Ontogenesis of the Human Temporomandibular Joint

1. Development of the Condyles

LOUIS J. BAUME
Institut de Medecine Dentaire, Universite de Geneve, Geneva, Switzerland

Endochondral growth activity of the mandibular condyles, in contrast to epiphyseal

plates of long bones, is responsive to induced stresses.1 2 This unique behavior of
condylar cartilage is accompanied by peculiarities in phylogenesis, histogenesis, and
metabolism which differ from the pattern of epiphyseal growth cartilages: (1) Phylo-
genetically, the temporomandibular joint represents a new acquisition of the order of
mammals; it replaces the articulation of higher reptiles between the articulate and
the quadrate, which, in mammals, become ear ossicles (malleus, incus). (2) Histo-
genetically, condylar cartilage accrues by surface apposition in a peripheral fibro-
cartilage, whereas epiphyseal plates proliferate interstitially. (3) Metabolically, epi-
physeal cartilages show much greater respiratory activity than found in articular car-
tilages.3 The hormonal regulation of condylar development also differs from epiphyseal
plates.4
The view has been expressed that this unique position of condylar cartilage among
endochondral growth centers may be explained on the basis of its specific embryonic
origin.5
A review of the literature recently presented by R. P. Cousin6 reveals only scanty
and contradictory information regarding the embryogenesis of condylar cartilage. Scott
and Symons7-to our knowledge-are the only textbook authors to present authentic
human material showing the first appearance of "secondary" condylar cartilage at the
48-mm. stage as a fringe on the dorsal end of the membranous dentale. This conforms
with the observations made by Mundaca,8 while Kjellberg, 1904,9 Vinogradoff, 1910,10
Harpman and Wollard, 1938,11 Giardino, 1950,12 Gasparini and Marchetto, 1954,13
place this event at much later stages, from 85 to 120 mm., attributing to the lateral
pterygoid muscle a decisive role. Politzer,14 therefore, states that the role of the sec-
ondary cartilage in mandibular morphogenesis still warrants further investigation.
Considering the great impact that a proper understanding of the physiology of the
mandibular joint has on oral surgery in general and dentofacial orthopedics in partic-
ular, we thought it worthwhile to pursue its morphogenesis in adequate human ma-
terial. This includes 25 fetuses from 15 mm. crown-rump length to full term and
5 infants: two days, six months, nine months, and four years of age. Analysis of the
serial sections has shown that 2 more or less independent blastemata-namely, the
condylar blastema and the temporal blastema-are operating in the histodifferentiation
of the joint structures.

This investigation was supported in part by a research grant from the Swiss Odontostomatological
Society.
1327

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1328 BA UME J. D. Res. November-December 1962
The prenatal and postnatal development of the entire temporomandibular joint will,
therefore, be reported, partly in collaboration with K. Hiiupl in a series of 3 papers in
the chronologic order of events: (1) the development of the condyles, (2) the develop-
ment of the temporal components, (3) the postnatal growth and transformation.
MATERIAL AND METHODS
Data of material and methods used for this study of condylar histogenesis are seen
in Table 1. Reference is made to a corollary study5 that gives low-magnification micro-
photographs of the sections here described.
TABLE 1
DATA OF MATERIAL* AND METHODS USED

Crown- Approx. Plane of

No. Rump Fetal Age Serial
Length (Week t) Sections
(Mm.)
1........ 15 6th-7th Sagittal
2........ 19 7th Coronal
3........ 24 8th Coronal
4........ 25 8th Sagittal
5.3.5...... 35 8th-9th Sagittal
6........ 35 8th-9th Coronal
7........ 44 9th Coronal
8.......... 50 9th-10th Coronal
f Coronal
9........ 55 10th
JSagittal
10 ....... 60 11th Coronal
11........ 72 12th fSagittal
J Coronal
* Immediately fixed in formalin, imbedded undecalcified in paraf-
fin, sectioned serially 5-8 p; each twentieth section stained with H.
& E.
t Crown-rump length measured before fixation. Fixation reduces
length by 2-3 mm. (Hamilton, Boyd, and Mossman, 1959).
According to B. M. Patten (1948).

RESULTS
At the 19-mm., crown-rump stage, the bony dentary* originated from bilateral intra-
membranous ossification centers in the integumental portions adjacent to the anterior
end of Meckel's cartilage (Fig. 1).
At the 24-mm. CR stage, these bilateral osseous centers have expanded posteriorly
along Meckel's cartilage toward the angle formed by the inferior dental nerve and the
lingual nerve (Fig. 2). The orientation of this very thin bony plate was best appre-
ciated by its relation to the primordia of the masticatory muscles, whose development
precedes bone formation. Coronal serial sections showed that the median pterygoid
muscle, partly connected with Meckel's cartilage, ended distant from the lower border
of the dentary; this, in a bend, tended to ossify in the direction of the muscle insertion
(Fig. 3A). At a more posterior level, below the auriculotemporal nerve (which
branches off the bifurcating mandibular and lingual nerves), the origin of the lateral
pterygoid muscle in its most primitive stage can be seen (Fig. 3B). The adjacent
* The term "mandible" will be used after formation of the condylar cartilage head.

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 Fic 1.-Coronal section of a 19 mm. crowxn-rump human fetus, showing bilateral intramembranous
ossification centers of the dentaries (D) in the paramedian integument ol the mandibular processes
St Stomodeumr. (Orig. mag. X36.)

FIG. 2. Sagittal paramedian section ot a 24-mm. crown-rump human fetus, evidencing the growth
of the dentary (D) in dorsal direction by peripheral bone apposition. MC =Meckel's cartilage;
T _ tongue. (Orig. mag. X36.)
FIG. 3. Coronal serial sections through the head of a 24-mm. CR human fetus showing, in an antero-
posterior sequence, the various sites of the primordia of the masticatory muscles and the ossifying
dentary. m = masseter, nipt = medial pterygoid muscle, t = temporal muscles, dn inferior dental
nerve, In lingual nerve, atn auriculotemporal nerve, P palatine process before fusion (posterior
end), TR = tubo-tympanic recess, MC Meckel's cartilage D = rlentary (mandible), and x = con-
dylar blastema. (Orig. mag. X36.)

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1330 BA UHE J. D. Res. November December 1962
mesodermal aggregation thus constituted the future area of the posterior portion of the
ramus and condylar process; it may be termed "condylar blastema."
At the 35-mm. CR stage, fusion of the palatine processes was proceeding. This for-
mation of a secondary palate was associated with a considerable downward displace-
ment of the mandible-tongue complex (Fig. 4). There were, however, no morphologic
indications that articulationing movements were involved. The chondrodesmosis be-
tween Meckel's cartilage (called articulatee" at its posterior broad end) and the quad-
rate, though present, lacked the structural differentiation of a functioning joint (Fig. 5 ).
The condylar blastema was found bilaterally in coronal sections which showed the

FISs. 4 8. Coronal serial sections of a 35-mm. crown-rump human fetus are shown. Fig. 4: Over-
all aspect of a coronal section, showing fusion of the palatal processes with nasal septum concomitant
with downward development of the tongue-hyoid complex. (Orig. mag. X36.) Fig. 5: The primordial
articulation of the mandibular arch formed by the end of Meckel's cartilage articulatee) and quadrate
in a region behind the outer ear anlage. A = articulate end of Meckel's cartilage, Q = quadrate,
o = otic capsule, RC - Reichert's cartilage of the hvoid arch, and ST ossification of the squamo-
temporal plate. (Orig. mag. X36.) Fig. 6: Over-all aspect of a coronal section illustrating the topog-
raphy of the condylar blastemata (x). Pch = prechordal plate of the cranial cartilage primordium,
OS = orbito-phenoid primordial cartilage, OR - oropharynx, and T ossification center of squamo-
temporal bone. (Orig. mag. X 13.) Fig. 7: Condylar blastema is shown at higher magnification. (Orig.
mag. X55.) Fig. 8: Immature myoblasts (arrow) of the lateral pterygoid muscle forming the medial
zone of the condylar blastema. D = distalmost ossification site of dentary. (Orig. mag. X210.)

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lVol. 41, No. 6 ONTOGENESIS OF TEMPOROMANDIBULAR JOINT. 1. CONDYLES 1331
oropharynx below the prechordal plate and flanked by the orbitosphenoid cartilages
which were perforated by the optic nerve (Fig. 6). The condylar primordium appeared
as a rectangular area of condensed cells, laterally to and above Meckel's cartilage and
adjoined medially by the lateral pterygoid muscle. In this region, a new skeletal ele-
ment in the form of the squamotemporal rod appeared; it extended from the post-
auricular region toward the upper masseter origin (Figs. 5-7). The dentary terminated
as a semilunar bony spicule in the center of the dense agglomeration of mesodermal
cells of the condylar blastema (Fig. 7). The lateral pterygoid muscle medial to the con-
dylar blastema was still little developed, as it consisted of immature myoblasts (Fig. 8).
At the 44-mm. CR stage, the dentary started to broaden, while posteriorly it further
accrued in height and length. Expansive rapid osteogenesis was observed at 3 sites: at

FICS. 911. Cor nal sections of a 4 -m . crowrn-rump human fetus show the thre sites of an
increased growth of the dentary. Fig. 9: Ra~pid osteogenesis occurring at the lower border, where the
masseter originated, and at the coronoid process. (Orig. mag. x36.) Fig. 10: A bulk of newly formed
boeappearing at the base of the condylar process.
blastema. muX mandibular nerve indicates site of The
dentary accrued rapidly within the condylar
future mandibular foramen. (Orig. mag. X36.)
Fig. 11: At higher magnification of Fig. 10, the pattern of intramembranous ossification prevailing
in the condylar blastema. A new mesodermal cell condensation (x) appeared at the lateral aspect
of the condylar blastema, below the temporal joint primordium (T). (Orig. mag. X 10.)

the lower border, the upper coronoid end (Fig. 9), and the basal portion of the con-
dylar process (Fig. 10). The posterior end, however, remained thin and bent inward.
The processes involved, as was seen at higher magnification, conformed with the pat-
tern of intramembranous ossification (Fig. 11). Concomitantly, further differentiation
of the condylar blastema took place; laterally in its outer layers, below the temporal
bone, a new zone of mesodermal condensation was forming.
At the 50-mm. CR stage, the entire periosteal circumference of the dentary showed
deposition of new bonle, but, at the 3 sites already previously noticed, it reached an
extraordinary extent. In the basal and coronoid portions, this growth spurt resulted
predominantly in a lengthening (Fig. 12), while, at the condylar end, widening was
engendered (Fig. 13). The entire bulk of the newly formed tissue consisted of imma-
ture fibrous bone (Fig. 14). The germinative layer at the cotndylar blastema was still
very wide, containing copious mesodermal cells, particularly at the laterosuperior as-

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/ 3,32 B UJ1I Ef 1J. 1). Res. Xo)veiubcr- Deceniber 1962
pect, which foreshadows the future cartilage center. A wide, loose mesodermal tissue
intervened the area below the squamotemporal bone.
At the 55-mm. CR stage, as seen in the sagittal section of Figure 15, the condylar
process is now topped by a condylar head, but with no cartilage present. The condyle
lay at a higher level than the coronoid process. At its dorsosuperior aspect, the meso-
dermal cell condensation of the condylar blastema xvas very wide. For the first time,
processes of modeling resorption were found in the periosteum of the ramus. It was
noted that no temporal elements of the future joint were present at this level.
The coronal section (Fig. 16) also showed that the dorsal end of the condylar
process had developed into a typically rounded condylar head. It consisted entirely of

Fios. 12 14. -Coronal serial sections through the developing dentarv of a 50-mm. crown-rump
human fetus are shown. Fie-. 12: The coronoid process growing above the level of the suture of the
temporal (T) and zvgomatic (Z) bones. At its top and the lower border, sites of expansive osteo-
genesis were seen (Orie,. mag. x36.) Fig. 13: Intramembranous ossification of the condylar process,
showing an extraordinary extent. (Oiig mag. X55.) Fig. 14: At higher magnification, the center
of rapid osteogenesis appearing at the laterosuperior aspect of the blastema (X). Note some osteo-
blastic activity of temporal bone rod. (Orig. mag. Xl10.)

young bone which contained a large amount of round osteoblasts in a copious eosino-
philic ground substance and a reduced amount of fibers (Fig. 17). This rapidly formed
bone tissue thus started to lose identity. At the dorsolateral aspect of the head a carti-
lage blastema became evident.
At the 60-mm. CR stage, the decisive event of histodifferentiation of secondary carti-
lage at the condylar head could be observed (Fig. 18). But, simultaneously, cartilage
also appeared at other sites of the mandible. In the anterior portion, lateral to the
symphysis, M/Neckel's cartilage was incorporated into the mandibular body by a process
of endochondral ossification (Fig. 19).
Cartilage also appeared at the 3 sites where previously rapid bone growth had
occurred (Fig. 20). At the lower border of the ramus, osteogenesis passed a stage of
secondary cartilage formation (Fig. 21). The fully differentiated chondrocytes were
irregular in size and arrangement; the basophilic matrix was abundant and contained
coarse fibers btit little hyaline. In its older )ortions, this secondary cartilage underwent

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Vlal. 41, NV. 6 0ONJOGENEiSIS OF 1'E.!J'OROS1.4 XDII3UL.4I JOINTX. 1. CONDIFLES 1333

calcification and eventually, resorption by marrow elements (Fig. 22); direct ossifica-
tion, producing a chondrosteoid tissue, also occurred.
A huge ovoid mass of more regular cartilage was seen on the dorsal end of the coro-
noid process (Fig. 23). Its base, however, consisted of newly formed trabecular bone,
while in the more proximal sections calcified cartilage was present. All these phenomena
were indicative of the great rapidity with which the ramus developed in an upward
and backward direction.
At the end of the condylar process, a fourth independent center of highly organ-
ized cartilage had appeared (Fig. 18). It occupied, in the form of a spheroidal mass,
the area that intervenes between Meckel's cartilage and the squamotemporal rod,
where the condylar blastema was previously found. Careful analysis indicated that the

Fws. 15-17. The development of an osseous condxlar head at the 55 mm. crown-rump is shown.
Fig. lt: A sagittal section showing the offshooting development of an osseous condylar head (H),
still topped by the condylar blastema (x). (Orig. mag. X30.) Fig. 16: The coronal section showing
the formation of a primary osteal head of the condyle, which develops in a laterodorsal direction.
Note relative aplasia of the temporal bastema. (Orig. mag. XS0.) Fig. 17: At higher magnification,
the persisting condyJar blastema of a large mesenchymal cell condensation (x) at the laterodorsal
aspect of the condvlar head; this was the center of imminent chondrogenesis. (Orig. mag. X98.)

cartilage-producing cells were located distant from the pterygoid insertion in the dorso-
lateral mesenchymal covering of the head (Fig. 24). The upper medial portion of this
covering produced a fibrous layer, which also assured the connection with the lateral
pterygoid muscle (Fig. 25), while, in the basal periphery, periosteal osteogenesis was
proceeding (Fig. 26). The condylar cartilage thus formed a cone whose top of the
germinative cells faced the squamotemporal bone and whose basis containing the oldest
chondrocytes faced Meckel's cartilage.
The squamotemporal bone opposite the newly formed condyle showed some osteo-
blastic activity (Fig. 18), but the differentiation of the temporal joint elements took
place in a more posterior region.
At the 72 mm. CR stage, serial sections made in both the coronal and the sagittal
planes disclosed the following events as far as they concern mandibular development:
At the basal portion of the ramus, secondary cartilage formation had completely
ceased (Fig. 27). Previously formed cartilage had disappeared, partly through resorp-

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1334 BeA UME J. D. Res. November-December 1962

tion, partly through ossification into a chondrosteoid tissue. Regular osteogenesis pro-
ceeded within the periost. At the coronoid process, secondary cartilage formation also
tapered off and was succeeded by rapid periosteal bone formation. Masses of a chon-
drosteoid tissue were found more mesially in the core of this apophysis.
At the condylar head, chondrogenesis, in return, showed accelerated activity. As seen
from serial sections (Figs. 27-30), it had resulted in a considerable elongation of a
cartilaginous condylar process. The oldest cartilage was found at the base of the
coronoid process (Figs. 27, 28), from where it extended backward and upward into the

FIGs. 18 23. Various sites of secondary cartilage appearing in the mandible at the 60-mm. crown-
rump stage. Fig. 18: A cartilaginous condylar head developing at the site of the previous condylar
blastema. The temporal bone shows increased osteogenetic activity. (Orig. mag. X36.) Fig. 19:
Endochondral ossification within the anterior portion of Meckel's cartilage in the 60-mm. fetus.
S - median symphysis, and II dental organ of lateral temporary incisor. (Orig. mag. X55.)
Fig. 20: A coronal section at the level of the coronoid process, revealing three sites of secondary
cartilage formation: at the top and the base of the coronoid process and at the lower mandibular
border. (Orig. mag. X36.) Fig. 21: Higher magnification of the lower mandibular border of Fig. 20,
showing irregular pattern of secondary cartilage formation. (Orig. mag. X210.) Fig. 22: The presence
of chondroclasts in the bone marrow, indicating processes of cartilage resorption. (Orig. mag. X280.)
Fig. 23: On top of the coronoid process more regular chondrogenesis proceeding; rapid osteogenesis
was seen at the basal portion. (Orig. mag. X 140.)

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Vol. 41, No. 6 ON TOGENESIS OF TEMPOROMANDIBULAW JOINT. 1. CONDYLES 1335
condylar head. Periosteal bone formation invested the old carrot-shaped cartilage into
the newly formed body of the ramus (Fig. 29). Rapid trabecular osteogenesis con-
tributed to the formation of a mandibular angle (Fig. 29). Chondrogenesis proceeded
from the dorsolateral portion of the mesodermal covering of the condylar head (Fig.
30). At its medial aspect, an enlarged tendinous attachment of the condylar head with
the lateral pterygoid muscle had developed. At the base of the condyle, in preparation
of endochondral ossification, capillary elements were seen to penetrate from the periost
toward the degenerating chondrocytes (Fig. 31). Serial coronal sections of the pos-
terior half of the condyle revealed a joint cavity that separated the condylar elements
from the temporal joint structures. Differentiation of an articular disk and a glenoid
fossa of the temporal bone was starting at a still more posterior level (Fig. 32).

chnrgnssfuda the laeourirmsnhal coeing() The meil portion of thi

coern prdue tednu elmet (2) whle the baa portios wer enae in periosteal bon
Fi 26 Te lateruprio cete of chndrgeeis. (Oig ml|ag.
X280.)-

Histodifferetiationm cavnrumtiage aotne tFgthe 4condylar head)i compex phenomen onai

relatd t h opogenesis
ofna h l the enteire m hmandbl aoernd its.conecteda porgaons Aonghi
Musern nearthebifurcation
formauedtionpeeesitammrnous
olmns() ofthe
mentaland nerves. Expansive
incisal
vietebsteogenesiss boneformationfirst~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~...
Aereenaseof fuue muscelboe ....
frattahmnt are.sOitesg90)Fg 5:Temdilcne of rapidbonformation.truhu h al fori.mativeXstages
Threisa6heltronlogpeicordcner in the aperagneis (ofi these sieso2 gothspr.ontn

aito orsl dreclition of development. The fis oneformhead in th humane bheodyaieso

fromlate single inrmogemrnous ossfica etioncnerrnte paramediann
inonetedogumnt,amnd
dhevelopgsbypeiheraaccretion intole an prsimiivlhi plae.l Giardinoase contentionlof
5 usc6
diferetmandblrosfction centesina emrs snot substagntisAted by sevid rene* neither
isttahmetae
iniaionsofScttanid Syonse thratio thiprmryugossiiction center fraise situated.

nea the bifurcation ofScthe mental tandincisa nerves.rExpasiveboeioration

fetriirtuae
* A similar conjecture is found in G. Dubreuil, Embryologie humane, Paris: Vigot, 1941, and
widely reproduced in the French literature.

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1336 B. 4 U-tl E 1..1.1) es. .Vzember
t Deccmebc 1962

occurs at the lower border in the vicinity of the ends of the masseter and internal
pterygoid muscles. Another growth spurt in the region of the temporal muscle then
produces the coronoid process. It is conceivable that primitive fibrillar muscular con-
tractions may be conducive to these increased osteal growths. Since the lateral ptery-
goid muscle remains undifferentiated, however, until the condylar head has developed,
a different mechanism must be operative in the formation of the condylar process.
Throughout the earliest stages, it was noticed that the dorsal end of the dentary
remained particularly rich in undifferentiated mesodermal cells. We have designated

Fwits. 27- 32. Coronal serial sections through the posterior portion of the mandible of a 72-mm.
crown-rump fetus are shown. Fig. 27: The ramus at the level of the coronoid process, showing com-
plete cessation of chondrogenesis at the lower and the middle portions. (Orig. mag. x36.) Fig. 28:
The oldest layers of calcified condylar cartilage appearing in the posterior base of the coronoid process.
Note rapid trabecular bone formation at the lower border. (Orig. mag. X36.) Fig. 29: The anterior
portion of the condylar head, showing calcifying cartilage invested in a periosteal bone splint. Note
wide tendinous attachment of the lateral ptervgoid muscle. (Orig. mag. X36.) Fig. 30: The center
of the condylar head, showing columnar arrangements of chondrocy tes originating from a germinative
layer at the laterosuperior surface (.X. Note presence of articular space and trabecular osteogenesis
at the mandibular angle (A). (Orig. mag. X36.) Fig. 31: At the lover periphery of the condylar
head, some capillary loops penetrating from the periosteum toward the degenerating chondrocytes.
This is the very beginning of endochondral ossification. (Orig. mag. X280.) Fig. 32: Above the
dorsalmost portion of the condylar head, a disk is forming. The temporal hone shows a site of second-
ary cartilage formation (x). (Ojrig. mag. X36.)

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Vol. 41, No. 6 ONTOGENESIS OF TEMPOROMANDIBULAR JOINT. 1. CONDYLES 1337
this area "condylar blastema," thus avoiding the hitherto used terms related to other
elements of a future temporomandibular joint,18 19 as they prove to have a different
origin. Since bone tissue never grows interstitially and processes of modeling resorption
do not occur at the early stages, it must be realized that any new bone formed in this
condylar area will ultimately become a part of the more anterior portion of the dentary.
Mandibular topogenesis engenders continuous changes in relationship, particularly to
the attached muscles.15, 20
Toward the end of the tenth week of fetal life, cartilage appears almost simul-
taneously at various sites of the mandible. Three different patterns may be distin-
guished:
1. At first, the anterior portion of Meckel's cartilage undergoes endochondral ossifi-
cation* and becomes incorporated into the mandibular body. Whether or not this invo-
lution of the primordial cartilage has any bearing on the appearance of secondary
cartilage warrants further study.
2. Then secondary cartilage formation occurs at the sites of the insertion of the
medial pterygoid muscle, masseter and temporal muscle, where previously rapid trabec-
ular osteogenesis was observed. The cartilage here formed is quite irregular and not
highly differentiated. This passage of osteogenesis through a secondary cartilage stage
may be explained on the basis of an extraordinary growth spurt. The very limited dura-
tion of secondary cartilage production, in spite of a continued growth spurt, however,
suggests that some internal conditioning factors or induction may be operative.
3. Histogenesis of condylar cartilage, from the very start, shows a highly organized
pattern. It is the result of the final differentiation of the condylar blastema into the
elements of a specific synovial articulation; chondroblasts originate from a germinative
zone which is situated in the dorsolateral portion of the blastema; the capsular struc-
tures and tendon of the lateral pterygoid muscle arise from the medial zone of this
blastema, while its basal layers continue to form osteoblasts.
Endochondral ossification does not occur until a relatively late period.5 The primor-
dial condyles first increase in size both by chondrogenesis at the dorsolateral aspect and
by osteogenesis along the neck, whereby the older calcifying cartilage is getting in-
closed into the core.
The developmental rates of the human head show an apparent reversal in the gen-
eral embryogenetic law, which states that growth and differentiation progress gradual-
ly in a caudal direction from rostrum to tail.23 Ford,22 however, has shown allometri-
cally that after the tenth week of fetal life the anterior, prechordal, part of the cranial
base elongates considerably faster than the posterior, chordal, part. The condylar
blastema is located at a level anterior to the notochord. The onset of condylar cartilage
formation coincides, in fact, with the growth spurt in the prechordal skeleton. The rela-
tively late appearance of the mandibular growth cartilage thus is a phenomenon which
is in perfect correlation with the delayed development of the prechordal segment of the
mammalian skull.
Although the mandible originates as a typical membrane bone, its final makeup in-
volves other mechanisms which add to the unique position of the mandible among the
* The onset of this mechanism of skeletal development implies a beginning thyroid-parathyroid
activity which also presides over modeling bone resorption (Baume, Becks, and Evans, Am. J. Orthod.,
39:623, 1953).

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1338 BA UME J. D. Res. November-December 1962
bones of the skeleton: (1) endochondral ossification of the mediolateral portion of
Meckel's primordium incorporates a cartilage-bone component, (2) differentiation of
the condylar blastema into a secondary cartilage center adds a unique osteogenic phe-
nomenon, and (3) growth and transformation of the mandibular symphysis, whose
details are little known, constitutes still another peculiarity.
Ontogenetically, the mandible, therefore, is truly a "mixed bone," as originally con-
ceived by Kblliker.23
SUMMARY
Embryogenesis of the mandibular condyles was assessed in a material of 12 human
fetuses of ascending stages from 15 to 72-mm. crown-rump length. It was found that
condylar cartilage formation was a phenomenon closely related to the morphogenesis
of the entire mandible and skull. Secondary cartilage appeared simultaneously at all
sites of the insertion of the masticatory muscles. At the same time, the anterior portion
of Meckel's cartilage was incorporated into the mandibular body by a process of endo-
chondral ossification. The condyle originated from a mesodermal condensation which
was situated at the dorsal end of the dentary, in the vicinity of the proximal end of the
lateral pterygoid muscle. This so-called condylar blastema appeared at the 24-mm. CR
stage; at the early stages it contributed to the growth of the muscle and the dorsal
prolongation of the dentary. At the 55-mm. CR stage, rapid osteogenesis led to the
formation of a condylar head with no cartilage present. At the tenth week of fetal life
(60 mm. CR), as the culmination of this growth spurt, chondrogenesis started at the
dorsolateral portion of the blastema; concomitantly, the tendinous attachment of the
lateral pterygoid muscle was differentiated, while, at the condylar neck, periosteal bone
formation was proceeding. Endochondral ossification was not primarily involved in the
formation of the condyles; it was still absent at the 72-mm. CR stage. Neither was
there, up to this stage, any obvious developmental relations between condyle and tem-
poral joint elements.
The late maturation of the condylar blastema was shown to coincide chronologically
with the growth spurt of the prechordal segment of the cranial base.
Ontogenetically, the mandible is a "mixed bone"; it originated as a pure membrane
bone, then incorporated elements of the Meckel's cartilage primordium, and finally
developed at various sites secondary cartilage. Of these, only the condylar cartilage
persisted to form ultimately the highly organized growth center of the mandible.
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