ORIGINAL A R T I C L E S

Role of the lateral pterygoid muscle and

meniscotemporomandibular frenum in spontaneous
growth of the mandible and in growth stimulated by
the postural hyperpropulsor
J. J. Stutzmann* and A. G. Petrovic**
Strasbourg, France

Our experimental studies in young rats represent an attempt to elucidate the respective roles of the
lateral pterygoid muscle (LPM), the temporomandibular frenum (TMF), and the postural
hyperpropulsor (Hp) in the control of the growth rate, the growth direction, and the growth amount of
the condylar cartilage. Even after surgical resection of the LPM, the growth of the condylar cartilage
and the lengthening of the mandible continue but are significantly decreased. Also, the stimulating
effect of postural hyperpropulsion on the condylar cartilage and mandibular growth is much less
intense after resection of the LPM. The TMF, which has a blood-circulating and a biomechanical
component, appears to be a mediator of the LPM in the control of the condylar cartilage growth.
After the surgical resection of the TMF, the growth the condylar cartilage and the lengthening of the
mandible continue but are significantly diminished. (AMJ ORTHODDENTOFACORTHOP 1990;97:381-
92.)

T h e r e is now considerable agreement among means of intermittent electric shocks (frequency, 5

laboratory researchers that the rate and amount of con-
dylar cartilage growth may be modulated orthopedically
by repetitive placement of the mandible in a more for-
ward position. 143 Variations in the growth rate of the
condylar cartilage appear to be adaptations to the vari-
ations of the interjaw relationship. Like other adapta-
tions, they require the existence of appropriate feedback
loops. 4,14,15
Four lines of evidence have suggested that the lateral
pterygoid muscle (LPM) plays a role in the physiologic
control of the condylar cartilage growth rate:
1. After surgical resection of the LPM 14,17"2°in the
growing rat, untreated or treated with a functional ap-
pliance, a relative decrease in the growth of the condylar
cartilage can be observed.
2. Electromyographic record of the LPM in the
monkey treated with a functional appliance s.~.2~ shows
increased electrical activity.
3. Microelectronic stimulation of the LPM in the
young rat produces an increased rate of condylar car-
tilage growth. 22 The LPM was directly stimulated by
*In Charge of Research, National Institute of Health and Medical Research
(INSERM-U,213),
**Director of Research: Director, Laboratory for Craniofacial Cartilage and
Bone Growth Regulation (INSERM-U.213).
8/1/13730
times per second; duration, 10 ms; potential, 0.55 v).
4. After treatment with the postural liyperpropul-
sor, there is a significant increase in the proportion of
fast nonfatigable fibers in the young rat's LPM ~a and a
significant decrease in the number of serial sarcomeres
in the same muscle. 24 Thus, even without an increase
in contractile activity, the mandible is maintained in a
more forward position.
Nevertheless, Goret-Nicaise, Awn, and Dhem 25,26
have questioned the regulating role of the LPM in the
growth of the condylar cartilage.
Another structure belonging to the feedback lo0p
causal chain, which regulates the growth rate of the
condylar cartilage, is the meniscotemporomandibular
frenum* or retrodiscal pad. 27 Preliminary experiments
have suggested that the frenum not only has a biome-
chanical effect but also has a blood-circulating effect.
This has been demonstrated in the young rat by place-
ment of a ligature around the temporomandibular com-
ponent of the retrodiscal pad. 27
The following experiment was designed to inves-
tigate simultaneously the roles of the LPM and the TMF
*The retrodiscal pad includes the temporomandibular frenum and a meniscal
portion, In our experiments, only the temporomandlbular frenum was reseeted
or ligated. The meniscal portion was always protected from surgery and
clamping.

381
382 Sttttzrna/lll and Petrovic Am. J. Orthod. Dentofac. Orthop.
May 1990

Fig. 1. Radioautograph of sagittal, histologic sections of the condylar cartilage of a 48-day-old rat.
(Original magnification, x 80; final magnification, x 320.) In all 4-week experimental treatments, the
five zones of the condylar cartilage are visible: (1) fibrous layer; (2) mitotic compartment including
skeletoblasts and prechondroblasts; (3) zone of maturation including functional and hypertrophied
chondroNasts and cartilaginous matrix; (4) zone of erosion including degenerated chrondroblasts; (5)
zone of endochondral ossification, a, Control, sham-operated, b, After treatment with a postural hy-
perpropulsor. Note large increase in number of 3H-thymidine-labeled cells (arrows); moderate increase
in thickness of mitotic compartment; important widening of maturation zone. Note also intense hyper-
trophy Of the chondroblasts, c, After resection of LPM. Note large decrease in number of 3H-thymidine-
labeled cells (arrows); no evidential change in thickness of mitotic compartment; important decrease
in zone of maturation, with hypertrophy of chondroblasts less pronounced; decrease in number of
degenerated chondroblasts, d, After resection of TMF. Histologic modifications are similar to those
observed after resection of LPM. e, After clamping of TMF. Note decrease in number of 3H-thymidine-
labeled cells (arrows); histologic picture is, in general, similar to that of the control animal.

and the stimulating effect of a functional appliance, the METHODS

postural hyperpropulsor, on condylar cartilage growth One hundred eighty 20-day-old male rats (Sprague-
and mandibular lengthening. The experimental design Dawley strain) were used; half were treated for 12 hours
makes possible an adequate quantitative study and an daily (between 7 AM and 7 PM) with a 2 mm postural
appropriate statistical analysis. hyperpropulsor of the mandible (treated category), 14
Volume 97 LPM, TMF. and postural hyperpropulsor in growth of mandible 383
Number 5
while the remaining rats were not treated (reference
category). The design and the mode of action of the
postural hyperpropulsor have been described previ-
ously.~4 The male rats, originating from 42 litters with
9 to 11 newborn rats in each litter, were randomly
distributed. The rats were weighed every day, and our
study showed that weight was not significantly different
among intact and experimental rats. At the age of 48
days, both intact and experimental rats weighed
156 ± 11 grams. All experiments were conducted in
the period from April to June. The surgical operations
were performed and the rats killed between 1 PM and
3 PM. Both the hyperpropulsor-treated and the non-
treated animals were subdivided into 6 groups, each
consisting of 15 male rats.
Thefirst group of animals served as a control. These
animals had an intact lateral pterygoid muscle (LPM)
and an intact temporomandibular frenum (TMF) but
were sham-operated.
In the second group of animals, the LPM was sur-
gically resected bilaterally.17'18'19
In the third group of animals, the TMF was clamped
bilaterally (i.e., an appropriate ligature was placed bi-
laterally around the TMF to reduce the blood flow). A
directional optic fiber light was used under the operating
microscope (Wild M3, Type 355110). The ligature was
progressively squeezed until the condylar part turned
white. Visual inspection of blood vessels through the
operating microscope showed that the blanching effect
of such a ligation corresponds to the restriction in blood
flow. All the animals' blood flow was restricted using
this procedure.
In the fourth group of animals, the TMF was
clamped bilaterally and the LPM was resected bilat-
erally.
In thefifth group of animals, the TMF was resected
bilaterally.
Finally, in the sixth group of animals, both the LPM
and the TMF were resected bilaterally.
The experiment lasted for 4 weeks. As in previous
experiments, the rats were killed when they were 48
days old (i.e., just after the pubertal growth rate
p e a k ) . 14`tS'19`z7 One hour before being killed, each ani-
mal received an intraperitoneal injection of tritiated thy-
midine (3 fxCi/gm of animal weight; specific activity,
25 Ci/mmol). The condylar cartilage of one half of the
mandible was fixed, embedded, and cut serially. Serial
histologic sagittal sections of the condyle, 5 Ixm in
thickness, were radioautographed and stained with ei-
ther toluidine blue or hematoxylin and eosin (Fig. 1).
Radioautography was performed according to the
technique described by Kopriwa and Leblond? 8 The
growth rate of the condylar cartilage was evaluated by
counting the total number of tritiated thymidine-
labeled Cells on every fifth histologic section. The total
number of dividing cells at the condylar cartilage is
considered a better parameter for evaluating the growth
rate than an index (the number of labeled cells per 100
cells), since a dividing-cells index would be misleading
when the total number of cells in the mitotic compart-
ment is increased or decreased in treated animals.
After removal of the soft tissue from the opposite
side of the mandible, the growth direction of the con-
dyle ("Stutzmann's angle") was estimated radiograph-
ically by measurement of the angle between the main
orientation of the newly formed endochondral bone tra-
beculae, located in the central area of the condyle im-
mediately under the condylar cartilage, and the man-
dibular plane '~, 29 (Fig. 2). (An opening of the angle
indicates more posterosuperior growth and closing of
the angle indicates a primarily vertical component.)
This was done by placing the mandible under fine-grain
film and exposing it to a collimated x-ray beam of low
energy (30 to 35 kV) produced by a fine-focused tube
with molybdenum anticathode. The growth amount (the
operationally defined "length of the mandible") was
evaluated by measurement of the distance between the
posterior border of the condylar cartilage and the mental
foramen on the photographs and radiographs of the man-
dible (original magnification x 10).
The experimental data were analyzed by a three-way
analysis of variance (Tables I, II, and III and Fig. 3).
First-order interactions and a second-order interac-
tion between the different factors were detected. On the
basis of standard error for the whole population, the
smallest difference between two means was calculated
for each parameter. In this way, the different means were
compared in relation to one another (T method3°'3~). The
role of the LPM and of the TMF (retrodiscal pad) were
quantitatively investigated in nontreated rats and in rats
treated with a postural hyperpropulsor.
RESULTS
Animals not treated with the
postural hyperpropulsor (reference category)
We have previously described, in intact 48-day-old
male rats, (1) histologic features of the condylar car-
tilage, (2) cell filiation in situ, in cell culture and in
transplantation, ~'14'16and (3) variations of Stutzmann's
angle between the ages of 23 and 58 days. 2~ The his-
tologic, radiographic, and statistical findings relative to
different experimental groups are summarized in Figs.
1 to 3 and in Tables I to III.
Resection of the LPM alone. Resection of the LPM
384 Stutzmann and Petrovic
a Table IA. Total number of tritiated
b LPM
H
d thymidine-labeled cells in the condylar cartilage, and
Fig. 2. Radiograph of the mandible of a 48-day-old rat. a, Con-
trol, sham-operated, b, After treatment with postural hyperpro-
pulsor, the condylar bone trabeculae became oriented more
posteriorly. ¢, After resection of LPM, the condylar bone tra-
beculae became also oriented more posteriorly, d, After resec-
tion of TMF, the condylar bone trabecuiae became oriented
more vertically; this change is only statistically detectable.
alone produced a marked decrease in the number of
dividing cells (growth rate) in the condylar cartilage
and in the lengthening of the mandible. Stutzmann's
angle simultaneously opened. Histologically, when
compared to the control, the condylar cartilage (Fig.
1, c) presented the following modifications: The mitotic
compartment was slightly decreased in height; the chon-
droblastic zone showed a reduction in height where the
shortening appeared to be the result of decreased num-
Am, J. Orthod. Dentofae. Orthop.
May 1990
thymidine-labeled ceils in the condylar
cartilage (fifteen 48-day-old male rats in
each group)
Means
TMF
953 1449
LPMo 475 558
TM~
LPM 780 1071
LPM. 464 542
TMN
LPM 482 566
LPMo 426 453
Standard error for the total population-25.93
bers of dividing prechondroblasts; and the chondro-
blastic hypertrophy started later and was much less pro-
nounced than normal. In addition, some chondroblasts
did not undergo the usual degeneration observed in
normal condylar growth.
Resection of the TMF pad alone. Resection of the
TMF alone greatly decreased the number of tritiated
the lengthening of the mandible also diminished. The
decrease in the growth rate of the condylar cartilage
was roughly equal in value to the decrease observed
after resection of the LPM. The decrease in the length-
ening of the mandible was even more pronounced.
There was a statistically significant difference between
the group in which the LPM muscle was resected alone
and the group in which the TMF was resected alone
(however, the difference between the two means is
barely significant). Resection of the TMF caused a
slight closure of Stutzmann's angle. The histologic
modifications of the condylar cartilage after resection
of the TMF (Fig. 1, d) were similar or even more
pronounced than those observed after resection of the
LPM. Both the mitotic compartment and the chon-
droblastic zone were reduced in height.
Simultaneous resection of the LPM and the TMF.
Simultaneous resection of the LPM and the TMF did
not produce any additional changes in rate or direction
of growth of the condylar cartilage or in the length of
the mandible.
Clamping of the TMF. Clamping of the TMF pro-
duced a decrease in the number of dividing cells in the
condylar cartilage and in the lengthening of the man-
dible. For both parameters, the decrease was much less
pronounced than after resection of either the LPM alone
Volume 97 LPM, TMF, and postural hyperpropulsor in growth of mandible 385
Number 5

Table lB. Analysis of variance

Sum of Significance
Source of variations squares df Mean square F level

(1) Hp effect 1,403,793.42 1 1,403,793.42 139.15 ***

(2) TMF effect 4,339,583.41 2 2,169,791.71 215.08 ***
(3) LPM effect 7,106,704.20 1 7,106,704.20 704.46 ***
Interactions
1×2 410,901.01 2 205,450.5l 20.37 * * *
lx3 582,087.20 1 582,087.20 57.70 * * *
2x3 2,720,410.23 2 1,360,205.12 134.83 * * *
lx2x3 237,075.70 2 118,537.85 11.75 * * *
Error 1,694,822.26 168 10,088.23
TOTAL 18,495,337.44 179
Smallest significant difference between two means
At the 5% level, 72 * Significant at 0.05
At the 1% level, 96 ** Significant at 0.01
At the 1% level, 122 *** Significant at 0.001
NS, Not significant

df, Degrees of freedom; Hp,, no appliance; Hp, postural hyperpropulsnr (2 mm forward); TMF, temporomandlbular frenum (intact); TMF,,
clamped temporomandibular frenum; TMFo, resected temporomandibular frenum; LPM, lateral pterygoid muscle (intact); LPMo, resected lateral
pterygoid muscle.

or the TMF alone. The clamping produced an opening Table IIA. Angle (in degrees) between growth
of Stutzmann's angle. Histologically, the condylar car- direction of the condyle and the mandibular
tilage (Fig. l, e) was only slightly modified. Almost plane (Stutzmann's angle) (fifteen 48-day-old
no change was detected in the thickness of the different male rats in each group)
zones. The only change observed was that the chon-
Means
droblastic hypertrophy was less intense.
Simultaneous clamping of the TMF and resection ~o
of the LPM. When clamping of the TMF was associated
TMF
with resection of the LPM, there was a further decrease
LPM 122.7 134.9
in the number of labeled cells in the condylar cartilage LPMo 139.0 149. i
and in lengthening of the mandible, as well as an open- TMFc
ing of the Stutzmann's angle. The growth rate of the LPM 128.8 141.7
condylar cartilage was not significantly different from LPMo 134.7 145.9
TMFo
that observed after the resection of either the LPM alone
LPM 120,5 123.0
or the TMF alone or after simultaneous resection of the LPMo 120.1 120.9
LPM and TMF. As for the lengthening of the mandible,
the decrease was equal in value to that observed after Standard error for the total population = 0.5l
resection of the LPM and was in the same range as that
observed after resection of the TMF, but it was signif-
icantly less pronounced than after the simultaneous re- Lengthening of the mandible was decreased in all ex-
section of the LPM and TMF. perimental groups. The greatest decrease was observed
In summary, when compared with controls, the in the two groups in which the TMF had been resected
growth rate of the condylar cartilage was decreased in (groups 5 and 6, i.e., in which Stutzmann's angle
all experimental groups. The slightest decrease was ob- closed).
served in group 3, where only the TMF had been
clamped. Stutzmann's angle opened in alI the experi- AnimaJs treated with the postural hyperpropulsion
mental groups where the TMF had not been resected (treated category)
(groups 1, 2, 3, and 4). After resection of the TMF The histologic, radiographic and statistical analysis
(groups 5 and 6), Stutzmann's angle closed. Comments relative to the control animals is summarized in Figs.
on these results will be made in the Discussion section. 1 to 3 and in Tables I to III.
386 Stutzmann and Petrovic Am. J. Orthod. Dentofac. Orthop.
May 1990

• ~ LPM (intact) ~" [ ] ~ LPMo (resected)

Y¢ ~rTMF ( i n t a c t ) • [] TMFc (clamped) I) 0 TMF0 (reaected)

I O; degrees

eo
-= 31
L 5- i t ~.5
tt t 4
"
J:
o
I

t "
~..>, 4" N~IO" 2

cu m
o3 M2;8.2
41
3" 3 M3=1.19

E
E
J) 1.
=
z- ................ _~_L=_LL
s ~5 2

5 6
2 4 5 ,~_ 5
o
z~
6
=
m
=~o tt
s
" 0-

Fig. 3. Effects of postural hyperpropulsor on growth rate of condylar cartilage (number of tritiated
thymidine-labeled cells), on growth direction of condyle (Stutzmann's angle) and on length of mandible.
Each point represents the mean of the differences (with the limits of the 95% confidence interval)
between the mean value calculated for animals that were not wearing the postural hyperpropulsor
(reference category) and the mean value calculated for animals that did wear the postural hyperpro-
pulsor (treated category). M1 value corresponds to the mean of the postural hyperpropulsor effect on
the number of tritiated thymidine-labeled ceils in the six experimental groups. M2 value corresponds
to the mean of the postural hyperpropulsor effect on Stutzmann's angle in the 6 experimental groups.
M3 value corresponds to mean of the postural hyperproputsor effect on the length of the mandible in
the 6 experimental groups. The effect of the postural hyperpropulsor on the number of tritiated
thymidine-labeled cells is more marked in groups 1 and 3 (especially in group 1) than in groups 2, 4,
5, and 6. The effect of the postural hyperpropulsor on Stutzmann's angle is more marked in groups
1, 2, and 3, and 4 than in groups 5 and 6. The effect of the postural hyperpropulsor on the length of
the mandible is more marked in groups 1 and 3 than in groups 2 and 4; it is much less pronounced
in groups 5 and 6.

Histologicfeatures. Histologically, the sagittal sec-
tion of the condyles of intact rats, which had not un-
dergone surgical resection of the LPM, the TFM, or
both (Fig. 1, b), depicted the typical condylar cartilage
of a hyperpropulsed animal as described previously 1.3
(i.e., the mitotic compartment showed an increase in
vertical thickness and the chondroblastic zone was
greatly enlarged). This change appeared to be the con-
sequence of an increase in the number of prechondro-
blasts, which later differentiated into chondroblasts.
The chondroblast hypertrophy started much earlier than
usual resulting in the large increase in the number of
hypertrophied chondroblasts. In addition, the hypertro-
phy of the Chondroblasts was so pronounced that the
surrounding cartilaginous matrix had almost vanished.
The analysis of variance showed that in intact rats
(i.e., animals that had not undergone the surgical re-
section of the LPM, the TMF, or both) the wearing of
the postural hyperpropulsor produced a highly signifi-
cant increase (p < 0.001) in the number of HL
thymidine-labeled cells and in the length of the man-
dible. There was also an opening Of Stutzmann's angle.
Resection of the LPM in animals treated with the
postural hyperpropulsor. In the animals that underwent
resection of the LPM, the wearing of the postural hy-
perpropulsor produced only a very small but still sig-
nificant increase in the number of HLthymidine -
labeled cells in the condylar cartilage as well as in the
length of the mandible. At the same time, Stutzmann's
angle increased.
Resection of the TMF in animals treated with the
postural hyperpropulsor. In the animals that were sub-
jected to resection of the TMF, the wearing of the pos-
tural hyperpropulsor significantly enhanced the growth
rate of the condylar cartilage and caused an opening of
Stutzmann's angle. These effects were not sufficient to
Volume97 LPM, TMF, and postural hyperpropulsor in growth of mandible 387
Nnmber 5

Table liB. Analysis of variance

Significance
Source of variations Sum of squares df Mean square F level

(1) Hp effect 3,100.05 1 3,100.05 784.35 ** *

(2) TMF effect 10,302.68 2 5,151.34 1,603.35 ** *
(3) LPM effect 1,811.34 1 1,811.34 458.29 ** *
Interactions
1 X 2 996,10 2 498,05 126.01 ** *
1 x 3 36.45 1 36.45 9.22 **
2 x 3 2,078.61 2 1,039.31 262.96 ** *
1 x 2 x 3 0.43 2 0.22 0.05 NS
Error 664.00 168 3.95
TOTAL 18,989.66 179
Smallest significant difference between two means
At the 5% level, 1,4 * Significant at 0,05
At the 1% level, 1.9 ** Significant at 0.01
At the 1% level, 2.4 *** Significant at 0,00l
NS, Not significant

df, Degrees of freedom; Hp,, no appliance; lip, postural hyperpropulsor (2 mm forward); TMF, temporomandibular frenum (intact); TMF¢,
clamped temporomandibular frenum; TMFo, resected temporomandibular frenum; LPM, lateral pterygoid muscle (intact); LPM,, resected laterat
pterygoid muscle,

cause a significant supplementary lengthening of the Table IliA. Length of the mandible (in mm):
mandible. distance between posterior border of the
Simultaneous resection of the TMF and LPM in condylar cartilage and the mental foramen
animals treated with the postural hyperpropulsor. In (fifteen 48-day-old rats in each group)
the animals in which the TMF and the LPM had been
Means
simultaneously resected, the wearing of the postural
hyperpropulsor did not significantly affect any of the
three measured parameters--growth rate of the con-
TMF
dylar cartilage, Stutzmann's angle, or length of the
LPM 14.08 16.07
mandible. LPMo 12.53 13.34
Clamping of the TMF in animals treated with the TMF~
postural hyperpropulsor. In the animals that were sub- LPM 13.33 15.39
jected to clamping of the TMF, the wearing of the LPMo 12.46 13,80
TMFo
postural hyperpropulsor stimulated, in a highly signif-
LPM 11.88 12.34
icant manner, the growth rate of the condylar cartilage LPM° 11.33 I 1,83
and the lengthening of the mandible and produced a
strong opening of Stutzmann's angle. Standard error for the total population = 0.23
Simultaneous clamping of the TMF and resection
of the LPM in animals treated with the postural hy-
perpropulsor. In the animals that were subjected si-
multaneously to clamping of the TMF and to resection the six experimentat groups by the difference between
of the LPM, the wearing of the postural hyperpropulsor the mean value calculated for animals that were not
still produced a significant acceleration in the growth wearing the postural hyperpropulsor (reference cate-
rate of the condylar cartilage and enhanced the length- gory) and the mean value calculated for the animals
ening of the mandible. The opening of the angle was that did wear the postural hyperpropulsor (treated cat-
even more pronounced than after clamping of the TMF egory) (Fig. 3).
alone. The statistical analysis of the data regarding the
Graphic representation of the effects of the postural length of the mandible has shown that the postural hy-
hyperpropulsor in different experimental groups. The perpropulsor was most effective in group t (in which
stimulating effect of the postural hyperpropulsor on the the LPM and the TMF were not resected) and in group
growth of the mandible can he represented for each of 3. There was no significant difference between these
388 Stutzmann and Petrovic Am. J. Orthod. Dentofac. Orthop.
May I990

T a b l e IIIB. Analysis of variance

. . . . . . Mean Significance
Source of variations Sum of squares df square F level

(1) Hp effect 63.1783 1 63.1783 80.72 ***

(2) TMF effect 165.6542 2 82,8271 105.83 ***
(3) LPM effect 76.8059 1 76.8059 98.14 ***
Interactions
1×2 11.8646 2 5.9323 7.58 ***
lX3 4.5188 1 4.5188 5.77 *
2x3 20.2660 2 10.1330 12.95 ***
lx2×3 3.1189 2 1.5595 1.99 NS
Error 131.4847 168 0.7827
TOTAL 476.8914 179
Smallest significant difference between two means
At the 5% level, 0.64 * Significant at 0,05
At the I% level, 0.84 ** Significant at 0.01
At the 0.1% level, 1.07 *** Significant at 0.001
NS, Not significant

df, Degrees of freedom; Hpo, no appliance; lip, postural hyperpropulsor (2 mm forward); TMF, temporomandibular frenum (intact); TMF~,,
clamped temporomandibularfrenum; TMFo, resected temporomandibularfrenum; LPM, lateral pterygoid muscle (intact); LPMo, resected lateral
pterygoid muscle.

two groups (1.99 mm in group 1, 2.06 mm in group
3). In both groups, the supplementary lengthening of
the mandible resulted from an increase in the number
of thymidine-labeled cells in the condylar cartilage and
from a strong opening of Stutzmann's angle. The stim-
ulating effect of the postural hyperpropulsor was less
pronounced in groups 2 and 4 than in groups 1 and 3,
but there was no statistically significant difference be-
tween the effects produced in groups 2 and 4. The
resection of the LPM produced a very significant
decrease but never a total arrest of the cell divi-
sions in the condylar cartilage. Finally, in groups 5
and 6 the postural hyperpropulsor no longer had a de-
tectable stimulating effect on the lengthening of the
mandible.
In summary, aside from the experiment in which
the TMF and the LPM were simultaneously resected
(group 6), the postural hyperpropulsor always amplified
the growth rate of the condylar cartilage and produced
an opening of Stutzmann's angle. However, the mag-
nitude of these supplementations varied greatly, de-
pending on whether the rat was intact or had been op-
erated on (either LPMo, TMFc, or TMFo). A supple-
mentary lengthening of the mandible was observed only
in the experiments in which the TMF had not been
resected (groups 1, 2, 3, and 4). The greatest effect of
the postural hyperpropulsor was observed in intact rats
(group 1). The effect was less pronounced in TMFc
(group 3) and lowest in TMFc + LPMo (group 4) and
in LPMo (group 2).
Interactions between the effect of the postural
hyperpropulsor (Hp), the TMF, and the lateral LPM
The effects of the three factors (Hp, TMF, LPM)
are not additive. The analysis of variance revealed the
existence of an attenuation type of interaction between
(1) the effects of resection or clamping of the T M F and
of the LPM; (2) the effects of resection or clamping of
the TMF and of the postural hyperpropulsor; and (3)
the effects of resection of the LPM and of the postural
hyperpropulsor.
When one or two factors were surgically sup-
pressed, the attenuation type of interaction detected cor-
responds to an amplification type of interaction between
these factors when they are operational.
DISCUSSION
To justify this study, it is necessary to make the
following three remarks. This is followed by the major
conclusions of the study.
First, isolated histologic pictures are only snapshots
of a given section of the condylar cartilage (i.e., they
may illustrate, but not demonstrate, the actual results of
the investigation). Only a precise, detailed quantitative
study of the three parameters investigated in this study
(the total number of dividing cells in the condylar car-
tilage, Stutzmann's angle, and the operationally defined
length of the mandible) may lead to valid statistical and
methodologic conclusions concerning the effects of re-
section of the LPM, of resection or clamping of the
TMF, and of the action of the postural hyperpropulsor.
Vohune 97
Number 5
Second, it must be emphasized that the operation-
ally defined length of the mandible partly depends on
both the growth rate and the growth direction of the
condylar cartilage. Thus, when the growth rate of the
condylar cartilage remains constant, a greater opening
of Stutzmann's angle will yield an increase in the length
of the mandible; conversely, when the growth rate of
the condylar cartilage is diminished, the lengthening of
the mandible, as measured between the posterior border
of the condylar cartilage and the mental foramen, is not
necessarily decreased. In this case, the opening of
Stutzmann's angle may compensate for the diminished
growth of the condylar cartilage. However, as our in-
vestigations clearly show, the compensatory opening of
Stutzmann's angle occurs only when the TMF was not
previously resected. It is necessary to mention that the
growth in length of the rat mandible depends not only
on the condylar cartilage but also on the growth of the
angular cartilage and on the growth at the posterior
border of the ramus. The growth of the mandible as a
whole also depends on the subperiosteal ossification*
and on the alveolar bone formation.
Third, it should be pointed out that, in the rat, the
forward movement of the mandible is produced not only
by the LPM but also by the anterior fibers of the mas-
seter muscle. Thus even after resection of the LPM,
the mandible can be protruded forward and thereby the
temporomandibular component of the retrodiscal pad
may be functionally stimulated.
Resection of the LPM produced a significant de-
crease but never an arrest of the cell divisions in the
condylar cartilage. In other words, after resection of
the LPM, the growth of the condylar cartilage and the
lengthening of the mandible persisted but to a lesser
extent. However, the LPM does appear necessary for
the optimal lengthening of the mandible and intermax-
illary adjustment, tga7
As mentioned previously, our first observation con-
cerning the role of the LPM in condylar growth cartilage
and in the action of functional appliancesl6' t7was further
corroborated by McNamara, s,6.21 by Kantomaa and
Rrnning, 2~ and by Whetton and Johnston. 2° However,
some authors disagree with these findings. According
to Awn, Goret-Nicaise, and Dhem, 26 the unilateral sec-
tion of the LPM in the growing rat does not alter con-
dylar growth. However, their experimental design (nu-
merous subgroups, a small number of individuals in
each subgroup, etc.) makes it difficult to detect statis-
tically significant differences between the control rats
*Our previous research in rat and human subjects have shown that there is a
positive correlation in tile variations of the subperlosteal ossification rate, al-
veolar bone turnover rate, condylar cartilage growth rate, and the clincial
effectiveness of an orthodontic, functional, or orthopedic appliance. 32'a3
LPM, TMF, and postural hyperpropulsor & growth of mandible 389
and the rats that have undergone sectioning of the LPM.
And the absence of evidence is not evidence of absence!
Resection of the LPMproduced a highly significant
opening of Stutzmann's angle (i.e., a "posterior growth
rotation" of the mandible). Once again, this effect was
observed only when the biomechanical effect of the
TMF persisted. Indeed, this posterior growth rotation
of the mandible was still observed after clamping of
the frenum but not after resection of the frenum.
Why does resection of the LPM result in the opening
of Stutzmann's angle? After LPM resection it was ob-
served that the anterior fibers of the masseter tended to
maintain the mandible in the usual occlusal relationship
in spite of the decreases in the condylar growth and in
the lengthening of the mandible. The physiologic ten-
dency to maintain the normal occlusal intermaxillary
relationship--that which Petrovic15 calls in his cyber-
netic language the "attractor"* effect--implies an in-
creased contractile activity of the LPM and of the an-
terior fibers of the masseter. 4,t4 As a result, there is a
supplementary solicitation of the TMF, partly through
the intermediary action of the disc. It follows that Stutz-
mann's angle will open. Since the mandible is shorter
than normal, the biomechanical traction-effect of the
TMF on the condyle appears to be enhanced (i.e., the
growth of the condyle and, consequently, of the ramus
becomes more posterior). Thus Stutzmann's angle
opens.
Resection of the TMF produced a reduction in the
condylar cartilage growth rate that equaled in value the
reduction observed after resection of the LPM. This
finding means that the TMF, also, is not absolutely
necessary for the growth of the mandible, but, once
more, when the TMF is missing, the growth of the
condylar cartilage becomes quantitatively deficient.
We may assume that this diminished growth of the
condylar cartilage results from the surgically induced
reduction in blood and lymph flow. The closing of
Stutzmann's angle appears to be due to the absence of
the usual biomechanical traction effect of the TMF.
When the LPM and the TMF were resected simul-
taneously, the decrease in the condylar cartilage growth
rate was of the same magnitude as that observed after
the resection of LPM alone. 27 When compared to the
experimental group in which only the TMF was re-
sected, the simultaneous resection of LPM and TMF
produced no supplementary change in either growth rate
*In tile child tile "attractor" situation as represented by a full interdigitafion
(full Class I, but to a certain extent also full Class lI or full Class III) corresponds
to a structurally stable steady-state in a dynamic system as represented by the
occlusal relationship. A cusp-to-cusp type of occlusal relationship corresponds
to the "repeller" situation (i,e., to an unstable equilibrium state).
390 Stutzmann and Petrovic
or length of the mandible. How can we account for this
finding? A definitive answer to this question is still
premature. Tentatively, we may deduce from our ex-
perimental findings (Tables I, iI, and III), that the TMF
is, in part, the physiologic mediator of the LPM; that
is, we are dealing not with two separate factors, but
with a causal chain leading to the condylar cartilage
growth. 27 Experiments now in progress, where either
the superior or the inferior head of the LPM has been
resected separately, logically suggest that the TMF, us-
ing the disk as an intermediate, is the mediator of the
upper head of the LPM.
Ligation of the TMF caused a significant lowering
in the growth rate of the condylar cartilage. However,
tlais lowering was much less pronounced than after re-
section of the TMF. This is easy to understand, as a
ligature placed around the TMF produced a decrease in
only the blood circulation and lymph flow. Thus, when
the TMF has been resected, the more pronounced low-
ering in the condylar cartilage growth rate may be ac-
counted for both by the complete suppression of the
frenum blood circulation and the lymph flow decrease
and by the suppression of the biomechanical action of
the frenum.
The modus operandi of the postural hyperpropulsor
has been described previously.14'27'29.34'35Cybernetically
speaking,4.15 when the postural hyperpropulsor is placed
on the upper incisors, everything occurs as if the upper
dental arch (the "constantly changing reference input")
were in a more anterior position. The "confrontation"
between the dental arches at the "comparator" of the
servosystem then produces a "deviation signal." The
deviation signal will be reduced by an appropriate for-
ward positioning of the lower dental arch through the
muscle propulsion of the mandible. This implies a sup-
plementary contractile activity of the LPM, inducing a
supplementary growth of the condylar cartilage and,
consequently, a supplementary lengthening of the man-
dible. Withtime, this supplementary lengthening will
result in a reduction of the anatomic relative "retro-
position'! of the mandible and thus in a decrease of the
!'deviation signal." The supplementary contractile ac-
tivity of the LPM and the supplementary condylar car-
tilag e growth rate will also tend to diminish as a result.
However, the mandible of the treated rat will be longer
than that of the untreated animal.
Three points should be emphasized. First, this in-
terpretation agrees with the findings reported by
McNamara 6.36and McNamara and Carlson 12in the mon-
.key.. Second, if the LPM is necessary for optimal stim-
ulation of the condylar cartilage growth rate, the new
experiments corroborate previous findings that resection
Am. J. Orthod. Dentofac. Orthop.
May 1990
of the LPM does not suppress totally the stimulating
effect of functional appliances on the lengthening of
the mandible. 19.35Finally, according to Tewson, Heath,
and Meikle, 37 an anterior mandibular displacement in
the growing rat does not stimulate cell proliferation or
matrix formation at the mandibular condyle. The ex-
cessive size of their appliance and the short duration of
their experiment (1 to 14 days) could account for their
negative results. In this context, it should also be
pointed out that, in our own experiments, when the
thickness of the postural hyperpropulsor reached 3 ram,
the number of dividing cells and the length of the man-
dible became smaller than in control animals, a4
According to the findings reported here, the wearing
of the postural hyperpropulsor brings about a supple-
mentary lengthening of the mandible only in the ex-
perimental groups in which the TMF has not been re-
sected. In the new experiments, a small stimulating
effect of the appliance on the lengthening of the man-
dible was again detected, even in the rat whose LPM
had been previously resected. In other words, the TMF
appears to be an essential link in the action of the
postural hyperpropulsor, whereas the LPM is only
partly, even when predominantly, mediating the action
of the postural hyperpropulsor on the lengthening of
the mandible. It should be recalled that after the resec-
tion of the LPM in the rat the anterior fibers of the
masseter muscles induce a more forward positioning of
the mandible and thus stimulate an increased activity
of the TMF.
As pointed out, the supplementary lengthening of
the mandible induced by the postural hyperpropulsor
results not only from an increased growth rate of the
condylar cartilage but also from the opening of Stutz-
mann's angle. Thus, the stimulating effect of the
postural hyperpropulsor on the lengthening of the man-
dible is as marked in group 3 (TFMc) as in the control
group. This is due to the intense opening of Stutzmann's
angle, which compensates for the lesser postural
hyperpropulsor-induced increase in the condylar carti-
lage growth rate.
A large opening of Stutzmann's angle was observed
in group 2 (LPMo) and group 4 (TMF~ + LPMo). Here,
however, this intense opening was not sufficient to com-
pensate for the slight postural hyperpropulsor-induced
increase in the condylar cartilage growth rate in groups
2 and 4. Hence the postural hyperpropulsor-induced
supplementary lengthening of the mandible remains less
marked than in the control group.
It should be recalled that the opening of Stutzmann's
angle induced by the appliance appeared to be only a
transient remedial occurrence. '-9'34 In the long run, the
Volume 97
Number 5
l e n g t h e n i n g o f the m a n d i b l e elicited b y the postural
h y p e r p r o p u l s o r o c c u r s substantially, if not exclusively,
t h r o u g h s u p p l e m e n t a r y growth.
In the child, also, a significant increase in overall
m a n d i b u l a r l e n g t h c a n b e a c h i e v e d with a functional
a p p l i a n c e ( L S U - A c t i v a t o r ) , b u t o n l y in p e r s o n s with a
h i g h tissue-level g r o w t h potential and r e s p o n s i v e n e s s
as d e t e c t e d biologially b y the m a n d i b u l a r subperiosteal
ossification rate a n d a l v e o l a r b o n e t u r n o v e r rate. 38
The expert technical assistance of Ms. D. George and
Ms. M. Morales is gratefully acknowledged.
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Reprint requests to:
Dr. Jeanne J. Stutzmann
INSERM U.213
Facult6 de Mrdecine
4 rue Kirsehlrger
67085 STRASBOURG Cede×
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