Dietary protein, endurance exercise, and human skeletal-muscle
protein turnover
Nancy R. Rodriguez, Lisa M. Vislocky
and P. Courtney Gaine
Curr Opin Clin Nutr Metab Care 10:40–45. 2007 Lippincott Williams & Wilkins.
Purpose of review
Whereas diet and exercise have been shown to influence
whole-body protein utilization, little is known about the
impact of these factors on skeletal-muscle protein turnover.
We highlight the role of dietary protein in modulating
skeletal-muscle protein turnover in response to endurance
exercise. Effects of endurance exercise on skeletal-muscle
protein metabolism are presented and the influence of
habitual protein intake on exercise-related protein responses
is discussed.
Recent findings
Skeletal-muscle protein turnover increases in response to
endurance exercise training and following a single
endurance exercise bout. Nutritional supplementation
postexercise favorably affects skeletal-muscle protein
synthesis and demonstrates amino acid availability as
pivotal to the skeletal-muscle synthetic response following
exercise. The level of habitual protein intake influences
postexercise skeletal-muscle protein turnover.
Summary
Dietary protein and exercise are powerful stimuli affecting
protein turnover. Since variation in habitual protein intake
influences skeletal-muscle protein turnover postexercise,
investigations are needed to determine what role protein
intake has in regulating skeletal-muscle protein metabolism.
Long-term, well controlled diet and exercise intervention
studies are essential for clarification of the relation between
protein intake, endurance exercise, and skeletal-muscle
protein turnover. Studies designed to characterize this
relationship should be attentive to habitual macronutrient
and energy intakes.
Keywords
amino acids, exercise, phenylalanine, protein, skeletal
muscle
Department of Nutritional Sciences, University of Connecticut, Storrs, Connecticut,
USA
Correspondence to Nancy R. Rodriguez, PhD, Associate Professor, Nutritional
Sciences, Unit 4017, University of Connecticut, Storrs, CT 0626-4017, USA
Tel: +1 860 486 0120; fax: +1 860 486 3674;
Copyright © Lippincott Williams & Wilkins. Unauthorized reproduction of this article is
prohibited.
e-mail: nancy.rodriguez@uconn.edu
Current Opinion in Clinical Nutrition and Metabolic Care 2007 ,
10:40–45
Abbreviations
BCAA branched-chain amino acid
EAA essential amino acid
FBR fractional breakdown rate
FSR fractional synthetic rate
SMPTO skeletal-muscle protein turnover
2007 Lippincott Williams & Wilkins
1363-1950
Introduction
Varying the level of dietary protein alters whole-body
protein metabolism [1–3,4]. We have shown an increase in
the oxidation of total protein with increasing protein intake
that can likely be attributed to increased amino acid
availability [3]. However, the response rates for synthesis
and breakdown to variations in protein intake are less clear.
Discrepancies may be due to differences in the level of
dietary protein, as well as the magnitude of change in level
of protein intake implemented in study designs. We
previously reported that level of dietary protein intake
influences whole-body protein turnover at rest, with
increasing protein intake resulting in increased rates of
whole-body protein breakdown and oxidation and an
increase in nitrogen balance [3]. In this paper, we focus on
how changes in habitual protein intake influence skeletal-
muscle protein turnover ( SMPTO ) following an acute bout
of endurance exercise.
Exercise has a profound effect on protein metabolism. Of
particular interest is the influence that exercise has on
SMPTO, given that skeletal muscle comprises
approximately 40% of all body protein stores [5]. For the
purpose of this review, SMPTO refers to the synthesis (
fractional synthetic rate, FSR) and breakdown (fractional
breakdown rate, FBR) of skeletal-muscle proteins and the
difference between these processes which determines net
gain or loss of protein. The mode, intensity, and duration of
the exercise performed, along with the training state of the
individual, influences the magnitude and direction by which
exercise influences SMPTO. In contrast to resistance
training, the SMPTO response to endurance exercise has not
been fully characterized. Endurance exercise and
skeletal-muscle
protein metabolism
 Skeletal-muscle protein turnover Rodriguez et al. 41

The impact of an acute bout of aerobic exercise on skeletal-

40
skeletal muscle reacts during a single session of aerobic
exercise. However, due to the relatively short period of time
spent exercising compared with that spent not exercising over
the course of a day, the postexercise skeletal-muscle
responses provide valuable information regarding the overall
impact of endurance exercise on protein metabolism.
muscle metabolism in humans is not completely understood.
No published data demonstrate how
at rest after 4weeks of aerobic training. The increase noted in
FSR following endurance training may reflect an increased
synthesis of proteins responsible for bringing about the
adaptations associated with this mode of exercise. These
Figure 1 Skeletal-muscle protein turnover before and after
endurance training in healthy adults

To date, FSR following endurance exercise in humans has

% . h −1 0.2
been examined in only three studies [6–8]. Carraro et al. [6]
and Sheffield-Moore et al. [7] both found FSR increased in * Untrained
0.15 Trained
skeletal muscle following treadmill walking at 40% VO2max.
Although not significant, Tipton et al. [8], employing a high-
*
intensity swimming workout, observed a 41% increase in 0.1
FSR. Definitive conclusions cannot be drawn from just three
studies that differ greatly in design. However, collectively
0.05
these findings provide evidence of FSR increases following
acute endurance exercise. NET
0
FSR FBR
Even less is known about the influence of chronic aerobic
trainingonSMPTO.Asexercisegenerallylastsarelatively short −0.05

period of time compared to the time spent not exercising,

studying SMPTO at rest can provide insight −0.1 *
regardingtheinfluencethatchronicaerobictraininghason
skeletal-muscle protein metabolism. We recently Mixed-muscle fractional synthetic rate (FSR), fractional breakdown rate
conductedaninvestigation with previouslyuntrained,healthy (FBR), and net protein balance (NET) in eight healthy men and women
men and women in which rates of both protein synthesis who underwent 2 weeks of dietary intervention and 4 weeks of aerobic
exercise training and dietary intervention at baseline (untrained) and
(FSR) and breakdown (FBR) were measured before and after
after training (trained). Values are expressed as means standard error.
training to determine how aerobic exercise training
influenced SMPTO at rest [9]. To minimize the influence of Different from untrained, P < 0.05. Figure taken from Pikosky et al. [9 ].
diet (specifically variations in energy and protein intake), a
eucaloric diet intervention providing a constant level of
protein throughout the 6-week exercise training program was
employed. This study, which tested the hypothesis that a 4-
would include increases in mitochondrial volume,
week aerobic training program would result in an
mitochondrial enzyme activity, capillary/muscle-fiber
upregulation in SMPTO as indicated by increases in both
ratio, capillary density, the number of capillaries around a
FSR and FBR, was unique given the assessment of both FSR
given muscle fiber, as well as myofibrillar proteins [11–14].
and FBR in human skeletal muscle in response to aerobic
exercise training while dietary control was implemented.
Measurement of FBR in our study provides further insight
into the effects of aerobic exercise training on SMPTO
Resting SMPTO increased subsequent to 4weeks of aerobic
since net protein balance (FSRFBR) can be calculated and
exercise training, as indicated by an increase in FSR, a trend
SMPTO fully characterized. This had not been determined
toward an increase in FBR, and a decrease in net protein
previously at rest following an endurance training program.
balance (Fig. 1). A recent study by Short et al. [10] provides
Our results showed that in addition to the noted increases
the only other data regarding skeletalmuscle protein
in FSR, there was a tendency for FBR to increase,
metabolism following aerobic exercise training in human
ultimately resulting in a more negative net protein balance
volunteers. These investigators observed a 22% increase in
following 4weeks of aerobic exercising training. The
FSR at rest, following 4months of aerobic training in
potential physiological significance of the tendency for
previously untrained subjects. This is consistent with findings
FBR to increase at rest in response to aerobic training
from our study in which FSR increased approximately 17%
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Unauthorized reproduction of this article is
prohibited.
should not be overlooked, given the role of FBR in
providing amino acids for protein synthesis in the fasted
state [5,15].
Since rates of breakdown exceed those of synthesis in the
fasted state, a negative net protein balance is expected when
exogenous amino acids are not available [16]. However, the
observation that net protein balance is more negative
following endurance training is in contrast to what is seen
following resistance exercise, where net protein balance is
improved due to increases in synthesis that are of a greater
magnitude and longer duration than the increases in
breakdown [17].
42 Protein and amino acid metabolism
In addition to our SMPTO findings, we observed a decrease
in plasma branched-chain amino acid (BCAAs), essential
amino acid (EAAs) and nonessential amino acid
concentrations in response to 4weeks of aerobic exercise
training [9]. We do not believe this was due to an increase
in the oxidation of amino acids since leucine oxidation
significantly decreased and substrate-oxidation data
suggested a trend for a decrease in protein oxidation
following training [18]. Perhaps the observed decline in
plasma amino acids reflected insufficient protein intake to
support increases in protein turnover associated with the
onset of aerobic training. Indeed, we noted that whole-body
leucine turnover tended to increase and was positively
correlated with increases in resting energy expenditure
following training [18].
Our findings of a more negative net protein balance and a
simultaneous decrease in plasma amino acid concentrations
in response to training may suggest that the protein intake
consumed throughout the study (0.85gkg1 of body weight)
could have been suboptimal for our exercising subjects
despite adequate energy intakes. Although it has been
suggested that habitual endurance exercisers do not require
additional protein if energy intakes accommodate the
increased energy demands of exercise [19–21], a
substantial amount of research has demonstrated that these
individuals’ protein needs likely exceed the current
recommended dietary allowance for protein [22,23,24].
Perhaps if subjects in our investigation had habitually
consumed a greater level of dietary protein, free amino acid
pools would have expanded, thereby reducing the need for
amino acids from endogenous protein sources, and
subsequently attenuating protein breakdown.
Amino acid availability and skeletal-muscle protein
utilization
It has been postulated that muscle-protein synthesis and
breakdown work in a coordinated fashion according to
intracellular amino acid availability, particularly that of the
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EAAs. Therefore, when amino acids are available, muscle-
protein synthesis increases, and when they are deficient,
breakdown increases to supply free amino acids [15].
Recently it has been suggested that it is the
concentrationsofplasma ratherthan intracellular
aminoacidsthat are responsible forregulating protein
synthesis [25]. When muscle-protein synthesis is elevated,
intramuscular amino acid concentrations fall as amino acids
become incorporated into protein. In the event that
synthesis is increased at a rate that exceeds the capacity of
the cell to import sufficient amounts of all amino acids into
the intracellular pool, breakdown might be stimulated in
response to a fall in intracellular concentrations. Therefore,
it has been postulated that in certain situations breakdown
is regulated by the need to replenish intramuscular amino
acid pools [15]. In the fasted state, the ‘recycling’ of amino
acids between the intracellular and extracellular amino acid
pools can be a futile or even catabolic cycle, resulting in no
muscle gain or even muscle loss. Therefore, for muscle
gains to occur, it is critical that the plasma amino acid pool
delivers amino acids to the intracellular pool, so that amino
acids are available for synthesis and reliance on breakdown
is attenuated [26]. Since dietary protein is a major source of
amino acids to the plasma free amino acid pool, habitual
protein intake will likely impact the supply of amino acids
to skeletal muscle and ultimately affect SMPTO.
Dietary protein intake and skeletal-muscle protein
turnover
It is generally accepted that in the fasted state net muscle
protein balance will remain negative. However, the acute
provision of nutrients (primarily amino acids) can influence
SMPTO, such that muscle protein balance becomes positive.
These studies provide substantial evidence that when amino
acids (alone or with carbohydrates) are administered
immediately following exercise, a shift to a positive muscle
balance results [27–29]. A similar positive net protein
balance was reported when a protein-containing supplement
was consumed following endurance exercise [30]. Therefore,
provision of exogenous amino acids creates a more anabolic
environment by increasing muscle protein synthesis and
decreasing the reliance on amino acids from muscle protein
breakdown.
We conducted a study to determine whether postexercise
skeletal-muscle protein synthesis would be modulated in
response to chronic consumption of increased dietary protein
[22]. Theoretically, increased or high protein intakes could
expand the free amino acid pool, decreasing reliance on
breakdown to supply the amino acids required for skeletal-
muscle protein synthesis. Therefore, data were obtained
across a wide range of protein intakes reflective of the current
dietary reference intakes [31]. The highest level (high
protein) was based on protein intakes recommended in
contemporary diet plans ( i.e. the 40/30/30 macronutrient

composition for carbohydrate, fat and protein, respectively),
the middle level ( moderate protein) reflected current
recommendations for protein intake for endurance athletes
[32,33] and the recommended dietary allowance for this age
group constituted the low level (low protein). We
hypothesized that habitual consumption of dietary protein
beyond what is recommended for these young men would
improve the skeletal-muscle protein synthetic response
following an endurance exercise bout, ultimately improving
SMPTO. This investigation was the first to demonstrate that
variations in habitual protein intake can directly impact
skeletal-muscle protein synthesis in the fasted state following
endurance exercise [22]. Postexercise FSR was significantly
higher for both low-protein and
Figure 2 Post-exercise skeletal-muscle protein turnover for low-protein (LP),
moderate-protein (MP), high-protein (HP) diets
% . h −1 0.2
LP
MP
0.15 HP
0.1
* BCAA, branched chain amino acid; EAA, essential amino acid; NEAA,
0.05
0
FSR FBR NET
−0.05
−0.1
This figure depicts mixed-muscle fractional synthetic rate (FSR), fractional
breakdown rate (FBR), and net protein balance (NET) following a 75-min run
at 70% VO2peak for low-protein, moderate-protein, and highprotein diet
interventions. FSR was significantly greater for low protein (0.083% h1) and
moderate protein (0.078% h1) than for high protein (0.052% h1; P< 0.05).
Data taken from Bolster et al. [22].
moderate-protein diets in comparison with the highprotein
diet (Fig. 2).
Upon initial observation of the data, it appeared that
increasing protein intake was of no benefit to skeletalmuscle
FSR. However, when considering the SMPTO data (FSR,
FBR and net protein balance), some intriguing differences
began to emerge. Although the turnover data must be
considered preliminary due to the limited number of subjects
for which FBR, and hence net protein balance, could be
derived (n¼4), high protein also resulted in the lowest FBR
and least-negative net protein balance following exercise. We
believe that increased circulating levels of EAAs, as a result
of increasing protein intakes, contributed to changes in
protein utilization by attenuating breakdown postexercise.
Copyright © Lippincott Williams & Wilkins. Unauthorized reproduction of this article is prohibited.
Skeletal-muscle protein turnover Rodriguez et al. 43
The amino acid data for increasing levels of dietary protein
are shown in Table 1.
To date, no other investigations have examined the impact of
varying levels of dietary protein in humans on SMPTO
following a bout of endurance exercise. However, several
studies have examined the effects of habitual protein intake
on the whole-body protein metabolic response to endurance
exercise. In brief, no significant differences exist between
rates of whole-body synthesis and breakdown in the fasted
state when protein intakes ranged from 0.9 to 2.5gkg1day1,
before, during, or after exercise [1,2]. Higher rates of leucine
oxidation with higher protein intakes are the only con-
Table 1 Plasma amino acid concentrations in trained runners at rest for
low-protein (LP), moderate-protein (MP), high-protein (HP) diets
Plasma amino acid concentration (mmol/l)
LP MP HP
Leucine 112 7 125 3 135 19
Isoleucine 59 3 62 1 66 11
Valine 177 10 225 9 254 29
Total BCAAs 348 16 402 7 470 68
Total EAAs 694 35 752 30 876 106
Total NEAAs 1536 63 1247 63 1318 95
nonessential amino acid. Values are expressed as means standard error.
Different from low protein, P< 0.05.
sistent responses that have been demonstrated [1,2]. While
it does not appear that dietary protein intake can impact
rates of whole-body protein synthesis and breakdown in the
fasted state, either before or after endurance exercise, it is
important to note that wholebody assessments of protein
utilization do not necessarily reflect skeletal-muscle protein
metabolism [6,34,35]. Our work clearly documents that
variations in habitual protein intake can impact SMPTO,
and protein synthesis in particular, during recovery from
endurance exercise.
Whereas the observation that high protein resulted in
decreased FSR in the postexercise period was in contrast to
our original hypothesis, feeding higher levels of dietary
protein appeared to increase free amino acid availability, as
suggested by the plasma amino acid data. Significant
differences were noted only in BCAAs between low and
moderate protein. However, the observed increase in EAA
and BCAA concentrations may have physiological
significance. An increase in amino acid availability
subsequent to increased protein intake could influence
protein turnover in one of two ways: (1) by increasing the
protein-synthetic response postexercise, similar to what has
been documented with acute amino acid feedings following
resistance training [16,17,27], or (2) by limiting the body’s
reliance on endogenous sources of these free amino acids,
thereby limiting proteolysis during exercise [15].
It is plausible that the increase in baseline BCAAs and
EAAs on the high-protein diet might have attenuated
protein breakdown during exercise. The potentially less
catabolic environment during exercise might have reduced
the magnitude of the anabolic response needed
postexercise, and may explain our observed decrease in
FSR postexercise when subjects were habitually consuming
the high-protein diet. Unfortunately, the low number of
subjects for whom these calculations were completed
require that observations regarding FBR and net protein
balance measures be considered as preliminary. However,
these findings provide supportive evidence that a stepwise
decrease in FBR postexercise may exist with
44 Protein and amino acid metabolism
increasing levels of dietary protein (0.068, 0.134, and
0.169%h1 for high, moderate and low protein, respectively).
Without question, further studies are needed to determine
whether habitual consumption of different levels of (i.e.
increased) dietary protein intake can modulate endogenous
proteolysis during exercise and recovery.
Conclusion
Aerobic exercise training upregulates SMPTO. The
observation that a negative net protein balance occurred
when energy balance was maintained in the trained
individuals was possibly due to training, an inadequate
level of protein intake, or a combination of both. Variations
in habitual protein intake influenced postexercise SMPTO,
demonstrating a potential role for dietary protein in
regulating skeletal-muscle protein metabolism. Overall,
findings suggest that rates of protein synthesis are, in part,
dependent upon the availability of free amino acids.
Aminoacids are procuredfrom the freeamino acidpoolfor
protein synthesis. In turn, amino acids are needed to
replenish this pool. In the fasted state, these amino acids are
derived solely from endogenous protein breakdown.
Therefore, the increase in protein breakdown that results
from endurance exercise can be viewed as necessary to
provide the free amino acids required to elicit the
concurrent increase in synthesis. The habitual provision of
exogenous amino acids from dietary protein may augment
this free amino acid pool, thereby facilitating protein
synthesisanddecreasingtheneedforaconcurrentincrease in
breakdown. Future initiatives should focus on the
interactions between level of dietary protein intake and
endurance exercise in the regulation of SMPTO to better
understand the contribution of habitual protein (i.e. amino
acid) intake to metabolic regulation.
References and recommended reading
Papers of particular interest, published within the annual period of review, have
been highlighted as: of special interest
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