MACDOUGALL, J.D.,D. G. SALE, S.E. ALWAY, AND J.R. phy of existing fibers rather than by hyperplasia. The
SUTTON. Muscle fiber number in biceps brachii in bodybuilders training mode in this study, however, was of the endur-
and control subjects. J. Appl. Physiol.: Respirat. Environ. Ex- ance low-resistance type rather than the high-resistance
ercise Physiol. 57(5): 1399-1403, 1984.-Muscle fiber numbers training used in the studies that have reported hyperpla-
were estimated in vivo in biceps brachii in 5 elite male body- sia (8, 9).
builders, 7 intermediate caliber bodybuilders, and 13 age- Competitive bodybuilders possess extreme degrees of
matched controls. Mean fiber area and collagen volume density
muscle hypertrophy and thus are excellent models for
means of a modified Gomori trichrome stain for myofi- cross-sectional muscle area
brillar adenosinetriphosphatase (following preincubation (corrected for connective tissue)
at several pHs) before being photographed under the numbers of fibers =
average fiber area
light microscope. Cross-sectional area for both type I and
type II fibers (as classified by the pH 10.0 preincubation) where
was measured by a computerized digitizer for a minimum average fiber area
of 200 fibers of each type per biopsy. Percent fiber type
was estimated by counting a minimum of 550 fibers/ (% type I)(mean type I area)
biopsy. The proportion of collagen and other noncon- -t (% type II) (mean type II area)
tractile tissue was calculated in the trichrome-stained =
100
sections by means of a 168 point, point-counting tech-
nique (20). and connective tissue correction equals the total muscle
Cross-sectional area of biceps was determined from area minus the percentage of muscle that is collagen and
CT scans (Ohio Nuclear, model 20-30) of the upper arm other noncontractile tissue.
with the elbow extended (Fig. 1). Several scans were To evaluate the reliability of this method for the
taken at l-cm intervals-one at the estimated point of estimation of fiber numbers, biopsies were also taken
greatest circumference for biceps and one proximal and from both arms of seven of the control subjects and fiber
distal to that point. The CT scan negatives were printed number estimated for both right and left arms.
to 17 X 22 in. prints and biceps area measured by
computerized digitizer. The scan yielding the greatest RESULTS
area was then selected.
Muscle fiber number was calculated as follows Reliability of fiber enumeration technique. Based on the
FIBER NUMBER IN BICEPS BRACHII 1401
assumption that there is an equal number of fibers in groups. Average fiber areas for both bodybuilder groups
the left and right biceps, the reliability of the technique were significantly greater (P c 0.02) than for the control
for estimating fiber numbers is illustrated in Fig. 2. The group. Although the average fiber area of the elite group
correlation coefficient for fiber number for left and right appeared to be -13% larger than that of the intermediate
biceps for the seven control subjects was 0.79 (P < 0.01) group, the difference was not statistically significant.
with an SEE of t 11%. Stated in other terms, the mean There was a very wide interindividual range in esti-
intrasubject difference between right and left arms with mated fiber numbers in all three groups. Fiber numbers
this technique was t 8.9% with a t 3.6% SD. ranged from 172,000 to 381,000 among the control sub-
PhysicaL characteristics. The physical characteristics jects, from 198,000 to 374,000 among the intermediate
of the three groups of subjects are presented in Table 1. bodybuilders and from 204,000 to 419,000 among the
Both groups of bodybuilders had been training inten- elite subjects. The mean results for each group, however,
sively for approximately the same period (6-8 yr). The did not differ (Table 1 and Fig. 3).
elite group was significantly heavier (P < 0.001) and had Connectiue and noncontractile tissue. The proportion
significantly larger biceps areas (P < 0.005) than both of connective and other noncontractile tissue was ex-
the intermediate and control groups. The intermediate tremely consistent between individuals and was -13% of
bodybuilder group also had significantly larger biceps the tissue sample. Of this, -6% was identified as collagen
areas (P < 0.005) than did the untrained control group. and 7% as other tissue.
Four of the five elite bodybuilders admitted to the use of
anabolic steroids, as did four of seven intermediate bod-
1. Physical description, biceps area, average fiber area, ana esttmatea numoer
, l ,
TABLE OJ Jzoers
for elite and intermediate bodybuilders and for untrained controls
Elite bodybuilders (n = 5) 23.7 6.2 178 90.3 2487" 13.0 105.6d 213.8 290.7
Al.2 to.7 Ik2.2 t2.9 t181 t3.6 k28.2 t53.2 k72.2
Intermediate bodybuilders (n = 7) 24.2 7.8 173 81.1" 1925b*C 12.1 93.7d 190.8 259.5
rt3.1 k4.4 t3.2 t4.0 k321 t5.4 k27.3 t51.3 t69.7
Untrained controls (n = 13) 22.5 0 179 78.2 1412b 13.4 67.0 204.8 278.5
to.5 k3.1 t2.3 t291 t2.3 k19.6 t44.6 k60.7
Values are means t SD. Fiber numbers are expressed both uncorrected and corrected for change in sarcomere. Length with needle biopsy
technique. a See text for explanation. bP -=I0.005. "P e 0.005. dP e 0.02. "P e 0.001.
1402 MAcDOUGALL, SALE, ALWAY, AND SUTTON
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CHAO. Muscles across the elbow joint: a biomechanical analysis. J. 12. HUXLEY, H. E. Structural evidence concerning the mechanism of