European Journal of Sport Science

ISSN: 1746-1391 (Print) 1536-7290 (Online) Journal homepage: https://www.tandfonline.com/loi/tejs20

Considerations for protein intake in managing

weight loss in athletes

Caoileann H. Murphy, Amy J. Hector & Stuart M. Phillips

To cite this article: Caoileann H. Murphy, Amy J. Hector & Stuart M. Phillips (2015) Considerations
for protein intake in managing weight loss in athletes, European Journal of Sport Science, 15:1,
21-28, DOI: 10.1080/17461391.2014.936325

To link to this article: https://doi.org/10.1080/17461391.2014.936325

Published online: 11 Jul 2014.

Submit your article to this journal

Article views: 2346

View Crossmark data

Citing articles: 13 View citing articles

Full Terms & Conditions of access and use can be found at

https://www.tandfonline.com/action/journalInformation?journalCode=tejs20
European Journal of Sport Science, 2015
Vol. 15, No. 1, 21–28, http://dx.doi.org/10.1080/17461391.2014.936325

REVIEW ARTICLE

Considerations for protein intake in managing weight loss in athletes

CAOILEANN H. MURPHY1,2, AMY J. HECTOR1,2, & STUART M. PHILLIPS1,2

1 2
Department of Kinesiology, McMaster University, Hamilton, ON, Canada; Exercise Metabolism Research Group,
McMaster University, Hamilton, ON, Canada

Abstract
A large body of evidence now shows that higher protein intakes (2–3 times the protein Recommended Dietary Allowance
(RDA) of 0.8 g/kg/d) during periods of energy restriction can enhance fat-free mass (FFM) preservation, particularly when
combined with exercise. The mechanisms underpinning the FFM-sparing effect of higher protein diets remain to be fully
elucidated but may relate to the maintenance of the anabolic sensitivity of skeletal muscle to protein ingestion. From a
practical point of view, athletes aiming to reduce fat mass and preserve FFM should be advised to consume protein intakes
in the range of ∼1.8–2.7 g kg−1 d−1 (or ∼2.3–3.1 g kg−1 FFM) in combination with a moderate energy deficit (−500 kcal)
and the performance of some form of resistance exercise. The target level of protein intake within this recommended range
requires consideration of a number of case-specific factors including the athlete’s body composition, habitual protein intake
and broader nutrition goals. Athletes should focus on consuming high-quality protein sources, aiming to consume protein
feedings evenly spaced throughout the day. Post-exercise consumption of 0.25–0.3 g protein meal−1 from protein sources
with high leucine content and rapid digestion kinetics (i.e. whey protein) is recommended to optimise exercise-induced
muscle protein synthesis. When protein is consumed as part of a mixed macronutrient meal and/or before bed slightly higher
protein doses may be optimal.

Keywords: Protein, weight loss, body composition, nutrition

Introduction levels are often considered to enhance appearance

Weight loss is not an uncommon goal in athletes and
is frequently motivated by factors relating to per-
formance issues germane to the sport in which the
athlete is engaged. More often than not, this involves
loss of weight to enhance performance (i.e. improv-
ing the ratio of body weight to strength, power or
endurance) or for aesthetic reasons. Weight loss can
be achieved by restricting energy intake, increasing
the volume/intensity of training or, most frequently,
a combination of both of these strategies. For many
athletes, the desire for “weight loss” translates to the
desire for fat loss, rather than the loss of lean tissue,
which is due to the recognised role that skeletal
muscle plays in sports performance. On the other
hand, fat loss in some athletes may result in physio-
logical benefits such as more effective thermoregula-
tion, reduced energy cost or a greater power-to-mass
ratio (O’Connor, Olds, & Maughan, 2007). In
aesthetically oriented sports, such as bodybuilding,
diving, gymnastics and figure skating, low body fat
and consequently confer advantage in competition
(Sundgot-Borgen & Garthe, 2011). In many cases,
optimising the fat-to-muscle ratio of lost weight
during caloric imbalance is important, and this
has been referred to as “high -quality” weight loss
(Churchward-Venne, Murphy, Longland, & Phillips,
2013; Josse, Atkinson, Tarnopolsky, & Phillips,
2011; Phillips & Zemel, 2011). A growing body
of evidence now shows that higher protein intakes
during periods of energy restriction (ER) can
enhance the retention of fat-free mass (FFM) (Helms,
Zinn, Rowlands, & Brown, 2013; Krieger, Sitren,
Daniels, & Langkamp-Henken, 2006; Wycherley,
Moran, Clifton, Noakes, & Brinkworth, 2012),
particularly when combined with exercise (Mettler,
Mitchell, & Tipton, 2010), although studies specif-
ically examining elite athletes are limited.
The aims of this review are to examine some of the
mechanisms underpinning the influence of higher
protein intake on FFM preservation during ER.

Correspondence: Stuart M. Phillips, Department of Kinesiology, McMaster University, 1280 Main St. West, Hamilton, ON L8S 4K1,
Canada. E-mail: phillis@mcmaster.ca

© 2014 European College of Sport Science

22 C. H. Murphy et al.
We also aim to explore the associated practical issues
relevant to the athlete undergoing weight loss.
Skeletal muscle protein turnover: regulation of
muscle protein synthesis and muscle protein
breakdown
The maintenance of skeletal muscle mass is achieved
with a net neutral balance between muscle protein
synthesis (MPS) and muscle protein breakdown
(MPB) (Rennie, Wackerhage, Spangenburg, & Booth,
2004). In the postprandial state, MPS is stimulated and
exceeds MPB, resulting in a net positive protein
balance; however, in the post-absorptive state, the rate
of MPB exceeds the rate of MPS and results in a net
negative protein balance. It is well established that
MPS can be stimulated by protein ingestion and
resistance exercise independently and synergistically
(Churchward-Venne, Burd, & Phillips, 2012). A
key protein in the regulation of MPS is the
mammalian (mechanistic) target of rapamycin com-
plex-1 (mTORC1). Upon provision of amino acids,
especially leucine, the increase in the downstream
phosphorylation of targets of mTORC1 [4E Binding
Protein-1 (4EBP-1) and S6K1] ultimately leads to
increased protein translation efficiency (Wullschleger,
Loewith, & Hall, 2006). For further details on this
pathway and its regulation, the reader is directed to
several reviews (Drummond, Dreyer, Fry, Glynn, &
Rasmussen, 2009; Kimball, 2014).
Consequences of ER on changes in FFM
MPS is an energetically expensive process (Browne &
Proud, 2002). During ER, the availability of energy
from dietary sources is limited, and therefore, the cell
is forced to prioritise energy demands. Therefore, it is
not unrealistic to expect that ER would result in a
reduction in MPS. Pasiakos et al. (2010) demon-
strated a 19% reduction in MPS as well as an
alteration in the phosphorylation of 4EBP-1 in the
post-absorptive state during a 10-day 20% energy
deficit in healthy individuals, despite daily protein
intake being almost twice the Recommended Dietary
Allowance (RDA). The ER-induced reduction in
MPS may suggest an adaptive mechanism to conserve
energy. Consequently, if the rate of MPS is blunted,
the balance between MPS and MPB may shift from a
state of balanced protein turnover to negative protein
balance, resulting in the loss of muscle mass. In fact,
meta-analysis revealed that ∼25% of the mass lost
during weight loss in healthy individuals undergoing
ER is FFM (Weinheimer, Sands, & Campbell, 2010).
Few weight loss studies have been conducted in
elite athletes (Garthe, Raastad, Refsnes, Koivisto, &
Sundgot-Borgen, 2011). During weight loss in nor-
mal weight individuals, it has been shown that FFM
loss during the early ER phase can be more rapid
and severe compared to the overweight/obese popu-
lation, depending on the level of ER (Heymsfield
et al., 2011). One explanation for the differences in
weight loss patterns between overweight/obese and
normal weight individuals appears to be the amount
of stored body fat. For example, Elia, Stubbs, and
Henry (1999) explained that subjects with low body
fat during total starvation excrete twice the urinary
nitrogen and use more energy from protein sources
than those with higher body fat. Considering that
elite athletes typically have lower body fat and
participate in short-term weight loss plans, this
population may be more vulnerable to FFM loss,
and further research in this area is required.
High protein diets and FFM preservation in
ER: potential mechanisms
Understanding the mechanisms affecting muscle
protein turnover during weight loss would allow the
design of more effective weight loss programmes for
maintaining FFM. The current literature on protein
intakes and weight loss in athletes is sparse and uses
fairly extreme dietary comparisons. For example,
Pasiakos et al. (2013) compared protein levels at the
RDA (0.8 g kg−1 d−1), or two and three times higher
than the RDA on FFM and MPS over a 21-day
period at 40% energy deficit in healthy and over-
weight volunteers. It was observed that subjects
ingesting two or three times the RDA for protein
retained significantly more FFM and lost more fat
mass during ER. The authors also demonstrated that
the sensitivity or the anabolic response to protein is
only maintained during ER when protein levels
higher than the RDA are consumed. For example,
after consumption of a protein-containing mixed
meal, the MPS response was maintained during ER
only in those consuming two and three times the
RDA, but was inhibited during ER for those con-
suming the RDA. This diminished MPS response
has also been demonstrated in states of disuse
atrophy, where interestingly, the possibility of accel-
erated MPB contributing to FFM loss has been ruled
out (Phillips, Glover, & Rennie, 2009). Although
there were no differences in the MPS responses and
FFM retention between the two and three times
RDA protein groups with the healthy/overweight
population, considering the differences in body
composition and physical activity in elite athletes,
protein at an intake of close to three times RDA
protein may prove to be beneficial, and more precise
dose–response studies are necessary to determine the
optimal dose of protein, which may be somewhere in
the middle of these extreme dietary comparisons. In
addition, there are other factors such as protein

timing and quality (discussed later in this review)
which can optimise the effects of protein quantity.
There are conflicting reports on the effects of ER
on post-absorptive rates of MPS. While a 10-day
(Pasiakos et al., 2010) and a 5-day (Areta et al.,
2014) ER period showed decreased post-aborptive
rates in MPS, after a 21-day period, there were no
differences in the post-absorptive rates of MPS,
regardless of dietary protein level (Pasiakos et al.,
2013). Villareal, Smith, Shah, and Mittendorfer
(2012) has also demonstrated a maintenance of
post-absorptive MPS rate in subjects after 3 months
on a weight loss diet, suggesting that the MPS
response may adapt to more prolonged weight loss.
Heymsfield et al. (2011) provide support for this
theory by suggesting that weight loss should be
viewed in phases instead of as a continuous process,
whereby the early phase, lasting from days to a few
weeks is characterised by rapid weight loss and
is likely the phase where lean mass loss is most
pronounced, while a plateau is reached as weight loss
progresses. Such a model is suggestive of early phase
adaptations and a new “steady-state” as weight loss
progresses. Interestingly, the reduced post-absorptive
rate of MPS following short-term ER is rescued by
resistance exercise and protein ingestion (Areta et al.,
2014), and these strategies could preserve FFM
during short-term ER.
Increasing dietary protein intake during ER results
in an increased expression of human vacuolar pro-
tein sorting-34 (Vps34) mRNA, which is involved in
amino acid sensing and stimulation of mTORC1
(Pasiakos et al., 2013). The increased markers of
MPB in response to ER are reduced by feeding. For
example, after consumption of a protein-containing
mixed meal (480 kcal, 20 g protein, 8 g fat and 82 g
carbohydrate), protein ubiquitylation and 26S pro-
teasome activity were reduced during ER, regardless
of whether the protein content of the diet was 0.8,
1.6 or 2.4 g kg−1 d−1 (Carbone et al., 2013).
However, it may not be the level of protein affecting
postprandial Ubiquitin Proteasome Pathway (UPP)
regulation, but rather the level of insulin, which
would have been constant, following the consump-
tion of a mixed meal containing 82 g of carbohyd-
rate. Greenhaff et al. (2008) demonstrated that
increasing insulin to 30 mU/L (equivalent to post-
prandial levels) halved MPB without further inhibi-
tion at higher insulin doses. Additionally, there were
no further increases in the MPS rate above 5 mU/L
insulin. At the same time, the authors observed a
decrease in the expression of ubiquitin ligase MAFbx
and the C2 proteasome subunits with insulin levels
similar to what is seen in the postprandial period.
Increased amino acid availability augmented MPS
beyond 5 mU/l insulin with no change in MPB.
Considerations for protein intake in weight loss 23
The mechanistic advantage of high-protein diets
to spare FFM during ER can be explained by
considering the roles protein plays in stimulating
MPS by maintaining muscle’s anabolic sensitivity as
evidenced by both the increased Vps34 (amino acid
sensing) expression and retained rates of postpran-
dial MPS during ER with protein levels two and
three times the RDA. Furthermore, it appears that
fluctuations in MPS are the main determinants of
FFM, as rates of MPS fluctuate 10 to 20-fold more
than rates of MPB between post-absorptive and
postprandial states (Tang & Phillips, 2009).
Practical aspects for athletes: how much
protein?
The optimal intake of protein for athletes is difficult
to define, however, general guidelines state that,
under conditions of energy balance, protein recom-
mendations for endurance and strength trained
athletes range from 1.2 to 1.7 g kg−1 d−1 (Rodriguez,
DiMarco, & Langley, 2009). What is less clear is the
optimal protein intake for athletes during periods of
ER, although a number of researchers have high-
lighted that these requirements likely increase while
athletes are consuming hypocaloric diets (Helms
et al., 2013; Mettler et al., 2010; Phillips & Van
Loon, 2011). In a recent review, Helms and collea-
gues (2013) evaluated the effects of dietary protein
intake on body composition in energy-restricted,
resistance-trained, lean athletes. However, highlight-
ing the lack of available research in this area, of the six
studies included in the review only two actually
compared well-matched groups of athletes consum-
ing different protein intakes (Mettler et al., 2010;
Walberg et al., 1988). Mettler and colleagues (2010)
reported that consuming protein at the level of
2.3 g kg−1 d−1 (∼2.7 g kg−1 FFM) during a two-
week period of marked ER (40% reduction in energy
intake) resulted in a relative preservation of FFM
and the loss of ∼1.2 kg fat mass in resistance-trained
athletes. In contrast, a control group consuming
protein intakes of 1 g kg−1 d−1 (∼1.2 g kg−1 FFM)
not only lost similar body fat but also experi-
enced a significant reduction in FFM. In amateur
body builders, Walberg and colleagues (1988) showed
that while protein intakes of ∼0.8 g kg−1 d−1 (0.9 g kg−1
FFM) resulted in negative nitrogen balance during
1 week of severe ER (∼50% reduction in energy
intake), positive nitrogen balance was achieved when
higher protein intakes of ∼1.6 g kg−1 d−1 (1.9 g kg−1
FFM) were consumed. Supporting the notion that
FFM loss can be attenuated by higher dietary protein
intake, the mean FFM loss in this study was 2.7 kg and
1.4 kg, in the lower protein and higher protein groups,
respectively. Although the difference between groups
did not reach statistical significance the authors noted
24 C. H. Murphy et al.
that this may have related to the large variability in
body composition between participants.
The changes in body composition experienced by
athletes undergoing ER may be affected by a number
of other factors including the relative energy deficit
during ER (Garthe et al., 2011) and initial body fat
levels (Elia et al., 1999). Also relevant is the mode(s)
of exercise training undertaken by the athlete. It is
likely that whole body resistance exercise, as a
fundamentally anabolic stimulus, is the most effect-
ive in promoting FFM retention. Thus it is possible
that these factors may interact with protein intake
and influence ideal protein recommendations in a
case-specific manner. Indeed, it has been postulated
that protein intake recommendations should be
higher for athletes attempting to lose body fat who
are leaner and/or are undergoing a severe energy
deficit (Helms et al., 2013); however, we propose
that there is currently no conclusive evidence to
support this hypothesis and studies are required
to examine optimal protein intakes during caloric
restriction in high-level athletes under a range of
different conditions (i.e. severe vs. moderate energy
deficit, higher vs. lower initial body fat and resistance
vs. endurance training).
In the absence of sufficient data necessary to make a
truly evidence-based recommendation, protein
intakes of ∼1.8 – 2.7 g kg−1 d−1 (or ∼2.3 – 3.1 g kg−1
FFM) have been proposed, although intakes towards
the higher end of this range are largely speculative
(Helms et al., 2013; Phillips & Van Loon, 2011).
Nevertheless, from a practical point of view, intakes
towards the higher end of the recommended range
may be appropriate in the case of athletes consuming
large preweight loss protein intakes. The body adapts
to relatively high protein loads by increasing the
capacity for amino acid (or at least leucine) catabolism
(Millward, 2001). Thus, it is likely that the habitual
consumption of a high-protein diet means the athlete
is “forced” to continue consuming greater protein
intakes so that fed-state gains can balance fasted-state
losses. Considering that very high-protein intakes in
the range of ∼2.4–3.2 g kg−1 d−1 are frequently
reported in some groups of athletes (Phillips, 2004),
avoiding a sudden reduction in these levels, especially
when undertaking a phase of ER, would likely be
advisable to attenuate FFM losses. Equally, the “more
is better approach” can become problematic as unne-
cessarily high-protein intakes could potentially inter-
fere with their ability to achieve adequate intakes of
other macro-nutrients within a tight energy budget,
possibly resulting in impaired training adaptations
and/or limiting the ability to train effectively.
Few studies have been conducted to examine
whether athletes are self-selecting protein intakes
within the recommended range during periods of
ER. Umeda et al. (2004) reported that over a 20-day
period of weight loss prior to competition male
judoists progressively decreased their protein intake,
along with caloric intake, such that protein intakes
declined from ∼1.5 g kg−1 d−1 at baseline to ∼1 g
kg−1 d−1 four days before competition. In another
study, body builders gradually decreased their con-
sumption of all three macro-nutrients over 11 weeks
prior to competition (Maestu, Eliakim, Jurimae,
Valter, & Jurimae, 2010); however, the magnitude
of the reduction in protein intake was less (from 2.7
to 2.5 g kg−1 d−1) compared to the findings of
Umeda and colleagues. The relative maintenance of
protein intake reported by Maestu et al. (2010) was
likely facilitated by the longer time frame allocated
for body mass reduction (11 weeks vs. 20 days) and
consequently necessitated a reduction in energy
intake which was considerably less severe.
Clearly, if a higher protein intake (1.8 – 2.3 g kg−1 d−1)
is recommended to preserve lean mass during ER,
this will necessitate a reduction in at least one other
macro-nutrient. Whether it is more appropriate for
athletes to reduce fat and/or carbohydrate intake is
currently hotly debated. It has been reported that low-
carbohydrate diets (≤ 35–40% energy from carbohyd-
rate) enhance fat and body mass loss, although they were
also associated with greater FFM loss unless protein was
higher (Krieger et al., 2006). The vast majority of the
available literature on carbohydrate restriction and fat
loss comes from studies conducted in overweight and
obese populations. Consequently, concern has been
raised regarding the influence of low-carbohydrate diets
on sports performance and training ability in athletes.
Paoli et al. (2012) recently reported no decrements in
explosive strength performance in elite male gymnasts
after 30 days of a high-protein, low-carbohydrate
(∼22.0 g/d) hypocaloric diet combined with normal
training. On the other hand, Walberg et al. (1988)
reported a reduction in isometric quadriceps endur-
ance in bodybuilders consuming lower carbohydrate
intakes of ∼2.3 g kg−1 d−1 during 1 week of ER,
whereas a group consuming slightly higher carbohyd-
rate intakes of ∼3.2 g kg−1 d−1 did not experience a
decrement. In endurance athletes, a number of stud-
ies have reported no impairment in endurance-based
performance tasks, following short-term (1–2 weeks)
high-fat, low-carbohydrate (∼2.5 g kg−1 d−1) diets
designed to achieve energy balance (Yeo, Carey,
Burke, Spriet, & Hawley, 2011), although decrements
in high-intensity sprint performance have been
demonstrated (Havemann et al., 2006). Importantly,
the responses of athletes to low-carbohydrate intakes
appear to be highly variable and can impair training
capacity in some athletes (Yeo et al., 2008). Thus, it
appears reasonable to suggest that the decision to
reduce carbohydrate and/or fat intake to facilitate
increased protein consumption should be made on a
case-by-case basis with consideration of the athlete’s

sport-specific training needs and personal tolerance,
along with the amount of time available to achieve
weight loss.
Protein quality
Another key factor to consider when providing
advice to an athlete desiring to achieve high-quality
weight loss is the quality of the protein in the diet.
Until recently, the Food and Agricultural Organisa-
tion (FAO)/World Health Organisation (WHO)
recommended the use of the Protein Digestibility
Corrected Amino Acid Score (PDCAAS) method to
evaluate protein quality. The PDCAAS of a food
protein is determined from the limiting amino acid
score (i.e. the content of the first-limiting essential
amino acid – EAA – of the target food protein as a
percentage of the content of the same amino acid in
a reference pattern of EAA) corrected for the
digestibility of the target food protein. Although
this method was widely accepted, it has been
criticised in the scientific community for a variety
of reasons (Schaafsma, 2000). A controversial con-
cern related to use of PDCAAS was its truncation at
1.0 even if proteins had values exceeding this score.
It was argued that truncation removes any nutri-
tional differences between higher-quality protein
foods such as milk protein and soy since both have
a PDCAAS score of 1.0 even though they have true
PDCAAS scores of 1.21 and 1.04, respectively
(Phillips, Tang, & Moore, 2009). In order to address
some of these shortcomings, a report published by
the FAO Expert Consultation in 2013 has recom-
mended that the Digestible Indispensable Amino
Acid Score (DIAAS) should replace the PDCAAS as
the preferred method of measuring protein quality.
According to the DIAAS method values above 100%
are not truncated for individual foods consumed as
part of a mixed diet and amino acid digestibility
is determined at the terminal ileum, which more
accurately reflects the extent of amino acid absorp-
tion. Importantly, and in contrast to the PDCAAS
method, dietary amino acids are treated as individual
nutrients and digestibility correction factors specific
to each individual amino acid are used (where
available) rather than a single value for crude protein
digestibility (Schaafsma, 2000). While the FAO
Expert Consultation has highlighted that further
research is needed to ensure full implementation
and adoption of DIAAS, this method has demon-
strated greater variation in protein quality between
plant and animal protein sources. For example,
while milk and soy protein share the same PDCAAS
value, they differ in DIAAS value: milk at 122 and
soy at 94.
A role for protein quality (encompassing amino
acid composition, digestibility and availability) in
Considerations for protein intake in weight loss 25
muscle protein turnover becomes apparent when we
consider that an increase in plasma amino acid
concentrations is a potent stimulator of MPS and
this effect is primarily attributable to the EAA (Tang &
Phillips, 2009). It follows that emphasis on higher-
quality protein sources may be advantageous in the
preservation of muscle mass under conditions of ER.
Protein distribution
A rationale for considering protein distribution
throughout the day comes from acute studies show-
ing that ∼8–10 g of EAA is sufficient to induce a
maximal stimulation of MPS both at rest and after
resistance exercise (Cuthbertson et al., 2005; Moore
et al., 2009). From a practical standpoint, 8–10 g of
EAA translates into approximately 20–25 g of high-
quality protein, or ∼0.25–0.3 g kg−1 d−1. The
protein feeding-induced enhancement of MPS
returns to baseline ∼2–3 hours after amino acid
provision, even in the presence of continuing amino
acid availability (Atherton et al., 2010). Thus, it has
been suggested that athletes aiming to preserve lean
mass during ER may benefit from consuming 4–5
evenly spaced feedings containing 20–25 g of high-
quality protein throughout the day. Providing some
support for this hypothesis, it was observed that both
whole body (Moore et al., 2012) and muscle (Areta
et al., 2013) protein synthesis tended to be greater
when 80 g of whey protein was consumed as 4 g ×
20 g doses every 3 h compared to the same amount
consumed as either two large (40 g) doses every 6 h,
or eight small (10 g) doses every 1.5 h over the 12 h
recovery period following a session of resistance
exercise. However, there are a number of important
points to consider when applying this concept to the
energy-restricting athlete. First, the dose–response
studies conducted to determine the quantity of
protein required to maximally stimulate MPS were
conducted in weight-stable participants consuming a
diet designed to ensure energy balance in the days
leading up to the trial. Consequently, how this value
may be altered in athletes undergoing weight loss
and prolonged ER remains unknown. Second, pro-
tein or free amino acids were ingested in isolation
in these studies, typically in liquid form. In reality,
protein is often in a solid (more slowly digested)
form and is often co-ingested with carbohydrate
and fat during meals, which would generally slow
amino acid absorption and subsequently the profile
of aminoacidemia (Burke et al., 2012). Studies have
shown that the achievement of a rapid and pro-
nounced increase in plasma EAA concentration,
particularly leucine, is associated with increased rates
of MPS compared to a slow rate of appearance of
these amino acids (West et al., 2011). Therefore, it
would appear that when protein is consumed as part
26 C. H. Murphy et al.
of mixed meal quantities greater than 20 g per meal
may be necessary to achieve the minimal threshold
for blood leucine concentration required to maxim-
ally stimulate MPS. Thus, in the post-exercise
period, athletes aiming to preserve FFM would likely
benefit from the consumption ∼20 g of protein from
source with high leucine content and rapid digestion
kinetics in order to optimise the exercise-induced
MPS response. A recent study characterising the
plasma amino acid responses to the ingestion of 20 g
of protein from a range of commonly consumed
protein-rich foods reported that skimmed milk pro-
duced a significantly higher and faster peak leucine
concentration than all other foods/liquids (soy bev-
erage, beef steak, egg and liquid meal supplement;
Burke et al., 2012). Peak values for EAAs and
leucine after the consumption of a soy beverage
tended to be lower than for the other protein
sources, perhaps reflecting the lower leucine content
of soy. In addition, ingestion of liquid forms of
protein achieved peak plasma amino acid concentra-
tions twice as quickly as solid protein-rich foods,
suggesting that post-exercise consumption of liquid
protein sources may be preferable, although more
research is needed to determine the physiological
relevance of food form in the regulation of MPS.
Unfortunately, there are few data available on the
meal-to-meal protein intakes of energy-restricting
athletes. Erdman, Tunnicliffe, Lun, and Reimer
(2013) recently assessed the distribution of macro-
nutrient intake of 324 Canadian athletes competing
at national or international levels. Although the
athletes in this study were consuming total protein
intakes (∼1.9 g kg−1 d−1) exceeding those typically
recommended for athletes in energy balance, meals/
snacks containing at least 20 g of protein were
only consumed three times per day on average. A
potentially important and perhaps underappreciated
opportunity to consume a protein-rich meal is before
bed. It has recently been demonstrated that the
ingestion of a larger protein dose (40 g of casein)
immediately before bed resulted in a greater stimu-
lation of MPS overnight (Res et al., 2012), presum-
ably offsetting the FFM losses that would be
expected to occur during the 8–10-hour fast that
typically accompanies sleep. Such a pre-sleep meal
would likely also facilitate recovery as it would allow
repair/remodelling of proteins during the overnight
period. Considering the prolonged duration of the
overnight fast, larger protein doses such as those
used in the study by Res and colleagues may
be optimal in the pre-bed meal. In contrast to these
recommendations, the athletes studied by Erdman
et al. (2013) reported protein intakes of ∼6–8 g in
the evening snack. While the latter study did not
specifically investigate athletes undergoing ER, these
findings indicate that athletes may have suboptimal
practices when it comes to the pattern of daily
protein consumption.
Summary
Athletes aiming to achieve high-quality weight loss
should be advised to consume higher protein intakes
in the range of ∼1.8–2.7 g kg−1 d−1 (or ∼2.3–3.1 g
kg−1 FFM) in combination with a moderate energy
deficit (−500 kcal) and the performance of resistance
exercise. The target level of protein intake within this
recommended range requires consideration of a
number of case-specific factors including the athlete’s
body composition, training regimen, habitual protein
intake, personal tolerance and broader nutrition
goals. A reduction in dietary fat (to ∼20% of total
energy intake) and carbohydrate may allow athletes to
achieve higher protein intakes without the excessive
restriction of any one macro-nutrient which may
adversely affect health and/or performance. Athletes
should be advised to focus on consuming higher-
quality protein sources, aiming to consume 4–5
protein feedings evenly spaced throughout the day.
Post-exercise consumption ∼0.25–0.3g/kg of protein
from source with high leucine content and rapid
digestion kinetics (i.e. whey protein or skimmed
milk) is recommended to optimise the exercise-
induced MPS response. When protein is consumed
as part of a mixed macro-nutrient meal and/or before
bed slightly higher protein doses may be preferable.
References
Areta, J. L., Burke, L. M., Camera, D. M., West, D. W.,
Crawshay, S., Moore, D. R., … Coffey, V. G. (2014). Reduced
resting skeletal muscle protein synthesis is rescued by resistance
exercise and protein ingestion following short-term energy
deficit. American Journal of Physiology: Endocrinology and Meta-
bolism, 306, E989–E997. Retrieved from http://www.ncbi.nlm.
nih.gov/pubmed/24595305
Areta, J. L., Burke, L. M., Ross, M. L., Camera, D. M., West, D.
W., Broad, E. M., … Coffey, V. G. (2013). Timing and
distribution of protein ingestion during prolonged recovery
from resistance exercise alters myofibrillar protein synthesis.
Journal of Physiology, 591, 2319–2331. doi:10.1113/jphysiol.
2012.244897
Atherton, P. J., Etheridge, T., Watt, P. W., Wilkinson, D., Selby,
A., Rankin, D., … Rennie, M. J. (2010). Muscle full effect after
oral protein: Time-dependent concordance and discordance
between human muscle protein synthesis and mTORC1 signal-
ing. American Journal of Clinical Nutrition, 92, 1080–1088.
doi:10.3945/ajcn.2010.29819
Browne, G. J., & Proud, C. G. (2002). Regulation of peptide-chain
elongation in mammalian cells. European Journal of Biochemistry,
269, 5360–5368. Retrieved from http://www.ncbi.nlm.nih.gov/
pubmed/?term=Browne%2C+G.+J.%2C+%26+Proud%2C+C.
+G.+(2002) doi:10.1046/j.1432-1033.2002.03290.x
Burke, L. M., Winter, J. A., Cameron-Smith, D., Enslen, M.,
Farnfield, M., & Decombaz, J. (2012). Effect of intake of
different dietary protein sources on plasma amino acid profiles
at rest and after exercise. International Journal of Sport Nutrition

and Exercise Metabolism, 22, 452–462. Retrieved from http://
www.ncbi.nlm.nih.gov/pubmed/23212318
Carbone, J. W., Margolis, L. M., McClung, J. P., Cao, J. J.,
Murphy, N. E., Sauter, E. R., … Pasiakos, S. M. (2013).
Effects of energy deficit, dietary protein, and feeding on
intracellular regulators of skeletal muscle proteolysis. FASEB
Journal, 27, 5104–5111. doi:10.1096/fj.13-239228
Churchward-Venne, T. A., Burd, N. A., & Phillips, S. M. (2012).
Nutritional regulation of muscle protein synthesis with resist-
ance exercise: Strategies to enhance anabolism. Nutrition and
Metabolism, 9(1), 40. doi:10.1186/1743-7075-9-40
Churchward-Venne, T. A., Murphy, C. H., Longland, T. M., &
Phillips, S. M. (2013). Role of protein and amino acids in
promoting lean mass accretion with resistance exercise and
attenuating lean mass loss during energy deficit in humans.
Amino Acids, 45, 231–240. doi:10.1007/s00726-013-1506-0
Cuthbertson, D., Smith, K., Babraj, J., Leese, G., Waddell, T.,
Atherton, P., … Rennie, M. J. (2005). Anabolic signaling deficits
underlie amino acid resistance of wasting, aging muscle. FASEB
Journal, 19, 422–424. doi:10.1096/fj.04-2640fje
Drummond, M. J., Dreyer, H. C., Fry, C. S., Glynn, E. L., &
Rasmussen, B. B. (2009). Nutritional and contractile regulation
of human skeletal muscle protein synthesis and mTORC1 signal-
ing. Journal of Applied Physiology, 106, 1374–1384. doi:10.1152/
japplphysiol.91397.2008
Elia, M., Stubbs, R. J., & Henry, C. J. (1999). Differences in fat,
carbohydrate, and protein metabolism between lean and obese
subjects undergoing total starvation. Obesity Research, 7, 597–604.
doi:10.1002/j.1550-8528.1999.tb00720.x
Erdman, K. A., Tunnicliffe, J., Lun, V. M., & Reimer, R. A.
(2013). Eating patterns and composition of meals and snacks in
elite Canadian athletes. International Journal of Sport Nutrition
and Exercise Metabolism, 23, 210–219. Retrieved from http://
www.ncbi.nlm.nih.gov/pubmed/?term=Erdman%2C+K.+A.%
2C+Tunnicliffe%2C+J.%2C+Lun%2C+V.+M.%2C+%26
+Reimer%2C+R.+A.+(2013)
Garthe, I., Raastad, T., Refsnes, P. E., Koivisto, A., & Sundgot-
Borgen, J. (2011). Effect of two different weight-loss rates on body
composition and strength and power-related performance in elite
athletes. International Journal of Sport Nutrition and Exercise Meta-
bolism, 21, 97–104. Retrieved from http://journals.humankinetics.
com.libaccess.lib.mcmaster.ca/AcuCustom/Sitename/Documents/
DocumentItem/02-Garthe_J3212%20IJSNEM_2010_0037%20
97-104.pdf
Greenhaff, P. L., Karagounis, L. G., Peirce, N., Simpson, E. J.,
Hazell, M., Layfield, R., … Rennie, M. J. (2008). Disassoci-
ation between the effects of amino acids and insulin on
signaling, ubiquitin ligases, and protein turnover in human
muscle. American Journal of Physiology: Endocrinology and
Metabolism, 295, 595–604. doi:10.1152/ajpendo.90411.2008
Havemann, L., West, S. J., Goedecke, J. H., Macdonald, I. A., St
Clair Gibson, A., Noakes, T. D., & Lambert, E. V. (2006). Fat
adaptation followed by carbohydrate loading compromises
high-intensity sprint performance. Journal of Applied Physiology,
100, 194–202. doi:10.1152/japplphysiol.00813.2005
Helms, E. R., Zinn, C., Rowlands, D. S., & Brown, S. R. (2013). A
systematic review of dietary protein during caloric restriction in
resistance trained lean athletes: A case for higher intakes. Interna-
tional Journal of Sport Nutrition and Exercise Metabolism. Advance
online publication. Retrieved from http://journals.humankinetics.
com/AcuCustom/Sitename/Documents/DocumentItem/Helms_
ijsnem_2013_0054-in%20press.pdf
Heymsfield, S. B., Thomas, D., Nguyen, A. M., Peng, J. Z.,
Martin, C., Shen, W., … Muller, M. J. (2011). Voluntary
weight loss: Systematic review of early phase body composition
changes. Obesity Reviews, 12, 348–361. doi:10.1111/j.1467-
789X.2010.00767.x
Considerations for protein intake in weight loss 27
Josse, A. R., Atkinson, S. A., Tarnopolsky, M. A., & Phillips, S. M.
(2011). Increased consumption of dairy foods and protein
during diet- and exercise-induced weight loss promotes fat
mass loss and lean mass gain in overweight and obese premeno-
pausal women. Journal of Nutrition, 141, 1626–1634. doi:10.3945/
jn.111.141028
Kimball, S. R. (2014). Integration of signals generated by
nutrients, hormones, and exercise in skeletal muscle. American
Journal of Clinical Nutrition, 99, 237–242. doi:10.3945/ajcn.113.
068387
Krieger, J. W., Sitren, H. S., Daniels, M. J., & Langkamp-
Henken, B. (2006). Effects of variation in protein and carbo-
hydrate intake on body mass and composition during energy
restriction: A meta-regression. American Journal of Clinical
Nutrition, 83, 260–274. Retrieved from http://www.ncbi.nlm.
nih.gov/pubmed/?term=Effects+of+variation+in+protein+and
+carbohydrate+intake+on+body+mass+and+composition+duri
ng+energy+restriction%3A+a+meta-regression
Maestu, J., Eliakim, A., Jurimae, J., Valter, I., & Jurimae, T.
(2010). Anabolic and catabolic hormones and energy balance
of the male bodybuilders during the preparation for the
competition. Journal of Strength and Conditioning Research, 24,
1074–1081. doi:10.1519/JSC.0b013e3181cb6fd3
Mettler, S., Mitchell, N., & Tipton, K. D. (2010). Increased
protein intake reduces lean body mass loss during weight loss in
athletes. Medicine and Science in Sports and Exercise, 42, 326–
337. doi:10.1249/MSS.0b013e3181b2ef8e
Millward, D. J. (2001). Protein and amino acid requirements of
adults: Current controversies. Canadian Journal of Applied Physi-
ology, 26, 130–140. Retrieved from http://www.ncbi.nlm.nih.gov/
pubmed/?term=Millward%2C+D.+J.+(2001).+Protein+and
+amino+acid+requirements+of+adults%3A+current+controver
sies doi:10.1139/h2001-048
Moore, D. R., Areta, J., Coffey, V. G., Stellingwerff, T., Phillips,
S. M., Burke, L. M., … Hawley, J. A. (2012). Daytime pattern
of post-exercise protein intake affects whole-body protein
turnover in resistance-trained males. Nutrition and Metabolism,
9(1), 91. doi:10.1186/1743-7075-9-91
Moore, D. R., Robinson, M. J., Fry, J. L., Tang, J. E., Glover, E.
I., Wilkinson, S. B., … Phillips, S. M. (2009). Ingested protein
dose response of muscle and albumin protein synthesis after
resistance exercise in young men. American Journal of Clinical
Nutrition, 89, 161–168. doi:10.3945/ajcn.2008.26401
O’Connor, H., Olds, T., & Maughan, R. J. (2007). Physique and
performance for track and field events. Journal of Sports Sciences,
25, 49–60. doi:10.1080/02640410701607296
Paoli, A., Grimaldi, K., D’Agostino, D., Cenci, L., Moro, T.,
Bianco, A., & Palma, A. (2012). Ketogenic diet does not affect
strength performance in elite artistic gymnasts. Journal of the
International Society of Sports Nutrition, 9(1), 34. doi:10.1186/
1550-2783-9-34
Pasiakos, S. M., Cao, J. J., Margolis, L. M., Sauter, E. R.,
Whigham, L. D., McClung, J. P., … Young, A. J. (2013).
Effects of high-protein diets on fat-free mass and muscle
protein synthesis following weight loss: A randomized con-
trolled trial. FASEB Journal, 27, 3837–3847. doi:10.1096/fj.13-
230227
Pasiakos, S. M., Vislocky, L. M., Carbone, J. W., Altieri, N.,
Konopelski, K., Freake, H. C., … Rodriguez, N. R. (2010).
Acute energy deprivation affects skeletal muscle protein synthesis
and associated intracellular signaling proteins in physically active
adults. Journal of Nutrition, 140, 745–751. doi:10.3945/jn.109.
118372
Phillips, S. M. (2004). Protein requirements and supplementation
in strength sports. Nutrition, 20, 689–695. doi:10.1016/j.nut.
2004.04.009
28 C. H. Murphy et al.
Phillips, S. M., Glover, E. I., & Rennie, M. J. (2009). Alterations
of protein turnover underlying disuse atrophy in human
skeletal muscle. Journal of Applied Physiology, 107, 645–654.
doi:10.1152/japplphysiol.00452.2009
Phillips, S. M., Tang, J. E., & Moore, D. R. (2009). The role of
milk- and soy-based protein in support of muscle protein synthesis
and muscle protein accretion in young and elderly persons.
Journal of the American College of Nutrition, 28, 343–354. Retrieved
from http://www.ncbi.nlm.nih.gov/pubmed/?term=The+role+of
+milk-+and+soy-based+protein+in+support+of+muscle+protein
+synthesis+and+muscle+protein+accretion+in+young+and+elde
rly+persons
Phillips, S. M., & Van Loon, L. J. (2011). Dietary protein for
athletes: From requirements to optimum adaptation. Journal of
Sports Sciences, 29, 29–38. doi:10.1080/02640414.2011.619204
Phillips, S. M., & Zemel, M. B. (2011). Effect of protein, dairy
components and energy balance in optimizing body composi-
tion. Nestle Nutrition Institution Workshop Series, 69, 97–108.
doi:10.1159/000329288
Rennie, M. J., Wackerhage, H., Spangenburg, E. E., & Booth, F. W.
(2004). Control of the size of the human muscle mass. Annual
Review of Physiology, 66, 799–828. doi:10.1146/annurev.
physiol.66.052102.134444
Res, P. T., Groen, B., Pennings, B., Beelen, M., Wallis, G. A.,
Gijsen, A. P., … Van Loon, L. J. (2012). Protein ingestion
before sleep improves postexercise overnight recovery. Medicine
and Science in Sports and Exercise, 44, 1560–1569. doi:10.1249/
MSS.0b013e31824cc363
Rodriguez, N. R., DiMarco, N. M., & Langley, S. (2009). Position of
the American dietetic association, dietitians of Canada, and the
American college of sports medicine: Nutrition and athletic
performance. Journal of the American Dietetic Association, 109,
509–527. Retrieved from http://www.dietitians.ca/downloadable--
content/public/noap-position-paper.aspx
Schaafsma, G. (2000). The protein digestibility-corrected amino
acid score. Journal of Nutrition, 130, 1865–1867. Retrieved
from http://www.ncbi.nlm.nih.gov/pubmed/10867064
Sundgot-Borgen, J., & Garthe, I. (2011). Elite athletes in aesthetic
and Olympic weight-class sports and the challenge of body
weight and body compositions. Journal of Sports Sciences, 29,
101–114. doi:10.1080/02640414.2011.565783
Tang, J. E., & Phillips, S. M. (2009). Maximizing muscle protein
anabolism: The role of protein quality. Current Opinion in Clinical
Nutrition and Metabolic Care, 12(1), 66–71. doi:10.1097/
MCO.0b013e32831cef75
Umeda, T., Nakaji, S., Shimoyama, T., Yamamoto, Y., Totsuka,
M., & Sugawara, K. (2004). Adverse effects of energy restric-
tion on myogenic enzymes in judoists. Journal of Sports Sciences,
22, 329–338. doi:10.1080/0264041031000140446
Villareal, D. T., Smith, G. I., Shah, K., & Mittendorfer, B.
(2012). Effect of weight loss on the rate of muscle protein
synthesis during fasted and fed conditions in obese older
adults. Obesity (Silver Spring), 20, 1780–1786. doi:10.1038/
oby.2011.280
Walberg, J. L., Leidy, M. K., Sturgill, D. J., Hinkle, D. E.,
Ritchey, S. J., & Sebolt, D. R. (1988). Macronutrient content
of a hypoenergy diet affects nitrogen retention and muscle
function in weight lifters. International Journal of Sports Medi-
cine, 9, 261–266. doi:10.1055/s-2007-1025018
Weinheimer, E. M., Sands, L. P., & Campbell, W. W. (2010).
A systematic review of the separate and combined effects
of energy restriction and exercise on fat-free mass in middle-
aged and older adults: Implications for sarcopenic obesity.
Nutrition Reviews, 68, 375–388. doi:10.1111/j.1753-4887.2010.
00298.x
West, D. W., Burd, N. A., Coffey, V. G., Baker, S. K., Burke,
L. M., Hawley, J. A., … Phillips, S. M. (2011). Rapid
aminoacidemia enhances myofibrillar protein synthesis and
anabolic intramuscular signaling responses after resistance exer-
cise. American Journal of Clinical Nutrition, 94, 795–803.
doi:10.3945/ajcn.111.013722
Wullschleger, S., Loewith, R., & Hall, M. N. (2006). TOR
signaling in growth and metabolism. Cell, 124, 471–484.
doi:10.1016/j.cell.2006.01.016
Wycherley, T. P., Moran, L. J., Clifton, P. M., Noakes, M., &
Brinkworth, G. D. (2012). Effects of energy-restricted high-
protein, low-fat compared with standard-protein, low-fat diets:
A meta-analysis of randomized controlled trials. American
Journal of Clinical Nutrition, 96, 1281–1298. doi:10.3945/
ajcn.112.044321
Yeo, W. K., Paton, C. D., Garnham, A. P., Burke, L. M., Carey,
A. L., & Hawley, J. A. (2008). Skeletal muscle adaptation and
performance responses to once a day versus twice every second
day endurance training regimens. Journal of Applied Physiology,
150, 1462–1470. doi:10.1152/japplphysiol.90882.2008
Yeo, W. K., Carey, A. L., Burke, L., Spriet, L. L., & Hawley, J. A.
(2011). Fat adaptation in well-trained athletes: Effects on cell
metabolism. Applied Physiology Nutrition and Metabolism, 36(1),
12–22. doi:10.1139/h10-089