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Effects of growth and speed on hindlimb joint angular displacement patterns in vervet monkeys (Cercopithecus aethiops).

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 81:441449 (1990)
Effects of Growth and Speed on Hindlimb Joint Angular
Displacement Patterns in Vervet Monkeys
(Cercopithecus aethiops)
JOEL A. VILENSKY AND EVA GANKIEWICZ
Department of Anatomy, Indiana University School of Medicine,
Fort Wayne, Indiana 46805-1499
KEY WORDS
Primates, Locomotion, Ontogeny, Hip, Knee,
Ankle
ABSTRACT
Hip, knee, and ankle joint displacement patterns are compared across both age and speed for five immature vervet monkeys sampled
approximately every 6 months over a 3 year period. The analysis indicated
that, as a group, the animals displayed no consistent changes in joint patterns
as they grew. However, individual animals showed consistent patterns. There
were also no consistent effects of size across animals at the walk-gallop
transition. This is contrary to McMahon’s prediction (J. Appl. Physiol.
39:619427, 1975) based upon his elastic-similarity model of animal scaling.
With increasing speed, when symmetrical gaits were used, all of the animals
tended t o show a decrease in the relative positions of the hip, knee, and ankle
maximum values. Furthermore, across the walk-gallop transition, the animals tended to show a decrease in the range of ankle and knee movements.
Previous analyses of the effects of increas- monkeys sampled every 6 months over aping speed andlor size on joint angular dis- proximately a 3 year period at the identical
placement patterns in primates are ex- speeds. The purpose of the study was to
tremely limited. Vilensky and Wilson (1986) determine if there were consistent changes
found some consistent changes in hip and in these patterns both across time (i.e., with
ankle joint patterns with increasing speed in increases in body mass) and across speed.
one vervet monkey. Reynolds (1987) re- Clearly, by using a longitudinal approach for
ported that hindlimb angular excursions de- this study, problems caused by intersubject
creased with increasing s eed for a variety of variability are controlled.
primates. Vilensky et a! (1988) found no
MATERIALS AND METHODS
consistent effects of body mass on hindlimb
angular excursions in vervet monkeys. HowThe raw material for this paper consisted
ever, they did find some tendency for the of the same 100 frame& films used in our
excursions to increase across slower speeds companion paper (Vilensky et al., 1990).
and decrease a t the walk-gallop transition, These films displayed the animals locomotafter which there was an increase with in- ing on a treadmill at the following speeds:
creasing s eed. McMahon (1984) reported 0.62, 0.89, 1.17, 1.44, 1.72, 1.99, 2.28, 2.58,
for four di erent-sized mammals (none pri- and 2.81 d s . From the “good” strides for
mates) that hindlimb excursions at the trot- each trial (see Vilensky et al., 19891, we
gallop transition equaled 74.4mas~-~.’~.chose one stride to digitize. Thus, for each
Thus the larger animals had smaller angular animal, the following numbers of strides
excursions at this transition. This was in were digitized: A l , 38; A2,52; A3,40; A4,44;
good agreement with his model that predicts A5,43 (see Table 1 in Vilensky et al., 1990).
angles at this transition scale proportionally The specific stride chosen from each trial
to
(McMahon, 1975).
was one that had a cycle duration equal or
For this report we have analyzed in detail closest to the mean cycle duration for all of
the hip, knee, and ankle joint angular displacement patterns in five immature vervet
Received March 27,1989; accepted August 31,1989.
1
@ 1990 WILEY-LISS, INC
442
J.A. VILENSKY AND E. GANKIEWICZ
the strides of that trial. Table 1 in Vilensky
et al. (1990) details the ages and masses of
the animals at each filming. Here, we will
only present the ranges for each animal:
A1 (female), 1.17-3.17 years, 1.27-2.22 kg;
A2 (female), 1.0-3.5 years, 1.31-2.5 kg; A3
(female), 0.83-2.83 years, 1.14-2.27 kg; A4
(female), 0.5-2.83 years, 0.64-1.73 kg; and
A5 (male), 0.33-2.33 years, 0.73-2.00 kg. It
is noteworthy that the youngest animal
filmed was 4 months of age. Because by 2
months of age vervets have the ability to
perform both symmetrical and asymmetrical
gaits in an adult-like fashion (Vilensky and
Gankiewicz, 19891, we do not discuss our
findings with regard to the relative neuromuscular maturity of the animals. That is,
we consider “size” to be the only variable
within individuals at the same speeds.
Once a stride was chosen for analysis, the
individual film frames were rear-projected
onto a digitizing tablet that enabled an experienced technician to estimate visually the
locations of the followingjoint centers (right
limb) for each frame of the stride: metatarsophalangeal, ankle, knee, hip, shoulder, elbow, wrist, and metacarpophalangeal. Specifically, using a cursor, the technician
inputed the estimated locations of these joint
centers into the computer. Subsequently, the
computer determined the ankle, knee, hip,
shoulder, elbow, and wrist joint angle values
(illustrated in Fig. 1) for each frame of film.
For this report, we will only describe the
results based upon the hindlimb angles. The
forelimb data will be reported in a forthcoming paper.
Once determined, the raw angular values
were smoothed using low pass digital filtering (Winter et al., 1974; Winter, 1979) with
cut-off frequencies rangingfrom 6 to 8. Next,
we determined the followingfive parameters
from each of the smoothed curves for each
joint: minimum angle, relative position of
minimum angle (i.e., the time of occurrence
in terms of percentage of cycle), maximum
angle, relative position of maximum angle,
and range. It is important to note that for
some animals at some speeds we changed the
location of the minimum or maximum angle
for the knee or ankle so that it was in agreement with most of the other trials. That is, as
evident from Figure 2, during a complete
locomotor cycle the knee and ankle have two
minima and maxima. For our purposes (and
usually the case), the “true” minimum occurred for both joints during the second half
of the cycle, and the “true” maximum oc-
Fig. 1. Drawing of one of the animals illustrating the joint angles computed. A Ankle; B:
knee; C: hip (to horizontal);D: shoulder (to horizontal);E: elbow; F: wrist.
443
EFFECTS OF GROWTH AND SPEED ON JOINT ANGLES
gaits as well as for the transition. S ecifically, for each animal, we analyze (i.e.,
computed correlation coefficients) across
ages only those speeds for which we had at
least three symmetrical or asymmetrical
gait trials. Similarly, across speeds we analyzed only those ages for which we had at
least three trials of either type of locomotion
(see Table 1 in Vilensky et al., 1990). However, there was an additional restriction on
the asymmetrical strides. We used only
strides that had the same leading limb (regardless of gallop type; i.e., transverse or
rotary). This was done because during galloping the leading and trailing limbs have
been shown to exhibit different angular relationships (Cohen, 1979; Vilensky et al.,
1988). Finally, we analyzed the relationship
between angular excursions and mass at the
walk-gallop transition and the changes in
angular patterns at this transition. In accord
with our companion paper, and because of
the voluminous amount of information produced by this study, we will discuss our
findings for all of the animals but present in
detail only the results for animal No. 5 (A5).
B
35
40
145T
25
KNEE
50
100
/"\
W
75
65
75
0
25
50
75
100
RESULTS
PERCEKT OF CYCE
Fig. 2. Plots of the smoothed hip, knee, and ankle
displacement patterns for A5 at 1.17 m/s at five different
ages (and mass values).
curred at around 50%of the cycle. However,
occasionally the actual minimum occurred
during the first half of the cycle or the maximum a t the beginning or end. We changed
these so that we were comparing similar
turning points for each cycle. We also did not
use the position of the hip minimum value
for any calculations, because the hip minimum was very closely associated with the
beginning of the stride (i.e., limb touchdown).
Because we found previously in vervets a
discontinuity in hindlimb angular excursion
at the walk-gallop transition, and because
back motion significantly contributes to locomotor effort during galloping (Hurov, 19871,
we will present separate discussions of the
data for symmetrical and asymmetrical
Symmetrical gaits
Figure 2 depicts plots of the displacement
patterns of the hip, knee, and ankle angles of
A5 across all ages at 1.17 d s . At this speed
the animal used only symmetrical gaits. The
figure shows that the positions of the minimum and maximum ankle angles tended to
increase with size, whereas the actual values
did not chan e consistently. However, the
maximum an le value was at its lowest at
the second youngest age. For the knee, the
minimum value, position of the minimum
value, and position of the maximum angle
tended to increase with size. Also, again, the
maximum was lowest at the second youngest
age. None of the hip parameters showed any
tendency to change in a consistent manner
with a e, and, in contrast to the ankle and
knee, t e hip maximum was highest
at the
youngest age.
To describe better the relationship between age and angular parameters a i 1.17
m/s as well at other symmetrical ait speeds,
Table 1 presents correlation coe ficients for
A5 between the listed angular parameters
and body mass at each of the different speeds
for which we had sufficient data. The only
relatively consistent trend among the speeds
is for the maximum ankle position to in-
a
a
B
444
J.A. VILENSKY AND E. GANKIEWICZ
TABLE 1. Linear correlation ualues for A5 for listed
angular parameters Vs. body mass at each speed
(symmetrical gaits)'
'lo-
HIP
<'
100-7
90
Speed (m/s)
Parameter
Min Ankle
Pos Min Ankle
Max Ankle
Pos Max Ankle
Range Ankle
Min Knee
Pos Min Knee
Max Knee
Pos Max Knee
Range Knee
Min Hip
Max Hip
Pos Max Hip
Range Hip
0.62
0.89
1.17
1.44
0.92
0.30
-0.52
0.89
-0.84
0.49
-0.43
0.70
-0.24
-0.65
0.16
-0.73
-0.11
-0.63
0.84
-0.01
-0.16
0.92
-0.72
0.88
0.01
-0.81
0.59
-0.98
0.26
-0.93
0.67
-0.70
0.14
0.91
0.54
0.84
0.53
0.93
0.81
0.40
0.88
-0.48
0.21
-0.14
0.60
-0.31
-0.22
0.63
0.79
0.59
0.58
0.42
0.75
0.62
0.31
0.21
-0.16
0.58
0.54
0.74
-
-.
/,: >..,
._
.82 m/s
3 0 m/s .
1.17m/s-.
):.
70.60-7
__-'
40
30
0
25
50
75
100
0
25
50
75
100
'There were five ages available for the computations at 0.62,0.89,
and 1.17 m/s and four at 1.44 m/s.
increase with mass. Similar tables for the
other animals indicated the following:' 1)
The only somewhat consistent trend shown
by A1 was for the minimum ankle angle to
decrease with mass; 2) A2 showed a tendency for the maximum ankle and knee positions and the minimum knee value to increase with mass; 3) A3 showed a tendency
for the minimum hip value to increase with
mass and, accordingly, for the hip range to
decrease; and 4)A4 showed tendencies for
the ankle, knee, and hip ranges to decrease
with mass, and for the minimum knee position also to decrease with mass.
Figure 3 depicts the angular displacement
patterns of the hip, knee, and ankle joints of
A5 at 1.33 years (1.54 kg) across all symmetrical gait speeds. The figure indicates that
the ositions of the maximum ankle and hip
anges and the hip minimum value decreased with increased speed. Additionally,
the ankle and hip maxima reached their
lowest values at the slowest speed (0.62d s ) .
Table 2 depicts the correlation coefficients
for A5 between the available speeds (symmetrical gaits only) and each of the listed
parameters at each age (and associated
mass). Two strong trends are apparent. As
noted for 1.33years in Figure 3, the positions
of the maximum ankle and hip angles decreased with speed for all ages. At 0.33,0.83
and 1.83 years, the position of the maximum
160
150i
ANKLE
\
.I
..u..
140
1 20
._:
80
70
0
25
50
PERCENT OF CYCLE
75
1 oc
Fig. 3. Plots of the smoothed hip, knee, and ankle
displacement patterns for A5 at an age of 1.33years (1.54
kg) across all speeds during which symmetrical gaits
were used.
f
'It should be noted that the trends listed for A1 to A4, as forA5,
were determined subjectivelybased on the consistent appearance
of relatively high and similarly signed (i.e., + or -1 r values.
Because of the low sample sizes, we did not considerit worthwhile
to base our results on only statistically significant values.
knee angle also followed this trend. There
was also a notable tendency for the hip maximum and range to increase with speed.
Evaluation of similar tables for the other
animals indicated the following: 1) In Al,
minimum ankle angle tended to increase
with speed; the positions of the maximum
ankle, knee, and hip angles decreased with
speed; and the minimum knee position also
tended to decrease with speed. 2) In A2,the
positions of the minimum ankle and knee
angles generally decreased with speed; the
maximum ankle angle, ankle range, maximum knee angle, and knee range tended to
increase with speed; and the positions of
445
EFFECTS OF GROWTH AND SPEED ON JOINT ANGLES
TABLE 2. Linear correlation values for A5 for the listed angylar parameters Vs. speed (symmetrical gaits only)
Parameter
Min Ankle
Pos Min Ankle
Max Ankle
Pos Max Ankle
Range Ankle
Min Knee
Pos Min Knee
Max Knee
Pos Max Knee
Range Knee
Min Hip
Max Hiu
Pos Ma; Hip
Range Hip
0.33 (0.73)
0.83 (1.32)
Years of age (mass in kg)
1.33 (1.54)
1.83 (1.77)
2.33 (2.00)
0.98
-0.60
0.65
-0.92
-0.51
-0.67
-0.87
-0.99
-0.99
0.18
-0.75
0.87
-0.96
0.86
0.59
-0.91
0.53
-0.97
0.49
-0.91
-0.64
-0.03
-0.99
0.57
0.36
0.35
-0.97
0.11
0.48
-0.60
0.64
-0.95
0.36
-0.77
-0.32
-0.21
0.38
0.20
-0.93
0.57
-0.99
0.86
0.53
-0.85
0.81
-0.96
0.50
-0.09
-0.90
0.89
-0.85
0.81
0.32
0.91
-0.98
0.78
-0.92
-0.25
0.89
-
-0.96
0.91
-0.68
-0.34
-0.45
-0.26
0.87
0.23
0.93
-0.87
0.81
'There were five speeds available for the computationsat ages 0.83 and 2.33 years, six for 1.83 years, four for 1.33 years, and three for 0.33
years (Vilensky et al., 1989).
TABLE 3. Linear correlation values for A5 for the
listed angular parameters Vs. body mass at each speed
(asymmetrical gaits)'
Parameter
Min Ankle
Min Ankle
Max Ankle
Pos Max Ankle
Range Ankle
Min Knee
Pos Min Knee
Max Knee
Pos Max Knee
Range Knee
Min Hip
Max Hip
Pos Max Hip
Range Hip
1.72
Speed (m/s)
1.99
2.28
2.58
-0.67
0.26
o,99
o,48
0.95
0.78
0.90
0.01
0.98
0.67
0.29
0.93
0.54
0.95
0.92
0.99
0.67
0.76
0.58 -0.70
0.55
0.63
-0.80
0.99
-0.31
0.78
-0.67
0.65
0.31
o,84
0.99
-0.37
0.88
0.75
0.44
0.89
0.93
0.47
0.26
0.89
0.16
0.90
2.81
-0.87
0.11
o.51
0,51
0.94
0.80
0 -0.39
1.0
0.63
0.63
0.90
0.86
0.80
-0.01
0.91
0.87
0.78
-0.76 -0.44
-0.98
0.73
0.86
0.96
0.87
0.25
0.97
0.46
'There were five ages available for the computations at 2.81 m/s,
four for 1.99 and 2.28 m/s. and three for 1.72 and 2.58 m/s.
the maximum ankle, knee, and hip angles
tended to decrease with speed. 3) In A3,
maximum ankle angle and knee range
tended to increase with speed; minimum
knee position and hip minimum tended to
decrease with speed; and the positions of
the ankle, knee, and hip maximum values
strongly decreased with speed. 4) In A4, the
only consistent trends evident for this animal were for the positions of the maximum
ankle and hip angles to decrease with speed.
Asymmetrical gaits
The displacement curves for the hip, knee,
and ankle angles during galloping did not
differ grossly in pattern from those during
walking, and thus no examples are given.
Table 3 presents the correlation coefficients for A5 across age in a similar manner
as in Table 1, but for asymmetrical gaits. The
table indicates that the maximum ankle,
knee, and hip angles generally increased
across increasing mass. The ankle range also
tended to increase, as did the minimum knee
and the positions of the maximum and minimum knee angles. Similar tables for the
other animals showed no consistent trends
(A1 galloped only on one date, so no analyses
were available for her).
Table 4 depicts the correlation coefficients
for A5 in a similar manner as in Table 2, but
for asymmetrical gaits. The table shows a
tendency for the maximum ankle position to
decrease with speed and for the maximum
hip value and range to increase with speed.
Similar tables for the other animals revealed
the following: 1) The only clear trend evident
for A2 was that the position of the maximum
ankle value decreased with speed; 2) A3
showed a tendency for minimum hip angle t o
increase with speed; and 3) for A4, the positions of the maximum ankle, knee, and hip
values decreased with speed, the positions of
the minimum ankle and knee angles also
decreased with speed, and the hip and knee
ranges tended to increase with speed.
Walk-gallop transition
Figure 4 depicts a log-log plot of the hip,
knee, and ankle ranges at the walk-gallo
transition (lowest galloping speed) for eac
animal for each date for which it galloped vs.
mass. Additionally, regression lines and associated r values are depicted for each animal. Furthermore, the line representing the
equation computed by McMahon (1984) for
hindlimb excursion at this transition (based
on a variety of mammals) is plotted on the
K
446
J.A. VILENSKY AND E. GANKIEWICZ
TABLE 4. Linear correlation values for A5 for the listed angular parameters Vs. speed at each age (asymmetrical
paits onhi'
Parameter
Min Ankle
Pos Min Ankle
Max Ankle
Pos Max Ankle
Range Ankle
Min Knee
Pos Min Knee
Max Knee
Pos Max Knee
Range Knee
Min Hip
Max Hip
Pos Max Hip
Range Hip
0.33 (0.73)
0.83 (1.32)
Years of age (mass in kg)
1.33 (1.54)
1.83 (1.77)
2.33 (2.00)
-0.01
0.32
0.48
-0.66
0.81
-0.99
-0.91
-0.66
-0.99
0.86
-0.98
-0.74
-0.99
0.67
0.49
0.55
0.94
-0.89
0.24
0.44
0.18
0.95
-0.62
-0.44
0.10
0.99
0.18
0.69
0.38
-0.71
-0.19
-0.91
-0.42
-0.04
-0.55
0.59
-0.29
0.41
0.32
0.90
-0.42
0.79
0.20
-0.50
0.41
-0.89
0.14
0.16
-0.58
0.70
0.08
0.81
-0.38
0.94
-0.66
0.89
0.88
0.52
0.35
-0.77
-0.94
0.99
-0.94
0.99
-0.77
0.96
-0.35
0.99
-0.99
0.98
'There were qix speeds available for t h e computations a t age 1.33 years, five for 1.83 years, four for0.83 years, a n d three for0.33 and 2.33
years (Vilensky et al., 1989).
hip range graph (see Discussion). Clearly, for
all three joints the relationship between
mass and walk-gallop transition value was
usually poor within animals ke., low r values). Furthermore, the regression lines are
highly inconsistent among the animals.
Table 5 depicts the differences in the noted
angular parameter between the highest
walking (running) speed and lowest galloping speed for A5. A negative value implies a
decrease from the walking to the galloping
speed. For the hip, there were no notable
differences. For the knee and ankle, there
appeared to be consistent (except for age 1.33
years) decreases in range with the transition
to galloping. For the ankle, this was associated with a larger minimum ankle value
(i.e., less ankle dorsiflexion). Tables similar
to Table 5 €orthe other animals indicated the
following: 1)Knee and ankle minimum values increased in A2, and, accordingly, the
ranges decreased. 2) The hip maximum and
range decreased in A3, and the knee and
ankle minimum increased, and the ranges
showed some tendency to decrease. 3) In A4,
the hip minimum increased, and the ankle
maximum and range decreased.
DISCUSSION
Angular parameters and size
The presented data indicate that vervet
monkeys, as a oup, do not use consistent
changes in h i n g m b joint angular parameters to adjust for the changes in their size
with ontogeny during either symmetrical or
asymmetrical gaits. Nevertheless, particular animals did show highly consistentpatterns. Although it is possible that the in-
dividual patterns we recorded occurred
spuriously, the high r values and the consistency across speeds makes this seem unlikely.
In contrast to the present report, as far as
we can determine, no prior studies of either
animal or human locomotion has ever examined ontogenetic changes in joint movements
at identical speeds. Thus these data are not
directly comparable with those from any
other study. However, one previous study
investigated joint angle changes at identical
speeds in different-sized children (i.e., a
cross-sectional study; Vilensky et al., 1987).
The authors reported good relationships
with stature for hip minimum, knee minimum, and knee maximum angular values.
Two studies have reported a remarkable
likeness in joint angular excursions in adults
and children (Foley et al., 1979; Sutherland
et al., 1980), but in neither of these studies
was speed controlled.
Angular parameters and speed
In contrast to the lack of similar relationships across increasing size, the five animals
showed some similarities across speed while
using symmetrical gaits. The animals exhibited almost uniformly a tendency for the
positions of the maximum hip, knee, and
ankle values to decrease with speed. For
example, for A5 at 0.83 years, the maximum
hip position occurred at 61% of the cycle at
0.62 d s but at 43% of the cycle at 1.72 d s .
Individual animals exhibited other tendencies as well, but these were not consistent
among them.
Across the higher speeds, which required
447
EFFECTS OF GROWTH AND SPEED ON JOINT ANGLES
A1
A A2-..
= AA 43---' '
T
1.5
As-I
1.04
I
r=.96. ..'
*A . '
1.44
-0.20
-0.10
0.00
0.10
0.20
0.30
0.40
I
0.50
LOG MASS (kg)
Fig. 4. Log-log plots and regression lines of hip, knee,
and ankle angular ranges at the walk- allop transition
vs. mass. The solid line on the hip grapgh corresponds to
the equation presented by McMahon (19841, which is
based on a variety of different sized mammals (see text).
asymmetrical gaits, the pattern of changes
in angular parameters associated with increases in speed was inconsistent. However,
A4 showed, in accord with the symmetrical
gaits, a decrease in the positions of the hip,
knee, and ankle maximum values.
The fact that the relative positions of the
maximum hip, knee, and ankle values decrease with speed during symmetrical gaits
is not surprising, as these peaks tend to
occur late during stance phase, which also
decreases with increases in speed (Vilensky
et al., 1990). Thus the change in the location
of these peaks probably does not represent
any fundamental change in limb mechanics
associated with increased speed.
Reports detailing the effects of speed on
joint angular displacement patterns in both
humans and animals are quite contradictory, although hip joint amplitude appears
clearly to increase during human walking
and running (Vilensky, 1987). For animals,
the prevailing view seems to be, at least for
hip movements, that there are no dramatic
changes with speed (Grillner, 1975). For example, Halbertsma (1983) stated that, above
1 d s , changes in joint angles in cats with
increases in speed are slight. Our data for A5
presented in Figure 3 suggest this to be true
also for vervet monkeys. That is, the hip and
ankle maxima at 0.62 m / s were notably less
than those at the higher speeds. This was
generally also true for the other animals
(including the knee maximum at midcycle).
Reynolds (1987) reported that the hindlimb
angular excursions of two chimpanzees, a
spider monkey, and a lemur decreased with
increasing speed. Clearly, this finding is not
supported by the current study (at least
TABLE 5. Differences in the noted parameters (in degrees) for A5 between the values at the lowest galloping
speed and the highest walking (runningj speed at each age
Years of age (mass in kg)
Parameter
Ankle
Min
Max
Range
Knee
Min
Max
Range
Hip
Min
Max
Ranae
0.33 (0.73)
0.83 (1.32)
1.33 (1.54)
1.83 (1.77)
2.33 (2.00)
19
1
-18
7
-11
-18
-8
0
8
5
-13
-18
18
-6
-24
16
1
-15
-5
-19
-14
-2
1
-1 1
-12
-4
8
4
-4
-4
5
-9
-5
2
-3
4
6
3
-2
-5
8
-12
2
4
2
448
J.A. VILENSKY AN11 E. GANKIEWICZ
relative to hip range) nor was it supported by seemingly consistent ways to account for the
our cross-sectional study (Vilensky et al., changes in body size associated with growth.
This clearly shows tremendous flexibility
1988).
and/or variability in the neural programs
Walk-gallop transition
that control locomotor behavior.
This study also examined the effects of
As noted previously, McMahon (1975) proposed, based upon his elastic similarity speed on hindlimb joint movements. Furmodel of animal scaling, that angular excur- thermore, the study involved sampling the
sions at the trot-gallop transition should same animals serially over time at the same
scale proportionally to masspo.125.He empir- speeds. This process should have minimized
ically determined that hindlimb excursions the possibility of spurious observations beat this transition are equal to 74.4ma~s-~-l’ing reported as the norm. Our analysis indi(McMahon, 1984). The data presented in cated that the most consistent changes with
Figure 4 do not indicate that, at least within increased speed are decreases in the maxianimals, joint excursions at the walk-gallop mum positions of the hip, knee, and ankle
transition decrease regularly with increases joints. These changes are undoubtedly assoin size. Furthermore, although we did not ciated with the decrease in stance duration
measure hindlimb excursion for this report, associated with increases in speed.
Finally, it is important to emphasize that,
this parameter should be similar to hip
range. Thus the hip excursion plot in Figure despite our many samplings of the animals
4 depicts the line corresponding to McMa- for this study, we only examined one “t phon’s empirically derived equation. It is ap- ical” stride for each animal at each spee at
parent that this equation did not predict each age. It would seem important now to
quantify the variability in these patterns at
accurately the vervet transition values.
In contrast to the present report, for our set speeds on a single date. Additionally, it
cross-sectional study (Vilensky et al., 1988) seems important to investigate the possible
we did measure hindlimb excursions in interrelations between joint displacement
vervet monkeys. Again, however, we could patterns (i.e,, are greater hip excursions asfind no evidence that larger animals have sociated with smaller knee excursions, and
smaller angular excursions at the walk- so forth?) Such projects are necessary to
gallop transition, although the actual range understand more fully the biomechanics and
of predicted values was not too far removed neural control mechanisms underlying prifrom actual values. Lastly, it is noteworthy mate locomotion.
that Alexander (1985) detailed both theoretACKNOWLEDGMENTS
ical and empirical problems associated with
the theory of elastic similarity.
We are grateful to Mr. G. Duncan and Ms.
In our cross-sectional study, we reported P. Wilson for assistance during filming, to
that hindlimb angular excursions tended to Ms. J. Kettelkamp for digitizing the films, to
decrease across the walk-gallop transition. Dr. M. Cartmill and three anonymous reThe present, more detailed analysis shows a viewers for constructive comments on the
fairly regular tendency for the range of knee original version of this paper, and to Ms. D.
and ankle movements to decrease. This is Jackson for typing the many drafts of the
often caused by an increase in the respective manuscript. Funds for this study were supminimum values of those joints (i.e,, less plied by the Indiana University School of
flexion). It seems reasonable to suggest that Medicine.
the increased back motion associated with
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ankle joints (Hurov, 1985,1987).
B
CONCLUSIONS
This study is the first report on the effects
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