoccipital bone slides back and forth in a trough on tIlE
caudal edge of the parietal. This is the metakinetic joint.
The metakinetic joint is regularly fused up through the development of a sutil&
contact between the front edge of the supraoccipital and the hind edge of the par1e
800
0.
thErruL:
801
pterygoids. Caudally the quadrates (quadrate
units) link the parietal unit to the
basal units. The quadrates receive the caudal
tips of the supratemporals and of the
squamosals in two collateral, shallow ai1ic
ular facets on the dorsal surfaces
of their
cephalic condyles. The epipterygoid bone is
the last component of the maxillary
segment of VER5LUY5 (1912). The foot of the
epipterygoid rests in the pterygoid or
coluuxellar fossa on the dorsal surface of the
pterygoid. The dorsal head of the epi
pterygoid is bound to the alar process of
the prootie and to the deseensus parie
talis
by fibrous ligaments.
B. The mandibular joint
ju
Fig. 2. The snout complex of T7eraans bciiyeleiisis (MBS. 2410). Key to abbr,viatioflh
as in Fig. 1. Scale equals 10 mm
par.c
gI.s
the
the frontal lies below the upper edge of the front end of the parietal. Towards
art
lateral edges of the suture, the parietal forms distinct anterior lappets which
eulate in lateral notches on the hind edge of the frontal. The ventral hinge, the hypo
lieS
kinetic joint, consists of a lateral and a mesial component. The mesial cmupoildhlt
the
een
betu
between the palatine and the pterygoid. The lateral component lies
the
ectop
Of
teryg
oid
maxifia and jugal of the muzzle unit on the one hand and the
basal unit (see below) on the other hand.
(1962)
The maxillary segment is broken down into five components by FnazZETTk
unit
The muzzle unit described above links the parietal unit dorsally to the basal
11toit0
ventrally. The parietal unit is made up from the parietal, supratemPorals
ecto
frontals and squaiuosals. The basal units are made up from the ptcrygoids and
I mc
S.
o
n
a
r
t
m
C
802
The structure of the mandibular joint results in a lateral rotation of the lower jaw
around its long axis as it is depressed and in a mesial rotation around the long axis
as the lower jaw is elevated. Such a rotation of the lower was proposed by BRADLEY
(1903) on the basis of similar arguments.
When the jaws are fully adducted the dentary (as well as the maxillary) teeth pohit
slightly inwards. As the jaw is depressed relative to the long axis of the quadrate it
rotates around the dceper mesial side of the glenoid surface: the teeth are rtated
outward. They thus assume a fully erect position so that they hit the prey with their
tips instead of with the lateral surfaces of their crowns. Convcrsly, as the jaw is ele
vated relative to the iong axis of the quadrate, it rotates aronnd the higher lateral
side of the glenoid surface. Consequently the teeth rotate back inward again and
thus produce a clamping effect on the prey held between the jaws.
The second feature of the mandibular joint which has to he emphasised is the strong
anterior elevation of the glcnoid surface on the articular bone. Anteromesial to the
glenoid surface the surangular is also elevated and forms a prearticular condyle. Thc
anterior elevation of the glenoid surface provides an increase of area on which the
lower jaw can he rotated during depression (TmwcK3ionToN, 1976). But it also
locks the lower jaw firmly against the mandibular condyle of the quadrate whea
retractive forces are transmitted from the lower jaw through the mandibular joint
to the quadrate. As it will be shown below, the depression of the mnzzle unit is func
tionally important during the swallowing process in Varanus. Depression of the
muzzle unit results from retraction of the basal unit which can be achieved by back
w-ard rotation of the quadrate around its dorsal suspensioii. Jaw adductor muscles
which insert obliquely into the lower jaw will exert a force which can be broken down
and into a
into an adduetive component working at a right angle to the lower
jaw.
retractive component which works along the long axis of the lower
jaw
803
and the quadrate. The small amount of
translational movement possible results
from
the elasticity of the ligaments of the
joint capsule.
F. The Analysis of Feeding Movements in
Varanus bengaknsjs
The present account deals with Varanus
Sen galensis of which four live specimens
were available for study. The animals
were regularly fed on mice. Five
feeding se
quences were filmed with an ordinary
super-S movie cassette at a speed of 40
frames
per second. Two further feeding
sequences were recorded on a 16 mm
black and
I white film at the speed of 64 frames
per second. An eighth feeding sequence
was
filmed with an X-ray camera at a
speed of 48 frames per second. Cursorial
obser
vations were also made on one specimen
of Varanus exantheniat ices albigularis
and
on one adult specimen of Varanus komodoensis.
When Farce us bengaiensi.s locates prey, it
immediately starts to pursue it, seizing
it wherever possible without any
preference for any particular part of the
body. The
pi.ey is vigorously shaken and
banged against the substrate, rocks,
logs,
etc. until
it is immobilized or dead. The
JTaran mar then usually drops the prey and
starts
searcli
iag for the preys head. It has been
shown by Loop (1974) that varanids
feeding on
small mammals show a signifieamrt
preference to swallow- the prey head first.
Once
the Varaijus has started to swallow- its
prey, kinetic inertial feeding (GAN5,
1961,
1969) is observed. thuS (1961:
218-219) describes kinetic inertial feeding
as fol
lows: The jaw-s release the prey
quite suddenly and the entire head shifts
forward
and laterally, then bites in a newposition. The inertia of the prey reduces
prey
shifting. The head continues to sluft
from side to side in a w-alking sequence
that
graduajly produces ingestion of
the prey
The sequence results in a pushing of
the head over the prey rather
than drawring the prey into the month.
GANs (1961)
i5 FRAZZEYn who noted that
the lateral excursion of the forward
stroke of the
head does not occur in all lizards
during kinetic inertial feeding. In Varanus
benga
hens the lateral excursion of the
head occurs only occasionally It may also
be
noted
that Jaranns bee gaiensjs combines
the kinetic inertial feeding method witli
the
pos
Sibihty to draw- or rather to push the
prey back into the throat through jaw
mobility
(see belowr)
-
804
Vara.nid
Dontition
805
48
N--
..-:
A
16
7/
50
--
8Uspectwrn
(MB$.
20
N
A
N
-1
/7A
extremely elevated position on frame 16, that is before the mouth is maximally
opened. Maximal mouth opening is achieved on frame 40. It is essential to note that
the
opening of the mouth is brought about by upward rotation of the head around
prey.
the
occipito-atlantal joint. The mandible remains stationary and supports
As the mouth is maximally opened, the forward stroke of the head characteTiSt1
an
of the kinetic inertial feeding follows. Initially, the head moves very rapidly in
across
anterovesitral direction. The gape is held constant and the head now shifts
phase
directed
a
is
there
horizontally
the prey (Fig. 5 B). Following this movement
oCCI
the
around
of the forward stroke of the head. The head now rotates downward
gape
the
pita.l conthle, and simultaneously the snout is depressed. This closure of
the
of
brings the maxillary teeth into contact with the prey again. The final phase
snoUt
forward stroke of the head is directed anteroventrally again. The tip of the
around
now closely approaches the substrate. The head is strollgly rotated downward
strong
This
the occipito-atlantal joint and the muzzle unit is being strongly depressed.
and
mouth
and final depression of the muzzle unit pushes the prey further into the
down the throat.
99
\
\
\
\
\
Fig. 7. The application of the quadrie crank chain model to illustrate the movements
as they occur in the head of Varanus ben galensis during feeding
806
0.
Varanus bengalensis does not perform a kinetic inertial feeding cycle with each
down the throat is shown. Tho
orbit. Often a different method of pushing the
animal presses the prey against the substrate. It opens the mouth, pushes hard against
the prey and closes the jaws tightly again, thereby depressing the muzzle unit.
During the kinetic inertial cycle, it is advantageous for the animal not only to
maximally accelerate the head but also the maximize the excursion of the head
during the forward stroke. The longer the excursion is, the more is the head likely to
shift across the prey. Varanid lizards, which are adapted to relatively large prey,
show an elongated neck which results from an increase in number of cervical verte
brae and from an increase in length of the individual cervical vertebrae (HOFE5TETTER
and GAse. 1969. and Fig. 6).
Fig. 7 shows the applieaioii of FRAzzETTAs (1962) quadrie crank chain model to
the analysis of another kinetic inertial feeding sequence of Varanus bengalensis as
it is observed on the X-ray film. The model as it is drawn in Fig. 7 does not correspond
to the actual movements of the head which are generated in the neck region. Rather
it is used in order to allow measurement of degrees of rotation. Joint X is considered
to be the fixed reference point within the quadrie crank chain (FAZzETTA, 1962),
and for the sake of easy comparison, the parietal unit is always drawn in a horizontal
position. The analysis shows that starting from the rest position (Fig. 7 A: frame 1)
the muzzle unit is elevated for 9 while the mouth is maximally opened. The qua
drate is protra etc ci for 210. The angle between the basal unit of the skull and the
dentary is 48 at the stage of maximal jaw opening.
Following the forward stroke of the head, the skull is rotated downward around
the oeeipito-atlantal joint while the muzzle unit is depressed. At the stage of maximal
150
depression of the muzzle unit (Fig. 7 D: frame 99) the latter has rotated around
compared to the stage of maximal muzzle elevation: thus the muzzle is 6 lower than
it is at the rest position. Correspondingly, the quadrate has rotated backward around
27 compared with the maximally protracted stage. This is 6 further backward than
it is at rest position. The analysis shows that during the final stages of each orbit,
the snout is depressed below its rest position, and that this final muzzle depression
807
prey
I-
808
I;
809
/1/
4!
I
/
/
4
/
L
I
4.
tooth. But due to the development of plicidentine, the replacement teeth of platY
notans develop in an interdental position. This results in the discontinuity of func
tion at any given site in the jaws of Varanws or other platynotans, a functiofl
disadvantage which must be compensated by a functional advantage of plieidentiTle
Varanus is a pleurodont lizard. The tooth becomes ankylosed to an obliquely sloping dental shelf on the tooth bearing bone. The teeth of Varanus show widely flared
bases which in some cases may be overgrown by bone (adult specimens of Var
niloticus) what gives the teeth further support. Striations on the bases of the teeth
result from the formation of plicidentine.
The crorn of the adult tooth of Varanus is a blade like structure, laterally C01fl
pressed with a recurved tip and an anterior and posterior serrated cuthng edge. It
is curved in two directions. The transverse section through a dentary tooth of Vart
saleator (Fig. 9) shows the lingual wall of the tooth base to buldge mesially, produdhh1
810
811
Table 1
Same values as in Table 2, but for the dentaries of two specimens of Ttcronus solvotor
with complete dentitior.. L: left, R: right
MBSspecimen
numbers
Tooth positions
10755 H
74
73.5
10755 L
76
72
6 138 H
77
78
71.2
6138L
82
80
77
58
62
10
60
59.8
59
60
48.5
46.5
60
55
66.2
66.8
63
73
64
63
73.8
60
67
71
63
67
72
55
57
Table 2
Degrees of reeorvation of the teeth along the tooth row in seven specimens (loft maxillaries) of
Voronus solvotor. The angle is measured between the tangential line indicating tho direction into
which points the tip of the tooth and a horizontal line representing the lower edge of the maxillary
bone (Fig. 23). An increase of the recurvation results in the decrease of the numerical value of the
angle
MBS-specimen
numbers
Tooth positions
1
3713
69.5
78.8
60.8
55.5
3834
79
74
53.2
67
10 755
78.8
5028
85.5
3 835
74.5
79
85.4
69.5
77
76
74.5
77.8
69
59.5
5901
68.5
63
67.2
55
50.5
10
65
11
60.3
61
57.2
55
Fig. 11. Recurvation of the teeth along the tooth row in, top: maxilla of Varenns .901voter (MBS. 5878), bottom: dentary of Forming indicus (MBS. 11099)
43
59
51
12
45.8
59
67.5
53.2
9
56.9
58
76
74
5878
66.5
65
54.2
62
53.5
53
56
meso
62
into a position so that it may hit the prey with its tip. Yet it is critical for the tooth
to hit the prey with its tip. In doing so, maximal stress is exerted on the preys
surface and its penetration is more likely to be accomplished. A second critical
factor of tooth function is the axial loading of the tooth. This can only be achieved
if the tooth hits the prey with its tip. Axial loading reduces the bending moments
to which the tooth is subjected during the strike.
To bring a revurved tooth into an erect position it is most economical to rotate
the tooth along the circumferential line which represents its longitudinal axis. The
stronger the tooth is recurved, the closer will the center of rotation approach the
posterior cutting edge of the tooth. The lower jaw is hinged on the mandibular joint
which lies far posterior to the tooth row. Yet, the posterior dentary teeth are closer
to the mandibular joint than are the anterior teeth, and hence the posterior tEeth
show an increase in degree of recurvature which compensates a decrease in absolute
meta
occ
hy
Fig. 12. Loft maxillary dentition of Voronns solvator (MBS. 10755) rotated around the
]iypokinetie joint only (by; solid line) or around the occipito.atlantal joint only (0cc;
line). The rotation around the hypokinetic joint matchos the roeurvation of
the
teeth. Abbreviations: hy, liypokinetic joint; moso, mesokinetic joint; motn, rnota
dasliod
kinotik joint; oec, occipital condylo. Scab equals 10mm
size.
812
0.
0.
The maxilla is hinged on the occipital condyle far back behind the functional
tooth row, but it is also hinged on the mesokinetic axis. Rotation of the muzzle unit
is made possible through the combination of the hypokinetic and the mesokinetic
joints (Fig. 12).
As the hypokinetic joint lies immediately behind the maxillary bone, a greater
degree of reeurvaftre of the maxillary teeth can be expected in comparison with the
dentary teeth. This is indeed the case, and again the more posterior teeth approach
ing the hypokinetie joint more closely show a relative increase in recurvature but a
decrease in absolute size in comparison with the anterior teeth. The increase in degree
of recurvature may be critical for the function of the maxillary teeth since it is the
upper tooth row which is more active during the feeding process.
The combined rotation of the upper jaw around the hypokinetie and mesokinetie joints
and around the occipito-atlantal joint not only allows an increase in degree of recurva
ture, but it also alters the line of action of a given tooth (Fig. 13). The line along which
a tooth tip of the upper tooth row will travel was studied with a simple mechanical model
of a TTaranus head, constructed according to the quadric crank chain model of FRAz
ZETTA (1962). If the maxilla rotates around the occipito-atlantal joint only, eery
maxillary tooth tip will describe the upper left quadrant of a circle line (with the
snout pointing to the left, Fig. 13 A). The gape required to bring the tooth into a
fully erect position is wide, and the tooth points backward at a very early stage of
jaw closure already. Thus the tip of the tooth may point backward as it hits the prey.
It may hit the prey with its anterior cutting edge. Moreover, the line of action of the
tooth contains a forward component of the strike ST which will affect the direction
of the resultant force produced by the tooth as it hits the prey. Thus, if a maxilla
813
J7eranus
Fig. 13. A: Maxillary tooth of Vera, us rotated around the oeeipito.atlantal jont only.
B: Maxillary tooth of Vereneis rotated simlIltaneoIsly around tIle oeeipitoatlt
joint and around the Itypokinetie and mesokinetie joint. ST: Forward componen f the
strike
I.
814
This will certainly increase the stability of the tooth base. The formation of pliciden.
tine in platynotan lizards may indeed be correlated with the large. size of their teeth
(PEnn. 1968).
To study the stress distribution within the tooth base, plate models were cut from
plates of Araldite B to the size 35 times the size ofthe original tooth. Their shape
represented transverse sections at various levels through the functional tooth. The
loaded and inspected with light passing through crossed polaroids
tooth models
between which the models were mounted in an iron frame. Isoclinics appear as dark
lines on the model connecting these parts of the model where tbe directions of the
principal stresses are either parallel or perpendicular to the optical axis of the pola
roids. If the polaroids are rotated stepwise around 90, a series of sets of isoelinics
will be observed wandering across the model. Each set of isoclinics corresponds to
each step of rotation of the optic axis of the polaroids. The observed sets of isoclinics
can be superimposed on an outline drawing of the model. and with the indication
of the correspondiug optical axis of the polaroids the direction of the principal stresses
all across the model can he found. The isoelinics by themselves do not differentiate
between tensile and compressive stresses, but these can be inferred by pressing ones
nail into the edge of the model. If the isoelinies are attracted toward this
finger
stressing edge, tensile stress is indicated. If the isoelinics withdraw from the stressing
edge, compressive stresses are indicated (Nageiprobe: FPPL and MNdH, 1959). Note
that for any point of the model the direction of one principal stress (tension or corn
pression) is by definition perpendicular to the direction of the other principal stress
(centrifugal stress resulting from the deformation of the model).
Models of transverse sections through the functional varanid tooth were loaded
at different angles which resulted in changes of the directions of the resulting prin
cipal stresses (Fig. 14). The results indicate that upon axial loading (Fig. 14 A), ten
sile stresses result on the lingual side of the tooth base while compressive stresses
build up on the labial side of it. Moreover, upon axial loading the one direction of
the principal stresses (tension and compression) exactly parallels the direction of the
dentine lamellae of the plicidentine (Fig. 9 B) and meets the tooth bearing bony 5Wface at right angles anywhere within the tooth base. The other principal stress (cen
trifugal stress resulting from deformation) by definition runs parallel to the surface
of the tooth bearing bone. Therefore, no shear stresses will occur along the site of
ankylosis, and the dentine lamellae will be loaded in pure tension and compression
815
were
in5.
th
bp
is presented in a transverse
section under varying loads. For
further explanations see text.
Abbreviations: th, Theka; bp, Basalplattc; ling,
lingually;
lab, labially
pro
816
Summary
817
A cinradiographic analysis of the feeding movements iii l7m-enus bengolenais produced the
following results. The mouth is opened by raising the head (upper jaw) rather than by lowering
the lower jaw. Starting from the resting position, the muzzle unit is elevated around 9 reintive
to the rest of the skull during jaw opening; the quadrate swings anteriorly around 21. During
jaw closure, the snout is depressed around 15 relative to the rest of the skull, hence 6 beyond
the resting position. The quadrate swings backwards around 27.
Amphikinesis is interpreted as allowing a stronger posterior recurvature of the maxillary teeth
in Verensis. This increases the holding effect of the teeth without increasing their length, an adap
tation of Vorcoius to capture relatively large prey.
The formation of plicidentine (dentino infolding) in the teeth of Vorenus increases the surface
of attachment of the teeth on the supporting bone. Moreover, the dentine lamellne take up ten
sile anti compressive stresses along their long axes upon axial or vertical loading of the teeth.
Tho slope of plourotlonty is modelled so as to minimalize shear stress on the surface of anky
losis upon axial or vertical loading of the teeth.
Zusammenfassuag
\4tiilirentl ties Frehoktes dffnet Voronus benqeien.sis das Maul rioreli Hebung dos Oberkiefers
und nicht dureh Senkong ties Unterkiefers. Wiihrontl ties Offnens tier Kieter wird the Sc loiauze
urn 9 relat iv zum Rest ties Sclikdels angeliohen; tlas Quadratum drelit sick tiahei um 21 nach
verne. Wkhrenti tier ScldieBung tier Kiefer wird die Sehnauze urn 15 reiativ ztim Rest des Sehd
dels gesenkt, tI. h. sic senkt sich urn 6 fiber die Ridielage hinaus. Das Quadrnttim dreht sich dabei
um 27 nach hinten.
Die ampliikinetiscbeii Bewegungen im Sehddel von Coronet eriatthten eine stdrkore Rdekwilrts
krfiminung tier Zkhne. Dabei wird tier Halteeffokt tier Ziilme vergrd6ert oline deren abseiute Lange
zo steigern.
(lie
Die Bildung von Pheidentin erhdht the Atifingefidehe cmos Waranzahnes. Zndem aehmen
eatngen
Druekspannu
untl
Ztigthe
Ziihne
tier
Dentinfalten bei axialer otler vertikaler Belastung
lang ihrer Liingsaehse en f.
Zahnc
Der Pleurotientiewinkel ist so eingeriehtet, tiall bci axialer other vertikaler Belastung der
t
werden.
Seherkrafte entiang der Atifiagefiiiche minimahsier
References
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(1968)
116
Postilla
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111 (1962)
FEAzzETTA, T. H.: A functional consideration of cranial kinesis in lizards. J. Morph.
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Kfinstiergasse 16
CR
28 73 19.
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