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performance
during
11 i/ham
M Sherman,
ABSTRACT
diets
J Andrew
The
on muscle
muscle glycogen,
7 d of training13
effects
glycogen
Dole,
David
of moderate-
and
R La,nb,
and
or high-carbohydrate
performance
in runners
and
and exercise
Richard
H Strauss
centration
cyclists
on training
uncertain
over 7 consecutive
days
biopsies
were performed
of training
were determined.
Muscle
on 4 separate
days before
exercise
for
(5 g carbohydratekg
I h at
75i)
consumption
hydrate
1-mm
sprints.
peak
or cycled
cyclists
After
the
training
to exhaustion
and
was
(V02)
session
at 80%
runners
peak
<
by five,
7, subjects
ran
glycogen
for
. Muscle
with
(P
reduced
30-36%
diet. All subjects
on day
V02
maintained
followed
with the
all training
0.05)
moderatesessions.
This
cogen,
7.
on trainAm J
athletic
training,
exercise
Introduction
exercise
at intensities
70-80%
substantially
(VOmax)
of maximal
lowers
muscle
ox-
glycogen
concentrations
( 1 ). Associated
with reduced
muscle
glycogen
is
the inability
to maintain
exercise
intensity
at 70-80%
VO2max
( I ).It is usually
assumed
that muscle
glycogen
must be restored
between
daily
pabilities
The
training
sessions
carbohydrate
content
concentration.
Costill
cogen
progressively
declined
training
spite
to facilitate
optimal
training
ca-
(2. 3).
cogen
when
of the
subjects
43%
ofathletes
diets
affects
muscle
gly-
et al (4) reported
that
muscle
gly-
over
selfselected
decline
3 consecutive
a 43%
in muscle
d of running
carbohydrate
glycogen
over
diet.
those
of fatigue
over the days
inferred
that consumption
amounts
ofcarbohydrate
and thereby
impairs
In
days,
of trainof a
sociation
appears
.,1n1
between
clear
J C/in Nuir
but
training.
In this rowing
study
rowing
ability
was maintained.
dietary
the
effects
1993:57:27-31.
carbohydrate
and
of reduced
Printed
effects
and
high-
) carbohydrate
and
and
diets
maximal
cyclists over
of moderate(10
on
g carbo-
muscle
gly-
performance
7 d of intense
capa-
training.
muscle
glycogen
Thus,
the asmuscle
muscle
in USA.
men
volunteered
to serve
according
an honorarium
to institutional
for their
as subjects
provided
This project
was approved
by the Biomedical
Subjects
Review
Committee
of The Ohio
Subjects
previous
those
subject
also
was
applicable
The physical
provided
termine
phase.
weekly
imposed
training
required
to achieve
exercise
mode.
and physiological
in Table
loads
by the experimental
a V02
Each
in the
Sciences,
State
Uni-
were
similar
protocols.
peak
characteristics
1. Hydrostatic
to
Each
4 L/min
of the
weighing
was
for
subjects
used
to de-
Repeated
measures
of vital capacity
were determined
until the variability
The
and
guidelines.
participation
means
for the
three
highest
and hydrostatic
of the readings
values
Siri
equation
(7) was used to calculate
Exercise
mode-specific
peak oxygen
was determined
by using a progressive
also allowed
determination
maximal
heart
rate. V02
open-circuit
volumes
Equipment,
(Warren
system
were
that
measured
Wilton,
E Collins,
was
VT)
Boston).
averaged,
V02
every
a gasometer
calibrated
Expired
and
percent
body fat.
consumption
(peak
workload
protocol
of the ventilatory
determined
with
calculated
with
were
weighing
was
4%.
the
VO2)
that
threshold
and
an automated
20 s. Inspiratory
(RAM-9200:
against
a Tissot
oxygen
and
Rayfield
spirometer
carbon
dioxide
lowers muscle
both training
ca-
pabilities
and maximal
exercise
performance
ability.
This thesis,
however.
was not supported
by Simonsen
et al (5) for rowers
who consumed
a moderate-carbohydrate
diet while undertaking
twice-daily
intense
was reduced
but
the
wt.d)
compliance,
or exceeded
are
consumption
wt1
consent
was paid
versity.
1993:57:27-31.
Endurance
to compare
body
kg body
Thirty-six
the
ygen
is
Subjects
study.
Human
has no apparent
benefit
exercise
performance.
exertion.
designed
bilities of runners
tains muscle
ing capability
Physical
was
training
informed
subject
WORDS
study
KEY
capabilities
Methods
C/in Nutr
performance
the high-carbohydrate
completed
glycogen.
but this
or high-intensity
exercise
glycogen
glycogen
1993 American
conSociety
I From
the Exercise Physiology
Laboratory,
School of Health, Physical
Education,
and Recreation,
The Ohio State University,
Columbus.
OH.
2 Supported
by the Gatorade
Sports Science Institute,
Chicago.
3 Address
Education,
reprint requests
and Recreation,
to WM Sherman.
School of Health. Physical
337 West 17th Avenue,
Columbus,
OH
August
43210.
Received
Accepted
for Clinical
Nutrition
12. 1992.
27
oxygen
and
(3).
28
SHERMAN
TABLE
Physical
ET
AL
sumed
and physiological
characteristics
Age
ofthe
subjects*
Body
fat
VO2max
L/inin
between
Raw
VO2max
body
wt
inL . kg
30 3
253
11 2
30
34
11
10
9 for each
treatment
4.2
91
0.2
59
61
60
4.30.2
583
4.2 0.2
4.5 0.1
Blood
SE. n
diet: MOD
CHO.
group.
moderate-carbohydrate
HI CHO.
Blood
measured
burgh,
and
with
LB-2:
calibrated
with
electric
analyzers
Beckman,
Fullerton,
a National
the concentrations
(S-3A/l
Bureau
ofwhich
CA).
Pitts-
analyzers
were
calibration
gas,
The
of Standards
were
Ametek,
confirmed
with
chemical
9 for each
and
design
diet
the effects
to assess
cle glycogen
and
dietary
to either
were
period,
ofdietary
to assess
alternating
legs
carbohydrate
content
over
both
exercise
and
intense
diet
tively. Runners
ran up a +6%
to favor concentric
contractions
incline
and
before
the start of the training
Velodyne
ergometers
(Schwinn,
this
were
40, 40,
respec-
control
Cyclists
exercised
IL) fitted with
phase,
subjects
During
provided
during
the control
period
with liquid
dietary
supplementation.
laboratory.
Subjects
exercised
period.
daily
followed
by 1-mm
remained
mL
250
cooled
During
oratory.
the
every
Heart
rate
period
consumed
During
the
or 830
training
mL (high)
environmental
Subjects
during
food
diets
meal
between
I 630
period,
subjects
ofthe
and
1 800
sprints
at
For runners
ratings
also
for
100%
the
then
samples
was
provided
containing
either
liquid
1 730.
consumed
supplement
Breakfast
conditions
consumed
and
in the
lab-
5 or
10 g
supplemented
(GatorLode).
660
with
lunch
1645
the
lateralis
or lateral
glycogen
head
in
were
from
without
proximal
the
ofa
liquid
glycogen
potential
subsequent
nitrogen
effects of
biopsy.
and
concentration,
suc-
(> 3 cm)
stored
frozen
at
muscle
into three
pieces
(5-10
mg) under
(to the nearest
0.01 mg at -20 #{176}C).
of the
synthase
utilization
obtained
in 2 mol
same
HCI/L
muscle
( 1 1 ) in
activity
analyzed
h at
100
sample
and
8.4%
The interassay
CV
per kg wet tissue.
the vastus
lateralis
in a subset
of four
on day 1 of the
was 2.2%.
enzymatically
available
kit
and interassay
(2
the glucose
concenenzymatically
with
of variation
(CV)
of the same
muscle.
results
were expressed
using a commercially
The duplicate
sample
at
7 of
vastus
lateralis
were
hydrolyzed
for duplicates
citrate
the
at progressively
for
(Boehringer,
CVs were
and
runners
training
glucose
by
Indianapolis).
1 .4% and 3.9%,
respectively.
TABLE 2
Characteristics
carbohydrate
for a typical
ofthe
(CHO)
subject
control,
moderate
(MOD). and high (HI)
diets during the control and training phases
Expe rimental
of per-
exercise.
carbohydrate
. kg
. d (Table
2). All diets were
with a beverage
containing
I 8% maltodextrins
erate)
2.2%
Blood
a
in the
and
and
muscle
20 mm
7-d training
Subjects
1600
muscle
biopsies
frozen
first divided
and weighed
was
on
to famil-
supervised
by five 1-mm
rest intervals.
of each sprint.
The
50% relative
humidity.
water
was
between
at +6%.
ceived
exertion
for the whole
after
15 and 45 mm during
before
cessation
were 25 #{176}C
and
exercise
from
the vastus
gastrocnemius
muscles
was analyzed
and four cyclists
in muscle
obtained
period.
The CV for duplicate
samples
on
their
consumed
sample
Muscle
on a motor-driven
treadmill
to minimize
muscle
damage
phase
(8).
Excelsior,
quick
for triplicate
samples
was 3. 1 %. Glycogen
diet containing
8 g carbohydrate
kg
d
(67% ofenergy
from
carbohydrates,
I 5% from protein,
and 18% from fat: Table
2).
An 1 8% carbohydrate
beverage
(GatorLode,
The Quaker
Oats
Co. Chicago)
was
iarize the subjects
Each
was
period,
obtained
obtained
1 . 3. 5, and
#{176}C)
and neutralized
with NaOH.
Finally.
tration
of the hydrolyzate
was determined
fluorometry
(10). The intraassay
coefficient
on mus-
Subjects
exercised
for 60,
VO2 for control
days 1-5,
During
in a
7 d of
supervised
in the laboratory.
20, and 20 mm at 75% peak
bicycles.
and
assigned
were
#{176}C.
To determine
samples
were
liquid
nitrogen
or high-carbohydrate
capabilities
training.
control
cyclists
randomly
a moderate-
performance
cycling
or running
During
the 5-d
control
group)
also
Muscle
Samples
on
designed
supplements
were
days
anal-
was
mL
(mod-
the evening
were
con-
Control/
control
Energy intake (J)
CHO energy (% oftotal)
CHO from typical foods
(gCHO/d)
(% total energy)
Liquid supplement
(gCHO/d)
(CHOas%totalenergy)
(mL/d)
14.7
67
phase
Training!
MOD CHO
14.7
42
and diet
Training!
HIGH CHO
14.7
84
539
61
234
565
27
65
46
6
230
132
15
660
166
19
830
exercise
(n
ofmineral
on
either
session.
from
study
the training
design
double-blind
treadmill
from
carbohydrate
the
biopsies
were
session
-80
a 5-d
dietary
that
session
training
tion
yses.
E.vperimnental
muscle
Biopsies
and
as snacks.
body weight
the potential
biopsies
exercise
period.
provided
workout,
minimize
about
informed
macc/c
and
the
of the gastrocnemius
and nine runners
were
own
samples
oxygen
the
After
and
before
training
consumption.
runners
analysis
daily
high-carbohydrate
were
were
the afternoon
100 g. To
knowledge
they
celery)
to determine
the effects oftwo
types
muscle
glycogen
and performance.
Runners
HI CHO
MOD CHO
and
before
nearest
of subject
performance.
Cyclists
HI CHO
MODCHO
(carrots
Before breakfast
and
was recorded
to the
influence
vegetables
DIETARY
To
measure
session
two
exercise
on day
maximal
haustion.
performance.
7. each
subject
two
after
rested
performance
The
the
for 5 mm
trials
performance
CARBOHYDRATE
at
80%
trials
normal
before
peak
were
ergometer
or to keep
pace
VO2
until
ex-
separate
the treadmill
of variance
measure
(time)
were used to assess
ments
on the dependent
variables
least-significant-difference
cant
pair-wise
175
the subject
revolutions
E
E
150
the effects
for cycling
procedure
differences
committing
a Type
are reported
as mean
was
among
I error
was
with
ofthe
and
dietary
running.
used
means.
Lu
125
0.05.
<
75
treatThe
signifi-
probability
DAY
of
Group
.f* #
_#
100
-J
C,
repeated
to locate
The
to P
held
(diet)
%%h1I
speed.
C-)
analyses
CYCLISTS-MOD
--.-.
200
C,
0
one-way
CYCLISTS-HI
a)
by 5 mm
Statistical imialisis
Two
29
GLYCOGEN
.--
training
separated
with
MUSCLE
undertaking
of rest. Exhaustion
was identified
as the time at which
was unable
to maintain
the prescribed
rate ofpedal
on the cycle
AND
data
SE.
Results
subjects
All
completed
d of the training
75% ofmaximal
the
prescribed
period.
On average.
heart rate and peak
daily
training
ofwhole-body
perceived
exertion
79% and
throughduring
the
on average.
responses
or leg perceived
for either
the
cyclists
(results
not shown).
Interestingly,
despite
during
the
or runners
similar
5-d control
period.
in cyclists
compared
with
in runners
for either
eralis
muscles.
I 12.6
8.9
trocnemius
For
exercise
muscle
consistent
104.0
6.4
and
vastus
lateralis
<
either
dietary
was
0.05).
1 . muscle
mmol/kg,
before
cyclists.
of runners
and after
muscle
and
exercise
glycogen
cyclists
lat-
and
73.7
between
4.2
the
mmol/kg.
pre-
and
postexercise
for the
significantly
reduced
was
<
0.05)
a significant
on days
difference
in muscle
second
Similarly,
perfor-
was no effect
for either
cy-
when
performance
second
trials were summed,
there was no
the cyclists
or runners.
Mean (SE)
total
for the
cyclists
on
the
moderate-
and
high-
(data
not
averaged
effects
ofthe
6 13 36 and
dietary
treatments
560
for
shown).
are
often
advised
to consume
a high-carbohydrate
to maintain
muscle
glycogen
(2). The results
of the present
concentrations
study
clearly
undertaking
diet
a constant
(5 g - kg body
-e--
RUNNERS-HI
--.-
RUNNERS-MOD
coefficients
glycogen
7 compared
and
For
the
concentra-
150
a)
runners.
diet, muscle
glycogen
7-d training
period
(Fig 1).
3, 5, and
not shown).
no significant
training
levels
demonstrate
daily training
mmol/kg,
106.7 3.9
10.9
were
diet,
were
glucose
Athletes
biopsies
moderate-carbohydrate
(data
times
diet during
at optimal
not change
significantly
over the
In contrast.
for the moderate-carbohydrate
significantly
difference
bohydrate
blood
synthase
period.
and after
correlation
muscle
0.8 1 (P
on the high-carbohydrate
significantly
over the
In contrast,
There
Spearman
or runners
times
effect
first
period.
There
performances
Discussion
muscles
for
28.2 3.2
muscle
The
averaged
citrate
training
sessions
were
186.7
respectively.
Corresponding
for the
Muscle
underwent
clists
There
higher
was
activity
was similar
(P > 0.05) in the vastus lateralis
the cyclists
and runners,
averaging
30.7 2.0 and
j.imol#{149}min
g, respectively.
A subset
recorded
erateand high-carbohydrate
106 5, respectively.
effect
respectively,
muscles.
diet
or vastus
glycogen
were
carbohydrate
diets
Total
performance
controls
65%
This
the gastrocnemius
on day
and
and
glycogen
(P
runners
runners
consuming
times
performance
exertion
for the daily training
sessions
(results
not shown).
There
were also no differences
in body weight
over the 7-d training
period
Exercise
mance
of diet
glycogen
with
glycogen
on days 5 and
diet, muscle
was
day
for
1.
the
7.
glycogen
7-d training
period
(Fig 2).
diet. glycogen
was
reduced
on days 5 and 7. There
was a significant
in muscle
glycogen
for the high- and moderate-cardiets on days 5 and 7.
E
z
I 25
100
Lu
C,
0
C.)
75
>-
-I
C,
50
0
DAY
on all 7
exercise
elicited
VO2, respectively,
30
SHERMAN
d
during
) significantly
days
5-7
containing
muscle
muscle
bohydrate
have
diet
training
load
decline
(40%)
body
similar
muscle
over
Costill
in muscle
wt
days
maintains
days
when
when
the
both
design.
runners
and
Subjects
carbohydrate(5
mm at 75%
self selected
consumed
and
and
Other
drate on
increased.
over
a diet
with
containing
a crossover
45%
of energy
there
as
daily for 60
1 3% over
were
no differences
in responses
examined
the
of dietary
between
cyclists.
muscle
Costill
glycogen
when
the training
load was
et al (1 3) doubled
daily swim training
subjects
consumed
had
runners
consuming
a self
double
either
5 of the study.
of the present
14) demonstrate
that
a diet
daily
even
doubled
and
by 1 .47-fold
duced
when
and
those
jects reported
the performance
training
muscle
sessions.
glycogen
euenergetic
dined
this time
Similarly,
carbohydrate
kg
containing
(4,
consumption
results
in lower
8 g carbohydrate
kg
petitive
endurance
events
during
the
by the
performance
final
experiment
stages
were
(3, 4, 12-14).
Importantly,
the present
study demonstrates
the
time course
of the effects
of consuming
different
amounts
of
dietary
carbohydrate
on muscle
glycogen
when
daily
training
of an endurance
hydrate
diet and
event.
resultant
did
maximal
load
high-carbohydrate
is constant.
tains
drate
Consuming
muscle
-
kg
5 through
glycogen,
.
whereas
significantly
day
Early
glycogen
10 g carbohydrate
consuming
reduces
kg
muscle
enhance
tions
in muscle
glycogen
do
Nevertheless,
high-carbohydrate
the relationship
between
low
and fatigue
during
endurance
muscle
perfor-
performance
of a healthy
ported
athletic
lifestyle
(16),
detrimental
performance,
and
mance
(1 , 1 5). This has led to the presumption
that reduced
muscle
glycogen
that resulted
from consuming
a moderateor
low-carbohydrate
diet over days or weeks
of training
would
de-
consume
a high-carbohydrate
The reasons
for the different
crease
capabil-
between
subjects,
period
ities.
training
compliance
As indicated
from
and
maximal
the results
performance
for a subset
of nine
training
on day 1 of the experimental
period
reduced
muscle
glycogen
by 1 1 7 and 33 mmol/kg
for the cyclists
and runners,
respectively.
Presumably,
this amount
of muscle
glycogen
or
less
was degraded
Interestingly,
concentrations
the subjects
day 7 after
on a daily
in spite
by days
completed
the daily
basis
during
of significantly
5 and
7 for
increased
cardiovascular
daily training
perception
strain
or the
both
training
muscle
cyclists
and
session.
glycogen
of effort
on
in
performance
tests between
diets. Subjects
neither
renor
exhibited
(eg, heart
rate) on day 7 during
maximal
performance
tests.
increased
either
are
same
diet
75%
ofthe
the
and
not
was
than
those
ofthe
there
muscles
status
muscle
glycogen
is possible
that
glycogen
for
the
ofthe
do not
two
been
no
on
to
period
the
intensity
the runners
However,
ran
was
uphill
the potential
in( I 7- 1 9). Trained
untrained
muscles
this
synthase
muscle.
It
were more
is not consistent
activities
in the
and cyclists;
thus. differences
to explain
the differences
between
conditions
modes
runners
did not
or
re-
experimental
control
exercise
VO2 . Also,
runners
appear
exercise
reduc-
concentrations
ofthe
the
relative
in citrate
concentration
the control
the
have
glycogen
During
runners.
similarities
the
to influence
diet-induced
peak
Perhaps
brief
the observed
quadriceps
in training
phases
a high-carboconcentrations
impair
athletic
performance.
diets are recommended
as part
diet.
muscle
and
mode-specific
present
these
carbohydrate
consumption
prudent
to advise
athletes
apparent.
consumed,
followed
too
has a higher
glycogen
content
than
be argued
that the cyclists
quadriceps
trained
with
cyclists
readily
to concentrically
fluence
of muscle
muscle
might
runners,
each
reduced
runners
bout
7 of the
performance.
was
dis-
day
or less simulate
because
effects ofhigh
it remains
fixed
training
short-term
not
present
exercise
these
conditions,
muscle
glycogen
or perhaps
the
intensity
on
regimen
athletic
day
The
endurance
dietary
carbohy-
7 of training.
studies
established
concentrations
not
to more
Under
higher
in the
to traverse
of higher
event.
from
glycogen
main-
used
athletes
undertaken
selected
reduced
or from
a period
of the
trials
required
ifthe
deficit
tasks
require
with
muscle
the
not appropriate
to detect
the
concentrations
on maximal
a close association
between
concentration
( 1 , 1 5). On
as fast as possible
when
intake
decreased
significantly
de-
energy
performance
the preexercise
glycogen
constant
power
exercise
tances
completing
of perceived
completed
the
12-
muscle
the
difficulty
ratings
(1 3) it is uncertain
the
ex-
influ-
was suddenly
energy
compliance
from
and other
studies
(4, 1 3, 14) were
effects ofdiffering
muscle
glycogen
performance.
Early studies
reported
when
training
Notably,
a diet
capabilities.
volume
consumption
inadequate
training
in this case
that
glycogen
on training
compliance
expenditure.
Although
sub-
subjects
capability
resulted
muscle
glycogen.
It is possible
kg
muscle
on training
training
oxygen
(14),
When
by 20%,
no
levels
declined
by 15% (1 3) or differed
still no systematic
effect of the re-
higher
reduced
(1 3). However,
training
reduced
distance
glycogen
in which
glycogen
there was
slightly
that
to increased
when
local muscular
fatigue
and
sessions
( 1 3) or increased
and
performance
reported
regard
ofexertion
muscle
glycogen
concentration
energy
intake
matched
energy
suddenly
volume
of others
over 3-10
d of training
or suddenly
increased.
muscle
glycogen
1 to day 3 of training.
On the other
not systematically
study
the effects
in muscle
( 14),
on maximal
with
days
muscle
or low-carbohydrate
et al (4)
observed
in studies
was
training
was
13% reduction
However,
glycogen
82 and
12 1 mmol/kg
for
diets, respectively,
before
5 g carbohydrate
than
their
study
dietary
ences
muscle
glycogen
volume
is either constant
containing
ofthe
and
Costill
successive
carbohy-
d . Muscle
glycogen
averaged
moderateand high-carbohydrate
ercise on day
The results
selected
kg . d.
During
I 30 to I 10 mmol/kg.
.
compliance
unusual.
pattern
with
of lowered
a moderate-
that
in
and cyclists.
It
equate
muscle
carbohydrate
is
8.2 g carbohydrate
declined
from
et al (14)
for 5 d while
effects
from
training
is not
exertion
have
10 d when
glycogen
study
studies
diet containing
muscle
glycogen
Kirwan
in a 3-d
cyclists
on daily
capabilities
offatigue
effect
resulting
decreased
by 40% from day
hand,
Pascoe
et al ( 1 2) did
a diet containing
43% of energy
from carbohydrate
over 3 d of
training.
The daily
training
was running
for 73 mm at 80%
VO2max
followed
by a VO2max
test. Similarly,
Pascoe
et al (12)
used
of a systematic
consistent
a progressive
subjects
lack
concentrations
diet
Two
of a moderate-carseveral
AL
The
diet
oftraining.
et al (4) observed
glycogen
30-36%
ofa
5-7
effects
glycogen
was constant.
by
consumption
during
reported
on
kg
concentrations
studies
glycogen
In contrast,
10 g carbohydrate
glycogen
other
reduces
oftraining.
ET
DIETARY
somehow
stored
Nevertheless,
cyclists
and
centration
etary
lack
differently
because
runners,
does
not
carbohydrate
ofeffect
between
comparisons
this
negate
intake
on training
cyclists
were
initially
the
CARBOHYDRATE
different
compliance
(20).
between
glycogen
of the effects
glycogen
and
runners
made
muscle
observations
on muscle
and
not
and
maximal
AND
conof di-
the resultant
performance
13
capabilities.
We acknowledge
the assistance
of Mike Wilson and Ashley
The cooperation
of the subjects in the study is appreciated.
Blostein.
References
9.
10.
1 1.
12.
13.
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JC, Sherman
WM, Lamb DR. Dernbach
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