Regulation of fiber size, oxidative potential, and

capillarization in human muscle by resistance exercise

H. Green, C. Goreham, J. Ouyang, M. Ball-Burnett and D. Ranney
Am J Physiol Regul Integr Comp Physiol 276:R591-R596, 1999.

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Regulation of fiber size, oxidative potential, and
capillarization in human muscle by resistance exercise
H. GREEN, C. GOREHAM, J. OUYANG, M. BALL-BURNETT, AND D. RANNEY
Department of Kinesiology, University of Waterloo, Waterloo, Ontario, Canada N2L 3G1
Green, H., C. Goreham, J. Ouyang, M. Ball-Burnett,
and D. Ranney. Regulation of fiber size, oxidative potential,
and capillarization in human muscle by resistance exercise.
Am. J. Physiol. 276 (Regulatory Integrative Comp. Physiol.
45): R591–R596, 1998.—To examine the hypothesis that
increases in fiber cross-sectional area mediated by high-
resistance training (HRT) would result in a decrease in fiber
capillarization and oxidative potential, regardless of fiber
type, we studied six untrained males (maximum oxygen
consumption, 45.6 6 2.3 ml · kg21 · min21; mean 6 SE) partici-
pating in a 12-wk program designed to produce a progressive
hypertrophy of the quadriceps muscle. The training sessions,
which were conducted 3 times/wk, consisted of three sets of
three exercises, each performed for 6–8 repetitions maximum
(RM). Measurements of fiber-type distribution obtained from
tissue extracted from the vastus lateralis at 0, 4, 7, and 12 wk
indicated reductions (P , 0.05) in type IIB fibers (15.1 6 2.1%
vs. 7.2 6 1.3%) by 4 wk in the absence of changes in the other
fiber types (types I, IIA, and IIAB). Training culminated in a
17% increase (P , 0.05) in cross-sectional area by 12 wk with
initial increases observed at 4 wk. The increase was indepen-
dent of fiber type-specific changes. The number of capillaries
in contact with each fiber type increased by 12 wk, whereas
capillary contacts-to-fiber area ratios remained unchanged.
In a defined cross-sectional field, HRT also increased the
capillaries per fiber at 12 wk. Training failed to alter cellular
oxidative potential, as measured by succinic dehydrogenase
(SDH) activity, regardless of fiber type and training duration.
It is concluded that modest hypertrophy induced by HRT does
not compromise cellular tissue capillarization and oxidative
potential regardless of fiber type.
training; resistance; hypertrophy; fiber type
ONE OF THE MOST CONSPICUOUS adaptations accompany-
ing high-resistance training (HRT) is an increase in the
cross-sectional area of the muscle cell (13). The cellular
hypertrophy appears to extend to both major fiber types
(type I and type II) and subtypes (IIA and IIB) (8, 21,
32, 33), although the magnitude of the increase appears
to be fiber type specific (8, 14, 33). The increase in fiber
size results in an increase in muscle force-generating
potential (13).
Somewhat unclear is the role of HRT in muscle cell
capillarization and oxidative potential, two properties
that appear to be intimately related in promoting
increased oxidative phosphorylation during exercise
(11, 27). Earlier studies based on comparisons between
weight lifters and age-matched, untrained controls
have indicated that fiber hypertrophy occurs in the
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absence of parallel increases in oxidative potential (2,
18, 36) and capillarization (37). However, other studies
(28, 30) have challenged these conclusions. The limited
number of longitudinal studies, which have employed
HRT for up to 20 wk, are inconclusive. Some studies
have reported no changes in capillary density, ex-
pressed as either capillaries per fiber or per unit fiber
area (16, 35, 38), whereas others (8, 21) have found
increases in the number of capillaries per fiber, both
type I and type II, without increases in capillaries per
unit fiber area. In general, changes in mitochondrial
potential have not been reported with HRT. Tesch et al.
Downloaded from ajpregu.physiology.org on November 22, 2011
(35) and Wang et al. (38), using the maximal activity of
citrate synthase in homogenates as a measure of oxida-
tive potential, have found no change in either women or
men after HRT. Such findings have also been confirmed
at the single-fiber level, regardless of fiber type (24).
These findings suggest that the absolute number and
size of mitochondria increase on an absolute basis to
maintain a constant relative potential. This hypothesis
has been confirmed by Wang et al. (38) by using
ultrastructural measurements. However, interpreta-
tion of the results of most studies is confusing because
hypertrophy was not demonstrated (16, 24, 35).
A number of stimuli have been identified in promot-
ing angiogenesis, and ischemia and hypoxemia have
been recognized as most potent (6, 15). In addition,
mechanical factors such as increased wall tension,
increased sheer stress, and increased intravascular
pressure appear to be important (12). What factors are
involved in inducing capillarization with regular exer-
cise is uncertain. It is known that prolonged, submaxi-
mal exercise training represents a major stimulus for
angiogenesis and mitochondrial biogenesis (12), suggest-
ing that sustained elevation of one or more of the
stimulating factors may be critical. It has been hypoth-
esized that the sustained elevation in blood flow which
occurs during prolonged exercise and which is designed
to meet the nutrient demands of a persistently contract-
ing muscle may be of importance in inducing changes in
stimuli necessary for capillary growth (12). If such is
the case, few repetitions of heavy resistance exercise,
typically employed for HRT, should have minimal effect
on angiogenesis.
In this study, we hypothesized that the HRT program
would lead to an increase in cellular cross-sectional
area in the absence of an increase in capillary number.
As a consequence, the capillaries subserving a defined
cellular area would decrease. Moreover, the decrease in
perfusion potential would be accompanied by a de-
crease in cellular oxidative potential. All of these
changes would be manifested in the major fiber types
and subtypes.
R591
R592 FIBER TYPE CAPILLARIZATION AND RESISTANCE TRAINING
METHODS
Subjects. Six healthy but untrained male volunteers (age
19.2 6 0.48 yr; wt 81.0 6 6.0 kg; means 6 SE) completed all
phases of the study. A substantially greater number (n 5 11)
were recruited, but with attrition and limitations in tissue
availability a complete data set for investigating histochemi-
cal features was restricted to six subjects. Based on the
information provided by questionnaire, none of the volun-
teers was ‘‘trained,’’ i.e., none was involved in exercise on a
regular basis or had been for at least 3 mo before the
beginning of the study. Peak aerobic power, determined
during a progressive cycle test to fatigue, was 45.6 6 2.3
ml · kg21 · min21. As required, all experimental procedures and
risks were described to each volunteer, and the study was
approved by the Office of Human Research.
Experimental design. The training model used in this study
consisted of performing three different exercises, specifically
chosen for their emphasis on the quadriceps muscle. The
exercises (parallel squats, incline leg presses, and leg exten-
sions) were performed 3 times/wk with supervision. At least
one day of rest was provided between each work session. Each
training session included three sets of each exercise with each
set performed for 6–8 repetitions maximum (RM). An approxi-
mate 2-min recovery period was provided between each set.
For safety, a warm-up set of 10 repetitions at 50% of each
subject’s 6–8 RM was included for each exercise. The training
extended for 12 wk with training intensity adjusted after 4
and 7 wk. On average, 10.1, 5.6, and 15.4 training sessions
were performed during each training segment. During the
first training segment, the weights (kg) used were 90.5 6 9.1,
165 6 10, and 40.9 6 2.7 for the squats, leg presses, and leg
extensions, respectively. For the final training segment, the
loads (kg) were increased to 146 6 12, 285 6 23, and 63.6 6 5,
respectively, for each of the exercises. Before the training and
at 4, 7, and 12 wk, samples of tissue were obtained from the
vastus lateralis muscle using the technique of Bergström (3).
The tissue obtained was oriented under a dissecting micro-
scope, mounted, rapidly frozen in isopentane cooled to liquid
nitrogen temperatures, and stored at 280°C until analysis.
Two tissue samples were obtained from each leg during the
study with the order randomized between legs after each
training segment. All tissue samples were obtained at least
24 h after the last training session.
Analytical procedures. Histochemical properties were deter-
mined on cross sections of tissue cut in a cryostat maintained
at 220°C. Muscle fiber-type identification and typing was
based on the procedure of Brooke and Kaiser (5) as modified
by Staron et al. (29). This modification allows identification of
type I fibers as well as several type II subtypes: IIA, IIB, IIAB,
and IIC. On average, ,200 fibers from each tissue sample
were used to determine fiber-type distribution. Staining for
fiber capillarization was performed on 8-µm-thick cross sec-
tions using a lectin (Ulex europeaus I ) technique described by
Parsons et al. (22). In addition, the oxidative potential of each
fiber was determined on cross sections (10 µm) stained for
succinic dehydrogenase (SDH) activity. Determinations of
SDH activity were based on the procedure of Pette (23) using
a reaction medium containing 100 mM sodium phosphate
buffer (pH 7.6), 1.0 mM sodium azide, 0.2 mM 1-methoxy-5-
methylphenazinium methyl sulfate, 50 mM succinate, and
1.5 mM nitro blue tetrazolium (NBT). Nonspecific reduction
of NBT was determined by incubating several sections in the
above medium with succinate. The reaction was allowed to
run for 10 min at 25°C. SDH measurements were based on a
single end point (7). For quantification, the optical density
(OD) measured from the blank, representing the mounting
medium, glass slide, and coverslip just outside the section
being measured, was subtracted from the OD measure in the
fiber. In addition, we also subtracted the change in OD due to
the nonspecific reduction of the reaction indicator NBT (4).
The value for this nonspecific reduction was obtained on
separate pieces of tissue that had been stained under condi-
tions as nearly identical as possible. The OD measured on 25
fibers of each type, where possible, was obtained using an
image analysis system (Image-Pro Plus; Media Cybernetics,
Silver Spring, MD). Fiber-type differences in SDH activity,
obtained before training, were very similar to those obtained
by others using a similar procedure and similar sex (24).
Fiber type-specific areas were obtained from the SDH stain
using the same fibers selected for the OD measurements. The
image analysis system, equipped with a video monitor, digitiz-
ing tablet, and tracing cursor, was used for area measure-
ments. Serial sections permitted the identification of fiber
area and fiber capillarization in the same fiber, which could
then be identified as to type.
Capillarization was determined using two approaches, i.e.,
at the level of the individual fiber and in a defined field of a
Downloaded from ajpregu.physiology.org on November 22, 2011
fixed, cross-sectional area. At the level of the individual fiber,
the capillarization indexes included the number of capillaries
in contact with each fiber type (I, IIA, IIAB, and IIB) or
capillary contacts (CC), and the number of capillary contacts
per fiber-type area. Capillary contacts per fiber were deter-
mined by counting the number of capillaries around each
individual fiber, subdivided by type, and then computing the
mean (9, 21, 25). The number of capillaries per fiber area was
estimated by dividing the number of capillaries in contact
with each fiber by the area of the fiber to which they were
adjacent and computing the mean number of capillaries per
fiber area for each fiber type (21, 25).
The indexes that were calculated from a defined cross-
sectional area of the tissue sample were the number of fibers
or fiber density (FD), the total number of capillaries or
capillary density (CD), the number of capillaries around the
fiber (CAF), and the sharing factor (SF). Both FD and CD
were obtained by simply counting the total number of fibers
and the total number of capillaries, respectively, in the set
cross-sectional area (21, 25). The CAF index was obtained as
the ratio CD/FD (9, 21, 25). The determination of SF (fibers/
capillary) was based on the quotient obtained by dividing the
number of capillaries around the fiber calculated from the
defined field into the average number of capillary contacts
(CC/CAF) (9, 25). For all of these measurements, discrete
regions with clear cross-sections free of artifacts were used.
The number of fibers and capillaries on the borders were
counted and then halved.
Statistical procedures. To determine the effects of training
time (0, 4, 7, 12 wk) and fiber-type (type I, IIA, IIB, IIAB)
distribution, area, and capillarization, a two-way ANOVA for
repeated measures was employed. Where only a single mea-
sure was obtained, for example with capillary density, with-
out regard to fiber-type differences, a one-way ANOVA was
used. Where significance was indicated, post hoc analysis
with the use of the Newman-Keuls technique was performed
to compare specific means. A 95% level of confidence was
accepted for all comparisons.
RESULTS
Muscle fiber-type distribution (%) for the vastus
lateralis muscle before training was 47.2 6 3.1, 28.4 6
4.8, 7.7 6 1.3, and 15.1 6 2.1 for type I, IIA, IIAB, and
IIB, respectively (Table 1; see Fig. 3). With the excep-
tion of the type IIB fibers, training was without effect in
 FIBER TYPE CAPILLARIZATION AND RESISTANCE TRAINING R593

Table 1. HRT and fiber-type distribution Table 3. HRT and capillary contacts per fiber
HRT Time, wk HRT Time, wk

0 4 7 12 0 4 7 12

Fiber-type dis- Fiber type

tribution, % I 5.38 6 0.22 5.61 6 0.34 5.94 6 0.23 6.38 6 0.19
I 47.263.1 50.063.5 49.262.7 48.962.1 IIA 5.54 6 0.44 5.27 6 0.38 5.84 6 0.33 6.18 6 0.24
IIA 28.464.8† 33.062.9 33.061.7† 32.561.4†‡ IIAB 4.16 6 0.28 5.92 6 0.49 5.41 6 0.30 5.77 6 0.46
IIAB 7.761.3†‡ 6.361.0†‡ 7.161.1†‡ 8.361.3†‡ IIB 4.20 6 0.51 4.25 6 0.34 4.97 6 0.24 5.23 6 0.41
IIB 15.162.1†‡§ 8.361.2*†‡ 9.961.8*†‡ 7.261.3*†‡
Values are means 6 SE no. of capillary contacts per fiber; n 5 6
Values are means 6 SE; n 5 6 subjects. HRT, high-resistance subjects. Main effects (P , 0.05) for both training and fiber type were
training. Significant differences (* P , 0.05): * from 0 wk; † from type I; found. For training, 0 wk 5 4 , 12 wk. For fiber types, type I 5 type
‡ from type IIA; § from type IIAB. IIA . type IIAB . type IIB. No interaction effect was found.

Additional indexes of fiber capillarization based on a

altering the percent composition. For the type IIB
fibers, a pronounced reduction ranging from 34 to 45%
was observed. The reduction was fully manifested by 4
wk of training. Although type IC and IIC fibers were
identified, these represented ,1% of the fiber pool and
defined field are presented in Table 4. Only one index,
namely CAF, was altered by training. In the case of
CAF, a 33% increase was observed, but not until the
12th week of training.

Downloaded from ajpregu.physiology.org on November 22, 2011

consequently were not included in the analyses.
HRT increased fiber area (Table 2). This effect, which
was not specific to fiber type, was observed by 7 wk of
training and persisted through the final 5 wk of train-
ing. After subjects were trained for 12 wk, fiber areas,
when averaged over all fiber types, were larger than at
4 wk of training. When comparisons were made be-
tween the fiber types, type IIA fibers were observed to
possess the largest area while type I fibers were the
smallest. No difference was observed between the type
IIAB and type IIB fibers in area. Training did not alter
the differences between the fiber types in area.
Training also had a significant effect in increasing CC
(Table 3; see Fig. 3). This effect, which was not specific
to fiber type, was not observed until the 12th week of
training. In general, type I and type IIA fibers pos-
sessed the largest number of capillary contacts per
fiber, followed by the type IIAB fibers, which were
intermediate, and the type IIB fibers, which were the
lowest. When the changes in fiber-type area were
considered, and the number of capillary contacts were
expressed relative to fiber area, no effect of training
was observed (Fig. 1). Type I fibers possessed the
highest capillary contacts-to-fiber area ratio compared
with the other types. No differences in this ratio were
found between the type IIA, type IIAB, and type IIB
fibers.
SDH activity, determined by microphotometry and
expressed in absolute units, was observed to be highest
in type I fibers, followed by type IIA and type IIB fibers,
and lowest in type IIB fibers (Figs. 2 and 3). The
oxidative potential was not affected by HRT, regardless
of fiber type and training duration.
DISCUSSION
We have been able to confirm, as hypothesized, that
HRT would result in an increase in fiber cross-sectional
area. This finding is not surprising given the many
previous studies (13) that have found fiber hypertrophy
with HRT. Our findings on the time course response are
also consistent with earlier work (31), namely that
increases in fiber area are a delayed response to
training, only observable after several weeks of train-
ing. In this study, significant increases were not ob-
served until the seventh week of training. Our HRT
program failed to produce a differential increase in area
between the different fiber types. By 12 wk of training,
increases in area between fiber types ranged between
14 and 24%. Although similar findings have been
reported previously (10, 14, 21), exceptions have been
noted (8, 33). Several studies have reported a differen-

Table 2. HRT and fiber-type areas

HRT Time, wk

0 4 7 12

Fiber-type area,
µm2
I 4,840 6 145 5,192 6 53 5,487 6 153 5,518 6 125
IIA 6,455 6 196 6,624 6 403 7,274 6 431 7,978 6 208
IIAB 6,065 6 658 6,165 6 248 6,758 6 618 6,901 6 392
IIB 5,577 6 194 5,882 6 173 6,348 6 198 6,355 6 421
Values are means 6 SE; n 5 6 subjects. Main effects (P , 0.05) for Fig. 1. Capillary contacts-to-fiber area ratios in untrained males
both training and fiber type were found. For training, 0 wk , 7 wk 5 during high-resistance training (HRT). Values are means 6 SE; n 5
12 wk; 4 wk , 12 wk. For fiber type, type IIA . type IIB 5 type 6. Only a main effect (P , 0.05) of fiber type was found: type I . type
IIAB . type I. IIA 5 type IIAB . type IIB.
R594 FIBER TYPE CAPILLARIZATION AND RESISTANCE TRAINING
Table 4. HRT and fiber capillarization indexes
in a defined field
HRT Time, wk
0 4 7 12
CD, capillaries/mm2 556641 564635 643648 625633
FD, fibers/mm2 30269.6 292611 288611 262614
CAF, capillaries/fiber 1.8060.14 2.0160.12 2.1260.13 2.4060.10*
SF, fibers/capillary 2.9160.15 2.6260.13 2.6960.16 2.5860.07
Values are means 6 SE; n 5 6. CD, capillary density; FD, fiber
density; CAF, capillaries per fiber; SF, sharing factor. * Significantly
different from 0 wk (P , 0.05).
tial hypertrophy between fiber types with the increase
in size more pronounced in the type II fiber subtypes (8,
10, 14, 33). The differences between the fiber-specific
response between studies may well reflect the specifics
of the different HRT programs (8, 14), including the
duration of the training. Indeed, the relatively brief
nature of previous training programs may be an impor-
tant factor in the inability to observe hypertrophy (16,
24). It must be emphasized that the objective of our
study was not to examine the efficacy of different HRT
programs in inducing fiber hypertrophy. Rather, we
were simply interested in selecting a program that
would result in hypertrophy based on previous pub-
lished reports. This was necessary so that we could
examine the relationship between hypertrophy, capillar-
ization, and oxidative potential.
HRT clearly resulted in an increase in the number of
capillaries in contact with each fiber. However, this
response was more delayed than the increases in fiber
area and not observed until the 12th week of training.
Similar to the fiber size changes, the magnitude of the
effect was not differentiated by fiber type. As with HRT
effects on fiber area, the delayed increase in capillariza-
tion may well explain the failure of a previous study
(16) to observe changes because this program was
shorter in duration than the present program, and of
others (8, 21) in which increases in angiogenesis have
been observed. Even with extended training, increases
in capillary number are not always a consistent finding
(38).
Fig. 2. Fiber-type oxidative succinic dehydrogenase (SDH) activity in
untrained males during HRT. Values are means 6 SE; n 5 6. Only a
main effect (P , 0.05) of fiber type was found: type I . type IIA 5 type
IIAB . type IIB. OD, optical density.
Despite the different time course that was observed
between the changes in fiber size and the changes in
fiber capillarization, capillarization as expressed by
capillary contacts-to-fiber area ratio was not altered
with HRT regardless of the period of training. Our
findings demonstrate that modest increases in fiber
area can occur in HRT without compromising the fiber
area served by a capillary regardless of fiber type.
Although previous longitudinal studies have reported
similar findings (8, 21, 38), there is also evidence that
HRT can compromise the capillary-to-fiber area ratio
(8). These discrepancies may be explained by the type of
HRT program. Training using concentric contractions
only, compared with training using concentric and
eccentric contractions similar to HRT with free weights,
does not increase capillary-to-fiber area ratios. Capillar-
ization was also examined from another perspective,
namely by examining capillary density in a defined
cross-sectional area of tissue. This procedure conveys
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the advantage of examining the changes from the
perspective of a defined field without regard to fiber-
type distribution. By using this procedure, neither the
total number of capillaries nor the number of fibers was
significantly altered. However, trends were clearly evi-
dent toward a decrease in the number of fibers and an
increase in the number of capillaries, as might be
expected. It is probable that the magnitude of the
hypertrophy was insufficient to reduce the number of
fibers occupying a given area. A more sensitive index,
namely CAF, was increased by 12 wk of training. The
change in this index was not accompanied by signifi-
cant changes in SF, indicating that, on average, HRT
was without effect in changing the number of fibers
sharing one capillary. An increase in this index could
conceivably compromise nutrient supply to specific
fibers.
It should be emphasized that the measures of capillar-
ization examined are only indexes that provide for the
description of muscle capillaries relative to fiber areas
and types. These properties may not provide the best
estimate of the potential for blood-to-tissue exchange in
skeletal muscle since they do not account for the
configuration of the capillary network (20). A new
property, described as the capillary-to-fiber perimeter
ratio, has been proposed that incorporates the effects of
capillary tortuosity and branching and may provide a
better functional measure (9, 20).
Oxidative potential, as measured by SDH, remained
unchanged throughout the training regardless of the
fiber type. Only one previous study (24) has examined
oxidative potential in the various fiber types in re-
sponse to HRT and arrived at the same conclusions.
However, in the study by Ploutz et al. (24), HRT failed
to result in fiber hypertrophy. These results are also
supported by others using determinations of maximal
enzyme activities of representative mitochondrial en-
zymes in homogenates prepared from whole muscle
samples (35, 38). Ultrastructural studies have provided
inconsistent results with either no change (16, 38) or a
decrease (17, 19) in the percentage of fiber volume
occupied by mitochondria reported. At present, the
 FIBER TYPE CAPILLARIZATION AND RESISTANCE TRAINING R595

Downloaded from ajpregu.physiology.org on November 22, 2011

Fig. 3. Cross sections of muscle samples extracted from vastus lateralis muscle at 0, 4, 8, and 12 wk of training.
Serial sections stained for myofibrillar ATPase (A), SDH (B), and capillarity (C). Muscle tissue obtained from a
single subject. See METHODS for further analytic details.

reasons for the differences between the ultrastructural
studies are not clear. However, the muscle group se-
lected for training may be involved. The work per-
formed by MacDougall et al. (17, 19) was based on the
triceps brachii, whereas others have examined the
quadriceps (16, 38).
As with other studies (8, 24, 31, 33, 38), we have
found reductions in type IIB fiber percentage with
HRT. This effect was observed very early in training
and before changes in fiber area and capillarization. It
is generally believed that the type IIB fibers, which are
composed of myosin heavy chain IIb (MHCIIb) (34)
undergo a sequential transformation in expression to
type IIAB (composed of MHCIIa and MHCIIb), then
type IIA (composed of MHCIIa), and ultimately ending
up as type I fibers expressing MHCI. In this study, the
increases in type IIA, type IIAB, and type I fiber
percentages were not significant. The failure of HRT to
elicit an increase in type I fiber percentage is a consis-
tent finding (8, 14, 24, 31, 33). Although increases in
type IIA fibers have been reported (1, 8, 33) such is not
always the case (31), conceivably because the relatively
low percentage of type IIB are transformed between
type IIA and type IIB.
Of particular interest is the implication of the trans-
formation of the type IIB fibers to the other fast
subtypes regarding the changes in other properties.
The transformation in itself, given the smaller fiber
area, lower capillary counts, and oxidative potential,
could have the effect of altering these properties in the
type IIA and type IIAB fibers independent of actual
changes in these properties. The magnitude of this
effect remains to be determined.
In summary, using a 12-wk program of HRT, we
demonstrated that modest fiber hypertrophy, typical of
that observed with many HRT programs, can occur
without compromising either the capillarization or the
oxidative potential of the different fiber types. Accord-
ingly, the hypothesis that we have proposed, namely
that decreases in both of these indexes would occur
with HRT-induced hypertrophy, must be rejected. The
specific mechanisms eliciting the absolute increase in
both angiogenesis and mitochondrial biogenesis with
HRT remain to be determined but would appear to be
R596 FIBER TYPE CAPILLARIZATION AND RESISTANCE TRAINING
associated with the elevated metabolic rate and blood
flow, and the cellular hypoxemia that occur (6, 15, 20,
26). Our findings do not exclude the possibility that
other manipulations of HRT may result in fiber hyper-
trophy accompanied by either increases or decreases in
capillarization and oxidative potential. These studies
remain to be done.
This work was supported by the Natural Sciences and Engineering
Research Council (NSERC) and Sport Canada.
Address for reprint requests: H. Green, Dept. of Kinesiology, Univ.
of Waterloo, Waterloo, Ontario, Canada N2L 3G1.
Received 10 July 1998; accepted in final form 22 September 1998.
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