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Transverse periosteal sectioning and femur growth in the rat.

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THE ANATOMICAL RECORD 207:339-348 (1983)
Transverse Periosteal Sectioning and Femur Growth
in the Rat
JAMES B. McLAIN AND PETER S. VIG
Bowman Gray School of Medicine, Wake Forest Uniuersity, Winston-Salem,
NC 27103 (J.B.M.) and Department of Orthodontics, the Uniuersity of
Michigan, A n n Arbor, MI 48109 R S . V)
ABSTRACT
Circumferential cuts through the periosteal covering of long
bones have been demonstrated to transiently increase epiphyseal growth. This
effect appears to be independent of vascular changes accompanying surgery
and has been hypothesized to relate to releasing tension in the periosteal
envelope. This study was designed to address problems of previous investigations by controlling for the effects of the surgical procedure and by using
regression analyses to analyze intra- and interanimal variations in the length
and proportionality of the femur in experimental, sham, and control SpragueDawley male rat littermates. Experimental animals received circumferential
periosteal sectioning of the right femur and no operation to the left limb. A
sham operation without periosteal sectioning was performed on the right
femur in the sham group. Right to left differences were analyzed using two
multiple regression models; one involved three absolute length measurements
as the dependent variables, while the other used the three ratios of these
length measurements as the dependent variables. The ratio measures were
utilized to reflect changes in bone proportionality.
Circumferential periosteal section was followed by a n alteration in the shape
of rat femurs a t 2 weeks postsurgery with a slight retardation of the length
dimension from medial epicondyle to head of the femur and a n overgrowth of
the length dimension from the lateral epicondyle to the greater trochanter.
The sham procedure produced a proportional decrease in all length measurements. The experimental procedure was also associated with surface bone
apposition at the site of section. At 3 weeks postsurgery, normalization of bony
contours between sham, experimental, and control groups had occurred; however, there were still some statistically significant decreases of length dimensions in the sham and experimental groups. In the experimental group the
length measurement involving the weight-bearing head of the femur remained
reduced a t 3 weeks postsurgery.
It is hypothesized that the functional demands of the long bone play a n
important role in the effect of periosteal regulation on growth. In situations
where a normal tensive force is exerted on the bone, the periosteal envelope
will act to restrain epiphyseal growth. When the bone is under a normal
compressive force the release of periosteal tension is not a quantitatively
significant stimulus to epiphyseal growth and the effects of surgical intervention and muscle trauma will play a more important role in the growth response
of the epiphyses.
Form and function interactions that play
a n important role in normal bony development must act through a variety of mediating factors from muscle and soft tissue
influences at the gross level to biochemical
processes at the cellular level. The role of the
0 1983 ALAN R. LISS. INC.
periosteal covering of bones in this interaction remains a n area of interest for investigation. It has been suggested that the
growing epiphyses place tension on the fiReceived January 21,1983; accepted May 13, 1983.
340
J.B. McLAIN AND P.S. VIG
brous layer of the periosteum and that essentially it is stretched over the underlying shaft
(Lacroix, 1951; Moss, 1978). The hypothesis
of a “neutral zone” suggests that a region
exists near the midshaft of a long bone where
the tension on the fibrous layer of the periosteum from the two growing epiphyses is
equalized and is therefore essentially zero. A
slower growing periosteum could then exert
a compressive force on the underlying epiphyses and thus control or regulate the
growth of the long bone.
Hueter (1862) and Volkmann (1862) noted
the relatively diminished growth of bones
that are under compressive forces. Others
have demonstrated the ability to retard longitudinal growth of long bones by the application of compressive loadings on the epiphyseal plates (Mueller, 1922; Strobino, et al.,
1952; Bylander et al., 1981, 1983). One
method of accomplishing this, by the stapling
of one epiphyseal plate, results in an overgrowth of the contralateral epiphysis (HallCraggs and Lawrence, 1969). It may be hypothesized that the stapling procedure moves
the neutral zone toward the stapled epiphysis, thus reducing the force on the opposite epiphysis and permitting more growth.
Roskjaer (1977) noted that the highest turnover of periosteal cells occurs adjacent to the
fastest growing epiphysis.
Attempts to stimulate long bone growth in
cases of limb inequality have been a stimulus to the study of periosteal stripping operations. Unlike a simple circumferential cut
through the fibrous layer of the periosteum,
most of these studies have involved partial
or complete stripping of the periosteum from
the shaft of the long bone. These methods
and others that have attempted to increase
endosteal blood flow have generally produced
unpredictable results (Wu and Miltner, 1937;
Lacroix, 1951; Brodin, 1955; Jansen and Langenskiold, 1956; Bei-trand and Trillat, 1948;
Jenkins et al., 1975; Larson et al., 1961; Sola
et al., 1963; Shui and Wong, 1964; Yabsley
and Harris, 1965; Pease, 1952; Frejka and
Fait, 1958; Chan and Hodgson, 19701.
The unpredictability of the stripping operations may relate to the possibility that two
control systems are being disturbed. In a
study of rabbits Whiteside and Lesker (1978)
showed that only in animals with traumatized muscle did periosteal dissection appear
to severely compromise collateral circulation
and that in animals with untraumatized
muscle neither extraperiosteal nor subperiosteal dissection reduced blood flow. There-
fore in those experiments involving a greater
degree of muscle trauma, it may be hypothesized that endosteal blood flow increased, and
greater stimulation of the growing epiphyses
would be expected.
If the periosteum is under tension and does
restrict epiphyseal growth, surgical release
of this tension should stimulate growth, at
least until the periosteum heals. Crilly (1972)
and Auer et al. (1982) observed measurable
clinical retraction of the margins of circumferentially cut periosteum in long bones. Hellquist (19721, in experiments which involved
stripping of the periosteum from rabbit maxillae, found consistent regeneration of the
resected periosteum that was more adherent
to the underlying bone, as well as transiently
more cellular and thicker. These histological
changes could easily affect the elasticity of
the fibrous layer of the periosteum and its
ability to stretch over the underlying bone.
In fact, Hellquist observed a deviation of the
snout toward the operated side when periosteum was stripped from the left nasofrontoand premaxillo-maxillary bones.
Crilly (1972) attempted to differentiate the
effects of periosteal tension, vascular damage, and muscle trauma on long bone growth
in a set of experiments utilizing the radius of
the chicken. At 17 days he found a n initial
overgrowth of 27.3% for transverse fracture,
2.8% for muscle trauma, 24.8% for transverse periosteal section, and only 1.9% for
longitudinal periosteal section. Results at 120
days demonstrated differences that were
much reduced-8.5%, 1.0%, 5.7%, and O.O%,
respectively. He concluded that the major
factor involved in longitudinal overgrowth
was the release of the periosteal tension and
that little was gained with the addition of
transverse fracture of the shaft. He recognized that there was some influence from
changes in endosteal blood flow, but concluded that this contribution was negligible.
The negligible results with longitudinal periosteal section coincide with Whiteside and
Lesker’s (1978) observations that the collateral circulation is not severely compromised
unless muscle is traumatized.
One severe limitation of Crilly’s work was
that experimental and sham radii were used
for comparison. This has the potential for
exaggerating the amount of overgrowth,
since both experimental and sham radii received surgery to place radiopaque implants.
This design precludes any assessment of the
effect of the surgical approach and inadvertent muscle damage on the growth of the ra-
TRANSVERSE PERIOSTEAL SECTIONING
dii. It is therefore possible that an observed
increase in growth may have been a feature
of the experimental design and not a valid
test of his hypothesis.
The purpose of the present study was to
differentiate the effects of surgery alone from
those of surgery together with periosteal section on growth of the rat femur. The rat femur was selected since it is a weight-bearing
bone. In contrast to Crilly's studies on chick
radii, we also sought to detect the possible
influence of function on the growth behavior
of a long bone.
MATERIALS AND METHODS
Six sets of six male Sprague-Dawley
outbred albino rat littermates were used for
this study. All 36 animals were 21 days old
at the time of surgery. The planned time of
sacrifice for three litters (18 animals) was 14
days after surgery (i.e., 35-day-old animals),
and for the remaining three litters (18 animals) the date of sacrifice was 21 days after
surgery (i.e., 42-day-old animals). The litters
were randomly assigned to 2- and 3-week
groups using a table of random numbers. In
each of the litters three animals were designated as experimental and three as sham
animals. The experimental or sham procedure was limited to the right femur, and for
each animal the left femur served as the
unoperated comparison for that animal.
Changes in the normal function of these contralateral limbs could not be accounted for in
this design; however, the animals continued
normal activity and diets throughout the
study period. The experimental procedure
consisted of a dorsal approach to the midshaft region of the femur. A sharp curette
was used to cut and gently reflect the perios-
Fig. 1. Length measurements used for the study. A)
the longest distance from the most proximal point on the
head of the femur to the most distal point of the medial
condyle; B) the shortest distance from the most distal
point on the neck of the femur to the most proximal
341
teum completely around the shaft. The cut
margins of the periosteum retracted leaving
an approximately 2-mm wide area of bony
shaft exposed. The completeness of section
was confirmed by the tactile feeling of the
curette against the bone. The area of section
approximated the region where the distal extent of the Adductor brevis and proximal end
of the Vastus intermedium muscles insert
into the shaft of the femur. This approach
appeared to be the most atraumatic available to the midshaft region. The skin was
sutured and the animals were returned to
their cages for routine care.
The sham procedure consisted of an identical surgical approach with reflection of all
muscles, but did not include periosteal section. No surgical procedures were carried out
on the left femurs. An additional series of
unoperated femurs from 42-day-old animals,
utilized in a similar surgical study of the
mandible, were utilized to gauge normal intraanimal variation between right and left
femurs.
Animals were sacrificed according to their
schedule at either 2 or 3 weeks with an overdose of intraperitoneal pentobarbital. Both
femurs and the mandibles from each animal
were recovered and defleshed by Sarcophagus beetle larvae. The bones with cartilage
intact were placed in warm tap water in
sealed containers. After approximately two
rinsings with water the articular cartilage
cleanly separated from the underlying bone.
The bones were then dried for examination.
Direct measurements were made on each
femur by the same investigator using a Helios caliper. Figure 1 shows the location of
the three length measurements (A, B, and C)
on a drawing of a 42-day-old rat femur. All
point on the intercondylar notch; and C) the longest
distance from the most proximal point of the greater
trochanter to the most distal point of the lateral condyle.
Shaded area indicates site of section.
342
J.B. McLAIN AND P.S. VIG
measurements were taken on three separate
occasions by the same investigator, and measurement error did not exceed 0.5% for any
dimension. Data were analyzed using the
Statistical Analysis System’s multivariate
regression analysis package (GLM). Each of
the three independent recordings of the measurements were kept separate for analysis
purposes and provided the pure error sum of
squares for the regression models generated.
Two separate statistical models were developed for this study. The first model included
the differences between right and left femurs
for the three length measurements A, B, and
C a s the dependent variables; the individual
animals served as the independent variables.
In the second model, which was used to test
for similar proportionality of the bones in the
experimental, sham, and control groups, the
dependent variables were the differences between right and left femurs for the ratios
A:B, A:C, and B:C, and the independent variables were the individual animals. By utilizing difference measurements between right
and left femurs for each animal, in effect
each animal served as its own control.
Separate comparisons of each group with
the other two, i.e., experimental to sham,
experimental to control, and sham to control,
were made using the “contrast statement”
capabilities of the GLM statistical program.
The control group allowed assessment of the
normally occurring intraanimal variation
between right and left femurs, while the use
of a sham group allowed separation of effects
due to sectioning and surgical procedure and
the effect of surgical intervention alone.
These factors combine to make this design
very sensitive to changes due to treatment.
Data for the femurs were analyzed separately for the animals sacrificed at 2 and 3
weeks postsurgery to eliminate statistical
problems associated with growth effects. For
the animals sacrificed at 2 weeks no intraanimal variation group existed, and sham and
experimental groups were compared. For animals sacrificed a t 3 weeks postsurgery, the
intraanimal limb variation was available
from animals taken from a parallel study
involving the mandible. The intraanimal
limb variation was quite small at 4 days of
age and one would not expect the magnitude
of this variation to have been larger a t 3 days
with smaller femurs.
RESULTS
Gross observations of the femurs taken
from animals sacrificed 14 days and 21 days
postsurgery are shown in Figures 2 and 3,
respectively.
At 14 days postsurgery both the experimental and sham animals demonstrate a decrease in length A on the operated side (Fig.
4); however, the experimental animals demonstrate less of a n effect than do the sham
animals with the difference significant at P
< 0.0001 (Table 1).For Measurements B and
C, the experimental group demonstrates a n
increased length on the operated side, while
the sham group demonstrates a decreased
length on the operated side. The length differences between the right and left sides between the two groups were significant at P
< 0.0001.
The ratio measurements A:B, A:C, and B:C
represent a n attempt to measure distortion
or “bending” of the femur attributable to the
interventions (Fig. 5). A reduction in ratio
A:B in the experimental group 14 days postsurgery reflects the relative overgrowth of B
and undergrowth of A. In contrast, the sham
animals showed reduction in B on the operated side, which was relatively greater than
the reduction of length A.
Ratio A:C demonstrates a decrease in A
relative to C in the experimental group. For
the sham group, lengths A and C have been
reduced in similar proportions so that the
ratio value A:C is the same for operated and
unoperated sides.
Ratio B:C remains the same for operated
and unoperated sides in the experimental animals, indicating that both lengths B and C
were increased in similar proportions. The
sham animals were characterized by a relatively greater reduction in length B than in
C, resulting in a decrease in ratio B:C. The
statistical analyses for the ratio measures
appear in Table 1.
To summarize, for animals aged 35 days,
which were sacrificed 14 days postsurgery,
the following observations were made. 1) In
experimental animals the operated side was
characterized by a deformation of the bone
with a slight decrease in length A (involving
Fig. 3. Dried femurs from 42-day-old rats, 21 days
following surgery. Periosteal sectioning was performed
in the midshaft region of the right femur. Left femur
served as intraanimal control. A) ventral view; B) dorsal
view; R) right (experimental side); L) left (control side);
a) the lateral ridge forming the insertion of the Vastus
lateralis and Vastus intermedius in the control limb; b)
the lateral ridge in the operated limb appears to extend
more proximally and laterally; c) some animals exhibited bony alteration at the site of section, while an equal
number show no alteration. The corresponding dorsal
surface appears essentially normal.
Fig. 2. Dried femurs from 35-day-old rat, 14 days following surgery. Periosteal sectioning was performed in
the midshaft region of the right femur only. Left femur
served as intraanimal control. A) ventral view; B) dorsal
view; R) right (experimental side); L) left (control side);
a) the lateral ridge forming the insertion of the Vastus
lateralis and Vastus intermedius in the control limb; b)
the lateral ridge in the operated limb appears more
proximal and exhibits some concavity; c) the site of previous periosteal sectioning appears as roughened new
bone formation on the ventral surface and a smoother
deposition on the dorsal surface.
Figure 3.
344
J.B. McLAIN AND P.S. VIG
22.00
19.00-
21.80
i8.8C18.60-
....
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.. 2-
18.40\
20.20-
18.00-
20 00-
17.80-
19 80-
17.60-
19.60-
17.40-
19.40-
~.-.
1
~
R
L
R
EXP
L
20.8020.6020.4020.20-
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tL
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SHRM
-.
-
17.20'
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21.20:1.00-
18.20-
R
L
L
R
R
DIP
EXP
A
Fig. 4. Length measurements A, B, and C for animals
sacrificed 14 days postsurgery (35 days old). Vertical
scale is in millimeters. Dots represent means, bars represent standard deviations, and dotted lines represent
SHAM
B
mv
L
C
maximum and minimum values. EXP, experimental animals (N = 8); SHAM, sham animals (N = 7); R, right;
L, left. Death following surgery, N = 3.
significantly different from each other, but
were significantly different from the control
animals a t P < 0.0001.
The ratio measurements for animals sacriVariable'
Sum of squares
F value
P > F' ficed 21 days postsurgery are presented
graphically in Figure 7. This figure indicates
A
0.49968
249.84
0.0001
B
2.74523
368.76
0.0001 that right and left femurs exhibited similar
C
2.18508
291.34
0.0001 ratios for the three length measurements for
A:B
0.00417
118.17
0.0001
both the sham and control groups. The experA:C
0.00117
63.33
0.0001 imental group was characterized by dissimiB:C
0.00036
10.34
0.0031
lar ratios for the right and left femurs.
:Degrees of freedom for each variable = 1
In the experimental group length A was
Probability of obtaining the F value by chance
significantly decreased relative to both
lengths B and C, and length C was also sigthe head of the femur) and with a n increase nificantly decreased relative to length B (Tain lengths B and C. 2) In experimental ani- ble 2).
mals measurements B and C were similarly
To summarize, for animals aged 42 days,
affected by the operation. 3) In sham animals which were sacrificed 21 days postsurgery,
the operated side was characterized with a the following was observed. 1)In both the
reduction in all three length measurements sham and experimental animals, the operwith no bony deformation. 4) In sham ani- ated side was characterized by a reduction in
mals lengths A and C were similarly affected all three length measurements. 2) The sham
by the operation, while length €3 appeared to group experienced a proportional reduction
be decreased by the operation to a greater in all three length measurements on the opdegree.
erated as compared to the unoperated side.
At 21 days postsurgery both experimental 3 ) The experimental animals showed a relaand sham groups showed decreased length tively greater decrease in length A than in
dimensions in the operated limbs compared lengths B or C on the operated side. 4)The
to the contralateral unoperated limb (Fig. 6). experimental group also showed a relatively
The control group demonstrated minimal greater decrease in length C than in length
right to left variation in unoperated femurs. B on the operated side.
Measurement A, involving the head of the
DISCUSSION
femur, was diminished to a greater extent in
the experimental group than in the sham;
The results are summarized and presented
however, both groups showed decreased in Figure 8. As predicted from reviewing prelengths when compared to the control group vious investigations, the stimulatory effect
(P < 0.0001; Table 2). For measures B and C of periosteal section on long bone growth was
the experimental and sham groups were not greatest nearest the time of sectioning (2
TABLE 1. Multiple regression analysis results
comparing right to left limb differences between
experimental and sham animals 14 days postsurgery
(35 davs old)
345
TRANSVERSE PERIOSTEAL SECTIONING
1.00
I
l
l
L
R
/
,9s
.R5
.90
an
R
R
L
EXP
,85
L
R
L
R
EXP
1
R
SHRM
EXP
SHRM
AIB
BIC
L
SHAM
A/C
Fig. 5. Ratio measurements A:B, B:C and A:C for animals sacrificed 14 days postsurgery (35 days old).
25.00
22.00
24.50
21.50
24.00
21.00
23.50
20.50
23.00
20.00
22.50
19.50
22.00
19,oo
21.50
18,Sfl
21.00
R
L
c
@
R
L
s
l
y
R
L
w
m
1R.flO
R
L
LXP
R
L
SHAM
A
R
L
W
CONTROL
maximum and minimum values. EXP, experimental animals (N = 8); SHAM, sham animals (N = 8); CONTROL, control animals (N = 20); R, right; L, left. Death
following surgery, N = 2.
TABLE 2. Multiple regression analysis results comparing right to left limb
differences between experimental, sham, and control anirnals 21 days
DostsurLrerv (42 davs old)
Variable'
A
B
C
A:B
A:C
B:C
Contras?
Exp-Control
Sham-Control
Exp-Sham
Exp-Control
Sham-Control
Exp-Sham
Exp-Control
Sham-Control
Exp-Sham
Exp-Control
Sham-Control
Exp -Sh a m
Exp-Control
Sham-Control
Exp-Sham
Exp-Control
Sham-Control
Exp-Sham
1ONTRol
C
B
Fig. 6. Length measurements A, B, and C for animals
sacrificed 21 days postsurgery (42 days old). Vertical
scale is in millimeters. Dots represent means, bars represent standard deviations and dotted lines represent
SilN
Sum of squares
F value
P > F*
4.90671
2.65219
0.24083
1.33203
1.66965
0.01333
1.72811
1.44586
0.00880
0.00169
0.00004
0.00086
0.00122
0.00034
0.00019
0.00000
0.00013
0.00007
457.82
247.46
22.47
65.09
81.59
0.65
326.00
272.76
1.66
17.20
0.39
8.69
41.31
11.47
6.47
0.04
2.73
1.47
0.0001
0.0001
0.0001
0.0001
0.0001
0.4222
0.0001
0.0001
0.2017
0.0001
0.5340
0.0043
0.0001
0.0012
0.0131
0.8375
0.1027
0.2299
'Degree of freedom for each variable = 1.
'Contrast statements comparing two groups at a time: Exp-Control = experimental vs.
Fontrol; Sham-Control = sham vs. control; Exp-Sham = experimental vs. sham.
Probability of obtaining the F value by chance.
346
J.B. McLAIN AND P.S. VIG
1.15
1.10
1.05
R
w
m
EXP
AIB
L
EXP
CONTROL
SHAM
B/C
R
L
SHAM
R
L
CUNTROl
A/C
Fig. 7. Ratio measurements A:B, B:C and A:C for animals sacrificed 21 days postsurgery (42 days old).
,SO
,401
I
30
,20#lo-
._._.
:.:.
......
........
.....
.....
.....
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...
0-
. .
.....
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.....
-.20-
-,30**
-.SO
J
Y
4.1
A
B
14 DAYS
*+
** *
C
A
B
21 DAYS
C
Fig. 8. Mean differences, right minus left femurs for
measurements A, B, and C in experimental (EXP),sham
and control groups, sacrificed at 14 and 21 days postsurery. **, P = 0.05; t i ,experimental and sham groups are
significantly different from one another when double
asterisks are outside the bracket. t , Double asterisks
indicate a significant difference between the starred
group and the control group.
weeks postsurgery), while the effect changed
from stimulatory to inhibitory a t 3 weeks
postsurgery (presumably following the complete healing and reattachment of the periosteum).
At 2 weeks, lengths B and C showed relative increases in the experimental limbs. This
is especially so when compared to the sham
group and is also supported by viewing the
distribution of intraanimal limb variation in
unoperated controls a t 3 weeks.
One may hypothesize that a relationship
exists between the functional demands on
particular segments of the epiphyseal plates
and the effects observed when the periosteum is cut. Bylaner et al. (1981) demonstrated a differential effect of Blount stapling
of human femur epiphyses with a greater
growth-arresting effect medially than laterally. It is interesting to note in this study
the decrease in length A in the experimental
group a t 2 weeks. This measure involves the
head of the femur-an area which one would
suspect is always under compressive loading
force during function in the rat. Measurement C involves the greater trochanter of the
TRANSVERSE PERIOSTEAL SECTIONING
femur, an area associated with the insertions
of the Gluteus medius, Gluteus profundus,
and Piriformis muscles. These muscles as
well as those inserting a t the lesser trochanter and intratrochanteric notch are aligned
in such a fashion a s to produce a natural
tension on the covering periosteum. Measurement B could be considered as middle
ground in terms of tension and compression
between measures A and C. Consistent with
these anatomical observations, length C appears more susceptible to stimulation with
periosteal section a t 2 weeks than does length
B, and length A shows a decrease in length
greater than normal intralimb variation.
This decrease in length A would be attributed to the surgical procedure itself and a
decrease in the normal function of the bone.
While the differences exhibited in this
study were significant, the magnitudes were
small. This is in contrast to Crilly’s (1972)
work using chick radii, where differences as
great as +27.0% for the experimental side
were observed. At 2 weeks the largest differences in this study were about + 1.0% for the
experimental side. Two explanations can be
offered to account for these differences. First,
Crilly compared experimental to sham limbs,
both having had surgery to insert bone
markers; thus he failed to account for the
inhibitory effect of surgery, muscle reflection, and scarring in his “control” limbs. Second, it can be hypothesized that the functional aspects of the bones involved varies
greatly between the two studies. The normal
compressive loading of the bones during
function may be quite different between the
chick radii and rat femur, thus providing
another covariable that is known to be of
importance in the regulation of bone growth.
While only two points in time were examined in this study, what was observed here
was consistent with other investigations indicating a peak of effect a t the time of periosteal reattachment and a decrease in effect as
growth continues. Local bony alterations also
diminished from 2 to 3 weeks postsurgery.
Better defined and larger new bone deposits
were evident a t 2 weeks, while at 3 weeks
remodeling had produced a recovery to essentially normal contour for most of the specimens.
It is tempting to propose that at 3 weeks
postsurgery not only has the stimulatory effect of periosteal sectioning ceased, but that
an inhibitory effect is also emerging related
perhaps to the healing and reattachment of
the periosteum to the femur shaft. Moss
(1978) suggests that in man collagen cross-
347
linking with age reduces periosteal elasticity
to the point that chondrogenesis ceases in the
epiphyseal plates. Further histological investigations are being carried out to determine which changes in the periosteum a t the
site of section posthealing might account for
this restrictive effect. Hellquist’s (1972) observations of thicker, more adherent periosteum in rabbit maxillae following sectioning
may hold the answer. Another observation,
the greater susceptibility of length A to decrease upon surgical intervention in both the
sham and experimental groups, may indicate
a greater dependence on normal function for
the maintenance of length A (involving the
head of the femur).
While supporting both Lacroix (1951),
Crilly (19721, and Moss (19781, who postulated that the periosteal covering of long
bones exerts a restraining force on the epiphyses, this investigation also demonstrates
that the functional demands of individual
long bones may modify the periosteal influence and may play a more dominant role in
the growth regulation of the epiphyses. These
are matters for further investigation.
ACKNOWLEDGMENTS
This project was supported through a National Research Service Award Individual
Fellowship, National Institute of Dental Research grant F32 DE05166.
LITERATURE CITED
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