Sports Medicine

https://doi.org/10.1007/s40279-025-02213-6

REVIEW ARTICLE

Nutritional Strategies to Improve Post‑exercise Recovery

and Subsequent Exercise Performance: A Narrative Review
Alireza Naderi1 · Jeffrey A. Rothschild2,3 · Heitor O. Santos4 · Amin Hamidvand5 · Majid S. Koozehchian6 ·
Abdolrahman Ghazzagh7 · Erfan Berjisian8 · Tim Podlogar9,10

Accepted: 13 March 2025

© The Author(s) 2025

Abstract
Post-exercise recovery strategies influence the body’s ability to restore physiological homeostasis, replenish energy stores,
repair muscle damage, and promote desired adaptations, which improve exercise performance. This narrative review examines
the impact of nutritional strategies commonly used for enhancing recovery and subsequent exercise performance, particu-
larly when athletes face short recovery periods. Carbohydrate ingestion is essential for glycogen replenishment, especially
within the initial hours post-exercise, with its impact dependent on the types, timing, and amount. Protein is essential for
accelerating muscle recovery and achieving a positive nitrogen balance, depending on the type and dosage. The co-ingestion
of carbohydrates with proteins or fats is explored for its role in maximizing glycogen resynthesis and muscle repair, with
evidence supporting the addition of protein to suboptimal carbohydrate intake for enhanced recovery. Moreover, this review
addresses the potential benefits of creatine and caffeine co-ingestion for accelerating glycogen synthesis and improving
subsequent performance. Hydration strategies, including the use of milk-based beverages and electrolyte solutions, are also
discussed, emphasizing their importance in maintaining fluid balance and optimizing recovery. This review also highlights
the emerging role of micronutrients such as omega-3 fatty acids, antioxidants, and sodium bicarbonate in reducing muscle
damage and improving acid–base balance. Evidence supports the tailored use of these nutritional strategies, particularly for
athletes managing tight competition/training schedules. Future research should focus on refining individualized approaches
for recovery and investigating the impact of novel supplements on subsequent performance.

Despite the growing body of literature, practical guide-

1 Introduction
lines for utilizing nutrition to maximize post-exercise recov-
ery are still evolving. The goal of this review is to synthesize
Effective post-exercise recovery is critical for restoring phys-
the available research on nutritional strategies to enhance
iological homeostasis, repairing muscle damage, replenish-
post-exercise recovery and subsequent exercise performance,
ing energy stores, and facilitating adaptations that enhance
defined as an exercise performance occurring within ~ 2–24
athletic performance [1]. This becomes especially important
h of a prior bout of exhaustive exercise. Specifically, we
when athletes are required to compete multiple times within
will explore the roles of dietary macronutrients (carbohy-
a short period. In addition, managing fatigue and training
drate, protein, and fat), micronutrients, creatine, caffeine,
load is essential for athletes aiming to achieve peak perfor-
fluids and electrolytes, and sodium bicarbonate in accel-
mance and prevent overtraining [1–3]. Nutritional strategies
erating recovery processes (Fig. 1). The literature search
play a pivotal role in the recovery process by replenishing
was conducted by using the following two databases: Pub-
energy stores, maintaining energy availability [4], and pro-
Med/MEDLINE and Google Scholar, using combinations
moting repair mechanisms and adaptive responses to exer-
of keywords and Boolean operators (“subsequent exercise
cise [5, 6]. Therefore, defining optimal nutritional interven-
performance”) AND (“carbohydrates” OR “carbohydrate
tions to promote recovery after exhaustive exercise is crucial
and protein” OR “protein”) AND (“dietary fats” OR essen-
for improving performance in subsequent bouts.
tial fatty acid) AND (“creatine” OR “carbohydrate and
creatine” OR) AND (“sodium bicarbonate OR NaH3OC”)
AND (“caffeine” OR “carbohydrate and caffeine”) AND
(“micronutrients OR vitamins”). The search was performed
Extended author information available on the last page of the article

Vol.:(0123456789)
 A. Naderi et al.

ingested carbohydrate (Fig. 2). Muscle glycogen demon-

Key Points strates a pronounced affinity for restoration during the ini-
tial 2 h window following exercise, a process that notably
Athletes should consume adequate carbohydrates, pro- occurs independent of insulin’s influence [10]. Although
tein, and fluids to optimize post-exercise recovery, which some muscle glycogen can be repleted in the absence of
is critical to maintain performance in the subsequent food ingestion [11], contemporary guidelines recommend
competition/training session when the recovery period is ingesting 1–1.2 g·kg−1⋅h−1 of carbohydrate in the initial
less than 24 h. 4-h post-exercise window [12]. In contrast, delaying post-
While creatine, caffeine, and sodium bicarbonate sup- exercise carbohydrate ingestion by 2 h can lead to reduced
plementation may improve post-exercise recovery and muscle glycogen concentrations 4 h after exercise compared
subsequent exercise performance in certain scenarios, with immediate ingestion [13] and may impair exercise per-
their intake should be personalized to accelerate recov- formance the following day [14]. Complete replenishment
ery for athletes preparing for the next training session or of muscle glycogen stores typically unfolds over a 24–36 h
competition. The role of dietary fat and micronutrient period post-exercise, provided that 7–12 g·kg−1 carbohydrate
ingestion during the post-exercise recovery period and are consumed per day [15–18]. Liver glycogen resynthesis
subsequent exercise performance should be investigated proceeds at a slower pace during the initial post-exercise
in future research. hours, achieving saturation only after 11–25 h, contingent
upon adherence to a high-carbohydrate diet [16].
Future studies should focus on sex differences in The type of carbohydrate consumed influences the rate
response to carbohydrate, protein, and ergogenic aids, of muscle glycogen resynthesis following strenuous exer-
including creatine, caffeine, and sodium bicarbonate sup- cise. Early research demonstrated that ingestion of glucose
plementation, during the acute recovery period in condi- alone increases muscle glycogen levels significantly more
tions such as real-world competition simulation, consid- than fructose alone [19, 20]. However, since glucose and
ering individualized approaches and training status. fructose are absorbed via distinct intestinal transporters,
it was hypothesized that their combined ingestion could
optimize carbohydrate absorption, minimize gastrointesti-
nal discomfort, and potentially maximize muscle glycogen
replenishment compared with glucose alone [21]. Wal-
through the titles, abstracts, and keywords of documents lis et al. [21] were the first to test this hypothesis, finding
indexed in the databases. We classified the paper into three that ingesting 90 g of carbohydrate (~ 1.2 g·kg⁻1·h⁻1) in a
categories: **grade I—strong evidence**: this category 2:1 ratio of glucose to fructose during a 4-h post-exercise
includes a significant number of well-controlled trials that recovery phase did not enhance muscle glycogen resynthesis
consistently demonstrate the effectiveness of the compound compared with glucose alone. This finding was confirmed
for improving post-exercise recovery; **grade II—moder- by Trommelen et al. [22], who reported that co-ingesting
ate evidence**: this category contains a substantial but less glucose polymers with fructose or sucrose at a higher dose
consistent body of research showing the effectiveness of the (1.5 g·kg–1⋅h–1) did not further augment muscle glycogen
compound for post-exercise recovery; and **grade III—lim- replenishment during a 5-h recovery window, despite a
ited evidence**: this category includes a small and/or highly notable reduction in gastrointestinal complaints associated
ambiguous body of research that indicates the effectiveness with sucrose or fructose compared with glucose alone. This
of the compound for post-exercise recovery. finding aligns with subsequent studies comparing glucose,
fructose, and sucrose in < 1 g·kg–1⋅h–1 doses during a 4–6 h
post-exercise timeframe [19, 23]. More recently, Podlogar
2 Grade I—Strong Evidence et al. [24] posited that a combination of galactose and glu-
cose could yield similar results as compared with glucose
2.1 Dietary Carbohydrates alone in promoting muscle glycogen resynthesis. Contrary
to their hypothesis, their findings revealed that glucose alone
2.1.1 Carbohydrate Timing and Type at the rate of 1.2 g·kg−1⋅h–1 facilitated a 1.3–1.6 times higher
post-exercise muscle glycogen synthesis rate than galactose
Replenishing glycogen stores through immediate post- alone or a glucose + galactose mix. In summary, current
exercise carbohydrate intake is crucial for recovery, espe- research suggests that glucose and glucose-derived carbo-
cially when athletes need to be fully prepared for another hydrates, with or without the addition of fructose, are the
training session or competition within 24 h [7–9]. The rate most effective sources for replenishing post-exercise muscle
of restoration is influenced by both the type and timing of glycogen within a recovery window ≤ 8 h, and that addition
Nutritional Strategies to Improve Post-exercise Recovery and Subsequent Exercise Performance

Fig. 1  Examples of nutritional strategies to accelerate the short-term recovery period between two exercise bouts to enhance subsequent exercise
sessions

of monosaccharide galactose results in inferior replenish- However, not all studies have shown positive out-
ment rates [24]. There is currently no evidence to recom- comes with a combination of glucose and fructose. For
mend consuming more than 1.2 g·kg−1⋅h−1 of carbohydrates, instance, in a study examining time trial performance
regardless of type [22, 25]. Nevertheless, co-ingestion of after a glycogen-depleting exercise, participants ingested
glucose with fructose is preferred owing to reduced gas- 1.2 g·kg −1 ·h −1 of a glucose–fructose mix or glucose
trointestinal complaints and the beneficial effect on liver combined with maltodextrin over a 4-h recovery period
glycogen replenishment, as discussed below [18, 22, 26] before undertaking 1 h of moderate-intensity steady state
(Supplementary Material 1). cycling followed by a ~ 40-min cycling time trial [35].
From a performance perspective, several studies have Despite higher carbohydrate oxidation rates (indicating
demonstrated that co-ingesting glucose and fructose-based higher carbohydrate availability) following the inges-
carbohydrates can enhance the recovery of exercise capac- tion of the glucose–fructose mixture compared with the
ity [29–32]. For example, Maunder et al. [33] compared the glucose–maltodextrin mixture, performance was not
effects of 90 g of maltodextrin mixed with either glucose or improved. A mechanistic explanation for this requires
fructose (in a 1.5:1 ratio) following a glycogen-depleting further research, whereas from the applied perspective, it
run. They found that the maltodextrin–fructose combination remains to be properly investigated if carbohydrate inges-
increased subsequent endurance running capacity by ~ 33% tion during subsequent exercise bout would alter the study
over the maltodextrin–glucose mix, highlighting the poten- results. In intermittent sports, Hengist et al. [36] found
tial of glucose–fructose to enhance endurance capacity. Sim- no improvement in either average running speed or the
ilarly, Gray et al. [34] reported that consuming glucose and number of sprints after participants consumed a recovery
fructose during the initial 4 h and again at 15 h of recovery drink containing glucose or a glucose–fructose mixture
from exhaustive exercise resulted in a notable increase in at 0.8 g·kg −1⋅h −1 plus 0.3 g·kg−1⋅h −1 of protein during
cycling performance capacity, ~ 27% longer compared with 3 h of recovery after high-intensity rugby exercise test.
glucose alone. Authors suggested beneficial effects to be In addition, consuming carbohydrate drinks containing
attributed to differences in hepatic glycogen storage on the glucose and fructose at doses of 30 and 60 g·h⁻1 did not
basis of existing literature (discussed in Sect. 2.1.2), but this improve subsequent taekwondo kick performance during
was not measured in these studies. the simulated tournament with 45 min recovery between
 A. Naderi et al.

a b
Effect of CHO timing Effect of CHO type on muscle glycogen
4-h muscle glycogen resynthesis

4-h muscle glycogen resynthesis

250 2 g ⋅ kg−1 glucose ingested immediately 250 g−1
1.2 g ⋅ kkg
1.2 −1
⋅ h−1
−1
of d
of differing
iffering ttypes
y pe
pes
s ffor
or 4 h
f llowing exercise
fo
or 2-h following
(mmol/kg/dry mass)

(mmol/kg/dry mass)
200 200

150
* 150
* *
100 100

50 50

0 0
Immediately post-exercise 2-h post-exercise Glucose Glu+Fru Glucose Galactose Glu+Gal
Ingestion time Carbohydrate type
Adapted from Ivy et al. 1988 Adapted from Wallis et al. 2008 and Podlogar et al. 2023

c Effect of Glycemic Index

d Effect of CHO type on liver glyocgen

* * *
24-h muscle glycogen resynthesis

6-h liver glycogen resynthesis

500
200
(mmol/kg/dry mass)

400
150
(mmol/L)

300

100
200

50
100

0 0
High-GI Low-GI Mal+Fructose Mal+Galactose Mal+Glucose
Glycemic Index Carbohydrate type
Adapted from Burke et al. 1993 Adapted from Decombaz et al. 2011

Fig. 2  Influence of timing (a) and type (b–d), of carbohydrate (CHO) on glycogen resynthesis *indicates significant differenes (p < 0.05) from
other groups within a given study. Adapted from [13, 21, 24, 27, 28]. Fru fructose, Gal galactose, GI glycemic index, GLu glucose

five repeated kick tests [37]. These mixed results suggest Carbohydrate types are often defined on the basis of their
that the type of exercise, its intensity and duration, the spe- glycemic index (GI), a metric comparing the postprandial
cific carbohydrate dosages used, and the glycogen deple- blood glucose response of a food to that of glucose [38].
tion protocols might play critical roles in determining the Therefore, high-GI foods, which rapidly deliver glucose into
efficacy of carbohydrate supplementation (Supplementary circulation, are thought to facilitate muscle glycogen resyn-
Material 2). thesis, a process enhanced by heightened insulin activity dur-
Therefore, on the basis of current research findings, it is ing the post-exercise recovery phase [27]. Supporting this,
inferred that the co-ingestion of glucose and fructose-based research has shown that ingesting high-GI carbohydrates
carbohydrates during the post-exercise recovery phase is at a dose of 10 g·kg–1·24 ­h–1 significantly enhanced post-
likely more beneficial than other carbohydrate sources, given exercise muscle glycogen restoration compared with low
its superior efficacy in faster gastrointestinal absorption and glycemic index carbohydrates (~ 106 versus 72 mmol·kg⁻1
glycogen stores and its association with reduced gastroin- wet weight, for high- and low-GI, respectively) over a 24-h
testinal discomfort when an optimal carbohydrate dosage recovery period [27]. However, the influence of GI on sub-
of 1–1.2 g·kg−1⋅h−1 is consumed [18]. sequent exercise performance appears inconsistent. While
Nutritional Strategies to Improve Post-exercise Recovery and Subsequent Exercise Performance

one study found enhanced endurance capability following However, it is challenging to assess the performance impli-
high-GI carbohydrate consumption [39], others found no cations of these changes in liver glycogen levels after con-
effect [40–42] or even superior performance with low-GI suming fructose or galactose-based carbohydrate sources
carbohydrate intake, attributed to improved glucose stability by measuring subsequent exercise capacity. Supporting
and increased fat oxidation due to lower insulin activity [43] this hypothesis, it was observed that the addition of either
(Supplementary Material 1). fructose or galactose to maltodextrin following exercise sig-
In summary, although high-GI carbohydrate ingestion nificantly enhanced the rate of hepatic glycogen synthesis
leads to higher glycogen content after 24 h of recovery [27], compared with a combination of maltodextrin and glucose
during very short recovery periods the total carbohydrate [28] (Supplementary Material 1).
dose appears more important than the GI. As discussed
earlier, when fructose, which has a lower GI, is combined 2.2 Protein
with glucose, it is as effective as glucose alone in promoting
muscle glycogen replenishment. In addition, co-ingestion 2.2.1 Protein Type and Dose
of carbohydrates with sources of fiber and fat can signifi-
cantly affect the glycemic response [44]. Therefore, further Protein ingestion after exercise is considered one of the
research is needed to thoroughly investigate the impact of pillars of recovery owing to its role in promoting muscle
different glycemic index foods on both muscle and liver protein synthesis (MPS), reducing muscle protein break-
glycogen replenishment during rapid recovery time lasting down, and facilitating muscle repair and adaptation fol-
2–8 h, and on subsequent exercise performance, particularly lowing exercise-induced damage [51–54]. Adequate pro-
using various glycogen-depleting exercise protocols within tein intake is particularly essential when rapid recovery is
short recovery timeframes. required between training sessions or competitions [55].
The optimal dose of protein for post-exercise recovery
2.1.2 Liver Glycogen has been extensively studied, with research suggesting
that 20–40 g of high-quality protein consumed after exer-
Owing to methodological constraints in measuring liver gly- cise maximizes MPS rates for several hours (i.e., 3–4 h),
cogen levels, most sports research has focused on optimizing the exact dose dependent on muscle mass involved dur-
post-exercise muscle glycogen synthesis, despite the impor- ing exercise [56–58]. While the importance of immediate
tance of both glycogen stores [45]. The liver plays a pivotal post-exercise protein intake remains debated in literature
role in metabolic regulation during exercise by controlling [59–61], this controversy largely arises from differences
blood glucose levels through glycogen storage and mobiliz- in study designs and varying recovery conditions [62, 63].
ing non-carbohydrate sources via gluconeogenesis [26, 46]. Nevertheless, it is commonly recommended to rapidly
Liver glycogen stores are depleted by approximately 40–60% increase amino acid availability following exercise to max-
during moderate to high-intensity exercise sessions that imize MPS rates and promote muscle recovery [55, 64],
extend beyond 90 min without carbohydrate ingestion [23]. with numerous studies demonstrating positive effects on
Therefore, implementing nutritional strategies that reduce recovery in endurance athletes [62, 63, 65]. Following the
liver glycogen utilization during exercise and/or promote its initial protein-based meal after exercise, it is advised that
replenishment post-exercise may be as imperative as strate- subsequent protein-rich meals follow at 3–4 h intervals
gies for muscle glycogen replenishment [8, 47]. [55, 66, 67]. However, it seems that more than the tim-
While carbohydrate ingestion during exercise does not ing of the meals, the total amount of protein plays a more
substantially affect muscle glycogen utilization [48, 49], important role than the timing itself [53, 68]. Endurance
it can reduce or even completely suppress liver glycogen athletes are recommended to consume ~ 1.2–1.8 g·kg –1
breakdown, thereby improving subsequent recovery [49, 50]. of high-quality protein [69–72], while those engaged in
Moreover, liver glycogen resynthesis is inherently slower power-based sports are advised to ingest 1.7–2.2 g·kg –1
than that of muscle, possibly owing to its higher rate of [73]. In addition, vegan athletes might require a higher
glycogen turnover and/or distinct glycogen synthesis path- daily intake to obtain sufficient amount of all amino acids
ways that are contingent on hormonal fluctuations (nota- [74]. Given the practical challenges athletes face in con-
bly insulin, glucagon, and epinephrine) [45, 47]. Research suming large quantities of protein in a single meal, distrib-
has demonstrated that the rate of liver glycogen resynthesis uting protein intake across multiple meals throughout the
within the initial 2-h post-exercise period is 30–50% lower day is a sensible approach. Considering the extended fast-
than during the subsequent 3–5 h [16]. Consequently, It has ing period during sleep, a pre-sleep protein dose of ≥ 40 g
been hypothesized that certain types of carbohydrates, such has been recommended to sustain MPS rates and minimize
as fructose and galactose, which are metabolized by the overnight muscle protein breakdown [75, 76]. However,
liver, might optimize the rate of liver glycogen synthesis. two studies investigating this did not observe any positive
 A. Naderi et al.

effects on recovery indices when total daily protein intake (VO2peak) when protein was co-ingested with carbohydrate
is sufficient [77, 78]. within 90 min following 5 h of recovery, attributed to higher
The type of protein consumed also significantly influ- muscle glycogen synthesis rates and positive nitrogen bal-
ences recovery efficacy. Amounts of essential amino acids, ance compared with carbohydrate alone in trained male
and especially leucine content, in protein sources seem to endurance athletes. In addition, exercise capacity during
be the strongest predictor of MPS [79, 80]. Whey protein, high-intensity interval cycling was significantly greater
a fast-digesting and leucine-rich source, has been repeat- with carbohydrate-protein co-ingestion (0.8 g·kg −1·h⁻ 1of
edly demonstrated to enhance MPS more effectively than carbohydrates and 0.4 g·kg−1·h⁻1 of protein) compared with
casein or soy protein owing to its rapid absorption kinetics carbohydrate alone (1.2 g·kg−1·h⁻1) following a 2-h recovery
and high essential amino acid content [81]. Casein protein, period [90]. These positive results were mainly attributed to
by contrast, provides a slower release of amino acids and greater muscle glycogen synthesis or positive nitrogen bal-
may be beneficial for sustaining protein synthesis over ance mediated by the additional protein [85].
longer recovery periods, such as overnight [81]. Plant- In contrast, recent studies have reported similar improve-
based proteins such as soy, rice, and pea protein, while ments in subsequent time-trial performance and high-inten-
effective [82], generally require higher doses (25–40 g) sity interval cycling capacity after either carbohydrates alone
to match the leucine threshold necessary for robust MPS or carbohydrate–protein co-ingestion during 2–5 h recovery
stimulation owing to lower essential amino acid content periods in both young and master endurance athletes [91, 92]
[53, 74, 83]. (Supplementary Material 3). These discrepancies may be
attributed to interindividual variations in training status and
2.3 Carbohydrate and Protein Co‑ingestion genetic factors, as well as differences in recovery durations,
glycogen depletion protocols, performance tests, amounts
Although carbohydrate intake is the cornerstone of glyco- and types of protein ingested, and methods of nutrient provi-
gen resynthesis, concomitant intake of protein has been sug- sion during recovery.
gested to confer additional benefits, primarily owing to the Taken together, combining protein with carbohydrates
insulinotropic effects of certain amino acids to reduce mus- during recovery periods shorter than 24 h likely has a small,
cle protein breakdown [84]. However, this benefit appears to but positive, effect on performance, ranging from 0.6 to 1.6%
occur only when protein is added to a suboptimal amount of [87]. Accordingly, it should be recommended as it may con-
carbohydrate (e.g., 0.3 g·kg⁻1·h⁻1 protein and 0.9 g·kg−1⋅h−1 fer additional benefits, including positive nitrogen balance
carbohydrate) [85]. Similarly, a meta-analysis confirmed leading to increased whole-body muscle protein synthesis
that ingesting sufficient carbohydrate should be the priority [93], increased protein content related to mitochondrial
when aiming to replenish muscle glycogen stores, report- biogenesis [94], reduced muscle damage markers [95], and
ing that co-ingestion of carbohydrate (0.86 g·kg−1⋅h−1) with accelerated post-exercise rehydration [96]. Therefore, co-
protein (0.27 g·kg−1⋅h−1) did not increase muscle glycogen ingestion of carbohydrates and protein during post-exercise
re-synthesis more than carbohydrate per se when ingested may be advantageous, especially for athletes engaging
at 0.95 g·kg−1⋅h−1 during short-term post-exercise recovery in multiple daily training sessions or with less than 24 h
(≤ 8 h) [86]. between bouts.
From a performance standpoint, co-ingesting protein
with carbohydrate during recovery between two bouts of 2.4 Fluid and Electrolytes
endurance exercise has shown small to moderate effects
on subsequent performance compared with carbohydrate Rehydration after exercise is another key aspect of recovery.
alone (e.g., improvements of ~ 1.6% in mean power output Exercising in a dehydrated state is well-established to nega-
during cycling time trials and 0.6% in competitive running tively affect both performance and health [97, 98]. During
performance [87]. However, even a 1% improvement can strenuous exercise in a hot or humid environment, athletes
be critical in elite sports, potentially making the difference may lose more than 2 L of sweat per hour, accompanied by
between finishing fourth and securing a podium position substantial electrolyte, extracellular, and intracellular water
in Olympic competitions [88]. Rustad et al. [89] reported loss that may negatively affect processes such as the rate of
that co-ingesting carbohydrate (0.8 g·kg⁻1·h⁻1) and protein post-exercise glycogen and muscle protein resynthesis [44,
(0.4 g·kg⁻1·h⁻1) during the first 2 h of an 18 h of recovery 99]. Therefore, re-establishing water and electrolyte balance
from strenuous cycling exercise led to ~ 14 min longer subse- during the post-exercise recovery period is important [100].
quent exercise capacity compared with carbohydrate intake Rehydration is particularly important before subse-
alone (1.2 g·kg−1·h⁻1). Similarly, Dahl et al. [63] observed quent training or competition when limited time is avail-
an ~ 8.5-min increase in exercise capacity during subsequent able between exercise bouts in environmentally challenging
cycling to exhaustion at 70% peak oxygen consumption conditions [101, 102]. Understanding pre- and intra-exercise
Nutritional Strategies to Improve Post-exercise Recovery and Subsequent Exercise Performance

fluid recommendations is fundamental before addressing simultaneously increasing glycogen resynthesis and stimu-
post-exercise hydration. Athletes are generally recommended lating muscle protein synthesis [114]. Interestingly, nonalco-
to slowly ingest 5–7 mL·kg⁻1 fluids (via food and liquids) holic beer has been studied as a post-hydration drink, since it
at least 4 h before exercise/competition [103]. Attempts to is rich in water, carbohydrates, and some minerals. However,
correct fluid loss induced by exercise should consider the as with other beverages, nonalcoholic beer could be consid-
sweat rate, which can vary from 0 to over 2 L·h⁻1, depend- ered on the basis of individual preference, but more studies
ing on factors such as environmental conditions, intensity, are needed owing to insufficient evidence [115].
duration, genetics, fitness level, acclimatization to the envi- Although water is one of the most common and acces-
ronment, and clothing [104, 105]. Although intra-exercise sible options for post-exercise rehydration, drinking water
hydration can mitigate fluid loss during exercise, post-exer- alone may not optimize hydration status, as it has a low BHI
cise hydration is critical for conditions involving subsequent value owing to the absence of osmotic nutrient content such
competitions. as electrolytes, carbohydrate, and protein [109]. Therefore,
If rapid recovery (< 24 h) is the aim or severe hypohydra- water alone may not be the best choice to maximize subse-
tion (> 5% body mass loss) is observed, intensive intake of quent exercise capacity [92, 116], especially considering that
fluids and electrolytes should be advised to enhance recov- other nutrients are also essential for recovery. Conversely,
ery before subsequent competition [106]. For example, daily factors such as an athlete’s eating behavior, thirst, palatabil-
water and sodium losses can vary significantly and be in the ity, and gastrointestinal comfort may influence the volume
range of 4–10 L and 3.5–7.0 g, respectively, in active ath- of other beverages consumed, leading to different recovery
letes under heat stress; thus, both need to be replaced to re- outcomes [117]. Studies examining the co-ingestion of vari-
establish normal total body water (i.e., achieve euhydration) ous foods with beverages ad libitum during 2–4 h of recov-
[106]. Beginning subsequent exercise in a dehydrated state ery from exercise-induced dehydration have indicated that,
may impair exercise performance. Therefore, rehydration regardless of beverage choice, similar recovery fluid balance
strategies between exercise bouts recommend that athletes [117, 118] and subsequent exercise performance [118] can
drink fluids equivalent to 125–150% of their body mass lost be achieved using milk-based formulations, sports drinks,
during exercise [107, 108]. or water along with different solid meals in both males and
Maughan et al. tested different beverages using the bever- females, despite varying dietary behaviors [117–119]. The
age hydration index (BHI), which compares the fluid reten- mechanism behind this may be related to delayed gastric
tion capacity 2 h post-ingestion compared with an equal emptying and increased plasma osmolality, which reduces
volume of water [109], finding oral rehydration solutions, urine output and enhances fluid retention [117] (Supplemen-
skimmed milk, whole milk, and orange juice had higher BHI tary Material 4, 5).
values compared with water. Notably, beverages such as cola
(regular and diet), tea (hot and iced), coffee, lager, sparkling
water, and sports drinks did not differ from water in terms 3 Grade II—Moderate Evidence
of BHI, suggesting that the volume of consumed liquid is
more important than its source for hydration. Therefore, it 3.1 Sodium Bicarbonate
is reasonable to use water or a mix of other beverages as
post-exercise hydrational strategies, tailored to individual Sodium bicarbonate ­(NaHCO3) supplementation before exer-
hydration needs and personal preferences [109]. cise is widely recognized for enhancing extracellular buffer-
Oral rehydration solutions and sports drinks are common ing capacity when administered at doses of 0.2–0.4 g·kg⁻1
strategies to improve fluid retention owing to their balanced body mass (BM) 1–3 h prior to activity [120]. This strat-
carbohydrate and electrolyte content [110]. However, stud- egy can improve high-intensity exercise performance last-
ies have observed no differences in fluid retention between ing approximately 30 s to 10 min [121]. The performance
post-exercise consumption of oral rehydration solutions and enhancements observed with N ­ aHCO3 supplementation are
sports drinks following exercise-induced dehydration, even attributed to increased extracellular buffering capacity, low-
in hot conditions [111, 112] or in subsequent endurance ered extracellular potassium ­(K+) levels, elevated sodium
performance [113]. Recent systematic reviews have found ­(Na+) levels before exercise, and a potential reduction in cel-
that dairy milk leads to a similar net fluid balance compared lular oxidative stress responses [122–124]. Another mecha-
with carbohydrate-electrolyte sports drinks and water when nism that may explain the ergogenic effect of post-exercise
consumed ad libitum and provides a higher net fluid bal- ­NaHCO3 ingestion is its hyperhydration effect, leading to
ance when consumed in similar volumes [114, 115]. Milk- an ~ 8% increase in blood volume through increased sodium
based beverages have also led to greater exercise capacity availability in the blood [125].
in subsequent exercise performance when compared with Initial studies indicated that N­ aHCO3 ingestion before
both noncaloric and energy-matched beverages, likely due to high-intensity exercise attenuated post-exercise blood
 A. Naderi et al.

acid–base imbalance during a 15-min passive recovery 3.2 Creatine and Carbohydrate Co‑ingestion
period compared with placebo [126] and facilitated a faster
restoration of acid–base balance [127]. In addition, ingest- Creatine (α-methyl-guanidinoacetic acid) is a widely used
ing 0.3 g·kg⁻1 BM of NaHCO₃ 180 min before exercise led nutritional supplement that has been extensively researched
to ~ 3.8% improvement in subsequent performance during over the past two decades [142]. It is well-documented that
four repeats of 2 × 4 km cycling time trials with 40 min pas- creatine supplementation can enhance adaptations to resist-
sive recovery conducted under hypoxic conditions as com- ance training, including increases in strength, power, and
pared with placebo [128]. More recently, research has shown muscular endurance [143]. In addition, creatine supplemen-
that administering ­NaHCO3 within approximately < 30 min tation may positively affect muscle glycogen storage in the
following exercise can restore extracellular acid–base bal- days following glycogen-depleting exercise [144].
ance during a 75–90 min recovery period between two bouts While the exact physiological mechanisms by which
of exercise and may improve subsequent exercise perfor- creatine augments post-exercise muscle glycogen storage
mance [129, 130] (Fig. 3), while others have reported that are not fully understood, several plausible mechanisms
a shorter recovery time of 35 min may be inadequate to have been proposed. Creatine may enhance the abundance
restored acid–base balance to improve subsequent exercise of glucose transporter (GLUT) 4, facilitating greater glu-
capacity [131] (Supplementary Material 6). cose uptake into muscle cells [145]. It may also mitigate
The lack of performance enhancement in some studies post-exercise inflammation, reducing markers of muscle
may be due to intraindividual variability in the time required damage and soreness [146]. This is important because exer-
to reach peak HCO₃⁻ levels and the occurrence of gastroin- cise-induced muscle damage (EIMD) can impede muscle
testinal discomfort [127, 132, 133]. De Oliveira et al. [121] glycogen replenishment for up to 10 days [147], suggesting
established a threshold increase of 4–6 mmol/L for circu- that creatine could facilitate glycogen resynthesis by alleviat-
lating ­HCO3⁻ levels to determine the ergogenic potential ing EIMD. Furthermore, as an osmotically active compound,
of ­NaHCO3 supplementation. This threshold depends on creatine can increase intracellular fluid volume [148], which
the time needed for circulating ­HCO3⁻ levels to rise and is may be beneficial because cell swelling has been identified
influenced by factors such as the dosage and timing of inges- as a potent stimulus for muscle glycogen synthesis [149].
tion [134], supplement form (i.e., solution versus capsules) However, creatine does not appear to affect muscle glyco-
[123, 135], potential adverse effects, genetic factors related gen recovery within the initial 6 h following exercise but
to hydrogen ion ­(H+) transporters, the individual’s training has been shown to increase glycogen storage over a 5-day
status, and exercise type [136]. period [144]. Other research has shown that creatine sup-
Taken together, administration of ­NaHCO3 after exercise plementation, combined with a high carbohydrate diet, led
can expedite the restoration of acid–base equilibrium fol- to 82% greater glycogen resynthesis within 24 h following
lowing intense physical activity, potentially improving per- exhaustive exercise compared with placebo [150]. Moreover,
formance during subsequent exercise sessions if an adequate Van Loon et al. [151] observed an 18% increase in mus-
recovery period of 30–90 min is provided [124, 129, 137]. cle glycogen content following 5 days of creatine loading
For athletes with longer recovery periods (> 90 min) who are (20 g·day–1), underscoring a notable glycogen storage boost
more susceptible to severe gastrointestinal symptoms, new without significant impacts on fasting plasma insulin levels
forms of NaHCO₃ such as enteric-coated, delayed-release or GLUT-4 mRNA/protein content. In addition, research
capsules and mini-tablet NaHCO₃ hydrogels could be con- by Nelson et al. [152] demonstrated that a moderate carbo-
sidered as they may mitigate gastrointestinal discomfort hydrate intake post-creatine loading could enhance muscle
[138–140]. In addition, administering ­NaHCO3 before exer- glycogen stores by 53% compared with a high-carbohydrate
cise can promote acid–base restoration and improve subse- diet alone, further indicating the efficacy of creatine sup-
quent exercise performance, particularly when the recovery plementation in enhancing glycogen storage, although the
period between sessions is less than 40 min [141]. However, exact mechanism remains unclear. However, an individual’s
further research is needed to examine the effects of differ- training status may influence creatine’s efficacy regarding
ent ­NaHCO3 supplementation forms and their impact on glycogen storage, as two studies focusing on highly trained
HCO₃⁻ kinetics during recovery between exercise sessions. athletes did not show significant enhancements in muscle
More research is needed to assess the impact of post-exercise glycogen levels following creatine supplementation [153,
­NaHCO3 supplementation in brief, high-intensity exercise 154]. Specifically, trained individuals might experience
bouts such as combat sports. reduced benefits from creatine supplementation in enhancing
glycogen storage owing to their already optimized glycogen
resynthesis capabilities following exercise, compared with
untrained individuals [155] (Supplementary Material 7).
Collectively, this evidence highlights the potential benefits
Nutritional Strategies to Improve Post-exercise Recovery and Subsequent Exercise Performance

a b
*
Control

* *
NaHCO3

Time to exhaustion (s)

300
7.5

7.4 200
pH

7.3 100

7.2
0
NaHCO 3 Placebo
t

ry

ry

ry
1

2
es

ve

ve

ve
e

e
R

is

is
co

co

co
rc

rc
re

re

re
xe

xe
-e

-e
in

in

in

Adapted from Gough et al. 2017

m

m
st

st
Po

Po
30

60

90

Fig. 3  Sodium bicarbonate (­NaHCO3) ingested 30 min follow- is highlighted in a. *Denotes significant differences between groups
ing exercise to exhaustion can increase pH (a) and improve time to (p < 0.05). Adapted from Gough et al. [129]
exhaustion during a subsequent exercise bout (b). Recovery period

and mechanisms of creatine supplementation in athletic per- of adenosine monophosphate-activated protein kinase
formance and recovery, although the impact of simultane- (AMPK), possibly through enhanced activation of the
ous creatine and carbohydrate consumption on subsequent ­Ca2+-calmodulin-dependent protein kinase (CaMK) and the
exercise performance remains debated. phosphorylation of acetyl-CoA carboxylase (ACC), which
In summary, the simultaneous ingestion of creatine and could collectively induce translocation of GLUT4 to the cell
carbohydrate can effectively augment muscle glycogen stor- membrane and promote muscle glucose uptake [162, 163].
age, particularly after the initial 6-h post-exercise period. However, the evidence regarding enhanced post-exercise
Despite these findings, further research is necessary to deter- glycogen synthesis with caffeine co-ingestion is mixed. For
mine the impact of creatine and carbohydrate co-ingestion example, no positive effect was observed when combining
on endurance performance in subsequent activities, with a caffeine (6 mg·kg–1) with glucose (~ 1 g·kg–1⋅h–1) on muscle
special focus on weight-bearing sports. It is also important glycogen replenishment rates during a 5-h recovery period
to note that most studies referenced involve male subjects. after exhaustive cycling [164]. In contrast, another study
Given that creatine metabolism exhibits sex-specific differ- reported that muscle glycogen restoration was ~ 66% higher
ences [156], there is a need for a better understanding of during 4 h of recovery with a very high dose of caffeine
creatine supplementation’s effects across sex and levels of (8 mg·kg–1) and moderate carbohydrate intake (1 g·kg–1·h–1)
athletic training. compared with carbohydrate ingestion alone in trained
endurance athletes [160]. However, both studies used carbo-
3.3 Caffeine and Carbohydrate Co‑ingestion hydrate doses below the optimal 1.2 g·kg⁻1·h⁻1 recommended
for maximal glycogen resynthesis [160, 164]. Another study
Caffeine (1,3,7-trimethylxanthine) is one of the most exten- examined the effect of caffeine co-ingestion with a more
sively researched performance-enhancing supplements and optimal carbohydrate dose, finding no significant differences
is known to enhance exercise performance when consumed in glycogen resynthesis when 1.7 mg·kg⁻ 1·h⁻1 of caffeine
before [157] and during prolonged exercise [158]. Recent was co-ingested with 1.2 g·kg⁻1·h⁻1 of carbohydrates (1:1
evidence suggests caffeine could also positively affect post- glucose and maltodextrin) during a 6-h recovery period,
exercise recovery owing to its ability to decrease muscle compared with carbohydrates alone. The authors con-
soreness [159] and, when paired with carbohydrates, accel- cluded that an optimal carbohydrate dose of 1.2 g·kg⁻1·h⁻1
erate muscle glycogen replenishment [160]. Loureiro et al. is sufficient to maximize glycogen replenishment rates, with
[161] proposed that caffeine might enhance muscle gly- no additional benefits from caffeine co-ingestion [165].
cogen synthesis by activating molecular pathways related Recently, Loureiro et al. investigated the impact of caffeine
to either energy sensing or muscle contraction. Specifi- co-ingestion through coffee combined with milk contain-
cally, caffeine co-ingestion could lead to higher activation ing 1.2 g·kg⁻1 of carbohydrates and 0.3 g·kg⁻1 of protein on
 A. Naderi et al.

muscle glycogen resynthesis, finding that the addition of caf- and a heightened risk of overnight insomnia [172, 173].
feine to milk resulted in approximately 153% higher rates of Therefore, further research focusing on glycogen storage
muscle glycogen synthesis compared with milk alone [166] measurements and involving larger sample sizes is necessary
(Supplementary Material 8). However, while the carbohy- to clarify the impact of caffeine–carbohydrate co-ingestion
drate dose was considered optimal, the carbohydrate com- on recovery and performance.
position (milk sugars) was not, as recent evidence suggests
that galactose–glucose mixtures are inferior as compared
with glucose-based carbohydrates when provided at a dose 4 Grade III—Limited Evidence
of 1.2 g·kg–1·h–1 [24] (Fig. 2). Therefore, the positive effects
of caffeine on muscle glycogen synthesis rates might be due 4.1 Dietary Fats
to suboptimal amount or type of carbohydrate provision.
Further studies are warranted to investigate caffeine and Fat is primarily stored as triacylglycerol (TG) in adipose tis-
glucose or glucose plus fructose combinations with different sue, with smaller amounts inside muscle fibers (intramyocel-
doses. Nevertheless, these results support the hypothesis that lular TG; IMTG) [174]. IMTG stores are used as an energy
components within coffee could stimulate enzymes asso- source mainly during early stages of endurance exercise,
ciated with muscle glycogen synthesis [161]. The positive with the reliance on them diminishing as exercise duration
effects of caffeine seem most apparent when muscle gly- extends in favor of plasma-derived substrate oxidation [175].
cogen stores are very low and/or carbohydrate availability IMTG are also utilized as a fuel source in the post-exercise
is suboptimal. Inconsistencies across studies may be attrib- recovery window [176]. However, their availability does not
uted to differences in exercise protocols, glycogen deple- seem to play a role in performance as interventions aimed
tion methods, recovery durations, dosages, and timing of at increasing IMTG have resulted in either no differences
carbohydrate and caffeine provision. [177] or worsened [178] performance in exercise trials
While caffeine–carbohydrate co-ingestion post-exercise lasting 20–140 min. In addition, there is no to very little
may enhance muscle glycogen repletion, its impact on sub- net utilization of IMTG during high-intensity intermittent
sequent exercise performance is unclear. For instance, Tay- exercise [179], which calls into question the necessity for
lor et al. [167] found that caffeine (6 mg·kg–1) combined replenishing stores prior to a competitive event. Although
with carbohydrate (1.2 g·kg–1·h–1 glucose) improved exer- a low-fat diet can attenuate repletion of IMTG following
cise capacity in an intermittent shuttle run test compared exercise [180], post-exercise plasma fatty acid availability is
with carbohydrate alone (~ 48 min versus 32 min). Con- increased independently of post-exercise dietary ingestion,
versely, Andrea-Souza et al. [168] did not observe signifi- mainly due to the hormonal response that stimulates adipose
cant performance enhancements in soccer players follow- tissue lipolysis and skeletal muscle blood flow during the
ing co-ingestion of caffeine (6 mg·kg⁻1) and carbohydrates recovery period [181]. However, there is a lack of evidence
(1.2 g·kg⁻ 1·h⁻ 1), possibly owing to insufficient glycogen to support targeted replenishment of IMTG to improve sub-
depletion in the testing protocol. Similarly, Barzegar et al. sequent exercise performance lasting less than 3 h, but future
[169] found no performance benefit with combined caffeine research in the context of repeated daily ultra-endurance per-
(6 mg·kg–1) and carbohydrate (0.6 g·kg–1) intake compared formance is warranted.
with carbohydrate alone or a placebo during a 5 × 500 m Ingestion of specific types of fats may have the potential
kayak test conducted 24 h after a 20-km time-trial paddle to aid in the recovery of muscle function and/or influence
used for glycogen depletion. Moreover, caffeine ingestion post-exercise glycogen resynthesis. Omega-3 fatty acids,
alone is sufficient to enhance performance in subsequent ses- specifically eicosapentaenoic acid (EPA) and docosahexae-
sions for shorter, high-intensity exercises such as wrestling noic acid (DHA) are types of fats that have reported anti-
[170] (Supplementary Material 9). inflammatory properties and the capacity for reducing EIMD
In summary, consuming caffeine alongside carbohydrates and enhancing muscle function following exercise [182],
during the initial post-exercise recovery hours may benefit providing additional rationale for their recommended use in
athletes who have multiple competitions in a single day or aiding exercise recovery in a dose- and duration-dependent
who employ training sessions with low carbohydrate availa- manner [183].
bility, such as the “sleep low, train low” thought to maximize Conjugated linoleic acid (CLA) refers to a mixture of
training adaptations [171]. Efficient repletion of glycogen polyunsaturated fatty acids that exists as various isomers
stores after a low carbohydrate-availability exercise session of linoleic acid (18:2), and chronic supplementation may
is crucial for subsequent training, especially those involv- positively influence glycogen resynthesis after exercise
ing high-intensity efforts [167]. It is important to note that [184]. Dietary sources include dairy and meat from rumi-
excessive caffeine intake can lead to adverse effects, includ- nant animals, which are primarily found as the cis-9 trans-11
ing tachycardia, anxiety, headaches, gastrointestinal issues, isomer; however, synthetic isomers have also been studied
Nutritional Strategies to Improve Post-exercise Recovery and Subsequent Exercise Performance

[185]. While most commonly studied in the context of body stimulate stress-related cell signaling pathways and increase
composition, humans have also shown an increase in post- the expression of genes encoding proteins, such as nuclear
exercise muscle GLUT4 protein expression and post-exer- respiratory factor 2 (NRF2), as well as pathways in skel-
cise muscle glycogen resynthesis rates following 8 weeks etal muscle, leading to increased mitochondrial biogenesis
of 3.8 g⋅day−1 of trans-10 cis-12 and cis-9 trans-11 CLA [206]. Over 8000 polyphenols have been identified, includ-
supplementation [184]. A study in rats supports these find- ing curcumin, resveratrol, quercetin, and catechins [206],
ings and provides some more mechanistical evidence for and are most commonly consumed in fruits, vegetables, and
these findings, namely showing increased muscle glycogen spices. Although most studies of antioxidants and exercise
synthase activity with a mixture of isomers (cis-9 trans-11, recovery have focused on markers of oxidative stress and/
cis-10 trans-12), but not with cis-9 trans-11 isomer, suggest- or muscle damage, rather than subsequent exercise perfor-
ing physiological effects are likely isomer-specific [186]. mance, quercetin supplementation for 7 days (1 g⋅day−1)
Although consuming dietary fat in the post-exercise meal improved time to exhaustion at 75% maximal oxygen con-
can prolong gastric emptying times [187], in the context sumption (VO2max) by 21.4% following a 3-h recovery period
of adequate carbohydrate ingestion, glycogen resynthesis is after 60 min of cycling compared with a placebo and also
not influenced by the addition of fat and protein in the 3-h reduced oxidative stress [207]. In contrast, 8 weeks of green
[188] or 24-h [189] post-exercise period. Therefore, athletes tea extract supplementation (500 mg⋅day−1) had no effect
can include protein and high-quality fat in the post-exercise on glycogen resynthesis, despite increases in postexercise
window without sacrificing muscle glycogen synthesis, bear- fat oxidation and exercise-induced muscle GLUT4 protein
ing in mind that very high-fat intakes in the post-exercise content [208].
window may attenuate some, but not all, signaling markers Free radicals produced via aerobic metabolism contrib-
[190]. ute both to exercise-induced fatigue as well as the adaptive
response to exercise [209]. This means there is potential for
4.2 Micronutrients high-dose antioxidants to blunt favorable training-induced
muscle adaptations. Despite attenuations in markers of meta-
Micronutrients consumed prior to exercise, including vita- bolic and/or mitochondrial adaptations, no impact on exercise
mins, minerals, and phytonutrients, also have the potential performance outcomes has been observed in human studies
to affect muscle recovery and therefore impact subsequent when supplementing with vitamins C and/or E during con-
exercise performance. This occurs primarily via reductions trolled endurance training [210]. However, this finding could
in EIMD and reductions in reactive oxygen and nitrogen be related to small sample sizes in studies. For example, two
species (RONS) leading to improved muscle contractile human studies showing a blunting of some markers of train-
function. ing adaptations also had the largest sample sizes [211, 212],
Several supplements may be effective in reducing EIMD, whereas a study showing no statistical effect of vitamin C on
including fruit extracts such as tart cherry [191], beetroot increases in VO2max reported increases of 10.8% in the supple-
[192], pomegranate [193], blueberry [194], watermelon mentation group (n = 5) and an increase of 22% in the group
[195], grape [196], and blackcurrant [197], as well as vita- (n = 9) without supplementation (Cohen’s d =  − 1.1, [95%
min E [198], curcumin [199], cacao [200], and vitamin D confidence interval (CI) − 2.2, 0.04]) [213]. Supplementa-
[201], with the strongest evidence surrounding tart cherry, tion with (-)-epicatechin impaired training adaptations during
beetroot, pomegranate, and vitamin D [202, 203]. These 4 weeks of cycling training, as increases in VO2peak (+ 22.6%)
benefits are likely due to antioxidant effects and have been and succinate dehydrogenase activity (+ 59.1%) were seen in
observed in both trained and untrained populations [203]. the placebo but not the supplement group, although no differ-
However, most studies have measured indicators of muscle ences in peak power or citrate synthase activity were observed
damage/inflammation rather than subsequent performance. between groups [214].
Despite reductions in muscle damage and/or inflammation, Supraphysiological doses of antioxidants may be benefi-
improvements in performance are not always observed [191, cial for enhancing acute recovery and subsequent performance,
192]. particularly during the competition period when adaptation is
Antioxidants have the potential to improve recovery and not the primary focus. However, long-term daily use should be
acute performance by reducing RONS, which impair muscle approached with caution, as research is inconclusive regard-
contractile force primarily through alterations in myofibrillar ing the extent of their effects on training adaptations, espe-
calcium sensitivity [204]. Often used as an umbrella term, cially during the muscle adaptation training phase [210]. This
antioxidants can exert their influence in different ways. Vita- could be related to the multiple pathways regulating skeletal
mins C and E share a common mechanism of donating a muscle adaptations to exercise. Beyond RONS, exercise adap-
hydrogen atom to a free radical [205], whereas polyphenols tations are also regulated by contraction-induced changes in
 A. Naderi et al.

Table 1  Summary of the evidence for the effects of supplements and nutrients on recovery and subsequent exercise performance
Supplement/nutrient Commentary

Carbohydrates 1–1.2 g·kg–1·h–1 of carbohydrate is recommended for initial 4-h post-exercise period with glucose and fructose in a
2:1 ratio
Most of the studies used 17–20% carbohydrate solutions
Small and more frequent doses are recommended
Fats No effect on subsequent exercise performance
Co-ingestion of fat and carbohydrates dose not impair glycogen resynthesis
Some signaling markers may be attenuated by very high fat intake
Micronutrients Effective in reducing exercise-induced muscle damage but no performance benefit
> 1000 mg of polyphenols per day optimize recovery between exercise bouts
Protein + carbohydrate 1–1.2 g·kg–1·h–1 of carbohydrate + 20–40 g highly digestive protein advised for effective muscle glycogen resynthesis
Beneficial effects may be due to higher muscle glycogen synthesis rates and/or positive nitrogen balance
Caffeine + carbohydrate 3–8 mg·kg–1 of caffeine + 1–1.2 g·kg–1·h–1 of carbohydrate is recommended
It may reduce muscle soreness
Little evidence shows that it may speed up muscle glycogen resynthesis more than carbohydrate alone
Creatine + carbohydrate 5–20 g·day–1 of creatine monohydrate + high carbohydrate diet may be more beneficial than carbohydrate alone for
muscle glycogen resynthesis
Hydration Replacement of 150–200% of lost sweat within 4–8 h is advised for effective rehydration
Oral rehydration solutions, milk, and orange juice are better than water
Cola, tea, coffee, lager, mineral water, and sports drinks are no different from water
There is little evidence to suggest that glycerol improves subsequent exercise performance
Sodium bicarbonate 0.3–0.4 g·kg–1 of sodium bicarbonate is recommended
It may improve subsequent exercise performance if recovery time is 40 min or less
New forms of sodium bicarbonate, such as enteric-coated and delayed-release capsules, are recommended to reduce
potential gastrointestinal discomfort

Table 2  Examples of post-exercise nutritional recommendations to enhance short-term recovery

Supplement/nutrient Less than 4 h between exercise bouts 4–24 h between exercise bouts

Carbohydrate 1–1.2 g·kg–1·h–1 1–1.2 g·kg–1·h–1 for first 4 h, followed by an individualized approach to
Small and more frequent doses are recommended achieve adequate glycogen restoration
Protein 20–40 g if there is adequate time for digestion 20–40 g immediately following exercise, followed by a total daily
intake of 1.2–1.8 g·kg–1 for endurance athletes and intake of 1.2–1.8
g·kg–1 for strength power athletes
–1
Caffeine 3–8 mg·kg 3–6 mg·kg–1 ~ 1 h prior to the subsequent exercise bout
Sodium bicarbonate 0.3–0.4 g·kg following the first bout of exercise 0.3–0.4 g·kg–1 following exercise and/or 2 h before the subsequent
–1

exercise bout
Fluid 150% body mass loss from exercise 150% body mass loss from exercise
Include electrolytes if food is not also consumed Include electrolytes if food is not also consumed
Can assess hydration status through urine color, frequency, and volume
or by measuring urine specific gravity and osmolality

mechanical strain, adenosine triphosphate (ATP) turnover, cal- per day either from food sources or supplements for at least
cium flux, redox balance, and intracellular oxygen pressure 3 days [203].
[215]. As an alternative to high-dose antioxidant supplementa-
tion, whole fruits and fruit-derived juices or extracts can also
provide a good source of antioxidants as well as micronutrients 5 Limitations and Future Directions
[216]. It is possible that interactions may occur when con-
suming between multiple sources of polyphenols, but practical Despite significant advancements in understanding nutri-
research is challenged by a large number of active compounds tional strategies for post-exercise recovery, several limita-
in a whole-food diet and the need for longer-term studies to tions within the current body of research should be acknowl-
see meaningful effects. To optimize recovery between exercise edged. First, much of the existing literature is based on
bouts, it is recommended to consume > 1000 mg polyphenols studies with male athletes, leading to a potential sex bias
Nutritional Strategies to Improve Post-exercise Recovery and Subsequent Exercise Performance

in the recommendations—a common problem within the engaging in training sessions with low carbohydrate avail-
sports science literature [217]. This disparity underscores ability to maximize training adaptations. However, caution
the need for more studies involving female athletes to ensure is advised owing to the increased risk of sleep disturbances
that recovery strategies are effective across the sexes. associated with high caffeine intake. Post-exercise creatine
Another limitation is the variability in exercise protocols and carbohydrate co-ingestion may enhance muscle gly-
used across studies, which complicates the generalization of cogen resynthesis but their role in improving subsequent
findings. For instance, research on carbohydrate replenish- exercise performance remains to be fully investigated.
ment often employs controlled glycogen-depleting protocols Milk-based formulations, sports drinks, oral rehydration
that may not reflect real-world exercise conditions when car- solutions, and water, along with solid meals, may opti-
bohydrates are also ingested during exercise or protocols are mize post-exercise rehydration and subsequent exercise
aimed at specific sports disciplines, and it is questionable performance. Post-exercise ­NaHCO3 supplementation may
how the findings translate into other disciplines. In addition, enhance acid–base balance restoration and improve sub-
the response to various recovery strategies, such as creatine sequent exercise performance, provided there is at least
and caffeine co-ingestion, appears to be highly individual- 30–90 min of recovery time between exercise bouts.
ized. Factors such as training status, genetic predisposition,
Supplementary Information The online version contains supplemen-
and diet playing key roles, suggesting a need for more per- tary material available at https://d​ oi.o​ rg/1​ 0.1​ 007/s​ 40279-0​ 25-0​ 2213-6.
sonalized approaches to recovery nutrition. However, lit-
erature on personalization is currently lacking. Moreover, Declarations
it is important to consider the interactions between different
performance and recovery-enhancing supplements, as they Funding No sources of funding were received to write this article.
often do not result in additive improvements [218]. Under-
standing these interactions is crucial for developing effective Conflict of interest Tim Podlogar has offered advice to RedBull
BORA hansgrohe Professional Cycling Team and Nduranz sports nu-
and safe nutritional strategies. Given that athletes such as trition brand. The other authors declare that they have no conflicts of
bodybuilders and powerlifters often engage in two training interest relevant to this paper.
sessions per day, future research is needed to explore this
training strategy. Such studies should focus on individuals Author contributions A.N., JR., HS., AH., M.S.K., A.G., E.B., and
T.P. wrote the first draft of the review; A.N., T.P., and J.R. revised the
involved in resistance training to assess their training quality original review. All authors have read and agreed to the published ver-
and muscle adaptation variables. Finally, a greater emphasis sion of the manuscript.
on the practical application of these strategies in real-world
sports contexts, including team sports and ultra-endurance Open Access This article is licensed under a Creative Commons Attri-
events, would help bridge the gap between laboratory find- bution 4.0 International License, which permits use, sharing, adapta-
ings and field performance. Future research should focus on tion, distribution and reproduction in any medium or format, as long
as you give appropriate credit to the original author(s) and the source,
translating experimental results into practical guidelines that provide a link to the Creative Commons licence, and indicate if changes
athletes and coaches can readily implement. were made. The images or other third party material in this article are
included in the article’s Creative Commons licence, unless indicated
otherwise in a credit line to the material. If material is not included in
the article’s Creative Commons licence and your intended use is not
6 Conclusions permitted by statutory regulation or exceeds the permitted use, you will
need to obtain permission directly from the copyright holder. To view a
Table 1 summarises the evidence regarding the nutrition copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
recommendations for optimizing post-exercise recov-
ery. Table 2 provides recommendations on the basis of
the duration between exercise bouts. Restoration of car- References
bohydrate stores within the body (i.e., muscle and liver
glycogen) appears to play the most important role in pro- 1. Kenttä G, Hassmén P. Overtraining and recovery. A conceptual
model. Sports Med. 1998;26(1):1–16.
moting recovery, followed by a sufficient protein intake 2. Kellmann M. Preventing overtraining in athletes in high-inten-
to promote tissue regeneration and adaptive responses to sity sports and stress/recovery monitoring. Scand J Med Sci
exercise. While the intake of fats and micronutrients may Sports. 2010;20(Suppl 2):95–102.
also aid recovery, the evidence supporting their efficacy 3. Halson SL. Monitoring training load to understand fatigue in
athletes. Sports Med. 2014;44(Suppl 2):S139–47.
is not as strong as that for carbohydrates. Co-ingestion of 4. Mountjoy M, et al. 2023 International Olympic Committee’s
caffeine with carbohydrates may accelerate post-exercise (IOC) consensus statement on relative energy deficiency in
muscle glycogen resynthesis and can be beneficial for ath- sport (REDs). Br J Sports Med. 2023;57(17):1073–98.
letes competing multiple times in a single day or those 5. Hawley JA, et al. Maximizing cellular adaptation to endurance
exercise in skeletal muscle. Cell Metab. 2018;27(5):962–76.
 A. Naderi et al.

6. Burke LM, Hawley JA. Swifter, higher, stronger: what’s on the 28. Décombaz J, et al. Fructose and galactose enhance postexer-
menu? Science. 2018;362(6416):781–7. cise human liver glycogen synthesis. Med Sci Sports Exerc.
7. Bonilla DA, et al. The 4R’s framework of nutritional strategies 2011;43(10):1964–71.
for post-exercise recovery: a review with emphasis on new 29. Alghannam AF, et al. Impact of muscle glycogen availability on
generation of carbohydrates. Int J Environ Res Public Health. the capacity for repeated exercise in man. Med Sci Sports Exerc.
2021;18(1):103. 2016;48(1):123–31.
8. Fuchs CJ, Gonzalez JT, van Loon LJC. Fructose co-ingestion 30. McCarthy DG, Spriet LL. Performance effects of carbohydrate
to increase carbohydrate availability in athletes. J Physiol. ingestion between bouts of intense aerobic interval exercise. Int
2019;597(14):3549–60. J Sports Physiol Perform. 2020;15(2):262–7.
9. Ivy JL. Regulation of muscle glycogen repletion, muscle pro- 31. Naderi A, et al. Carbohydrates and endurance exercise: a narra-
tein synthesis and repair following exercise. J Sports Sci Med. tive review of a food first approach. Nutrients. 2023;15(6):1367.
2004;3(3):131–8. 32. Burke LM, van Loon LJC, Hawley JA. Postexercise muscle glyco-
10. Ivy JL, Kuo CH. Regulation of GLUT4 protein and glycogen gen resynthesis in humans. J Appl Physiol. 2017;122(5):1055–67.
synthase during muscle glycogen synthesis after exercise. Acta 33. Maunder E, Podlogar T, Wallis GA. Postexercise fructose-malto-
Physiol Scand. 1998;162(3):295–304. dextrin ingestion enhances subsequent endurance capacity. Med
11. Fournier PA, et al. Post-exercise muscle glycogen repletion Sci Sports Exerc. 2018;50(5):1039–45.
in the extreme: effect of food absence and active recovery. J 34. Gray EA, et al. Postexercise glucose-fructose coingestion aug-
Sports Sci Med. 2004;3(3):139–46. ments cycling capacity during short-term and overnight recovery
12. Thomas DT, Erdman KA, Burke LM. American College of from exhaustive exercise, compared with isocaloric glucose. Int
Sports Medicine joint position statement. Nutrition and athletic J Sport Nutr Exerc Metab. 2020;30(1):54–61.
performance. Med Sci Sports Exerc. 2016;48(3):543–68. 35. Podlogar T, Wallis GA. Impact of post-exercise fructose-malto-
13. Ivy JL, et al. Muscle glycogen synthesis after exercise: effect dextrin ingestion on subsequent endurance performance. Front
of time of carbohydrate ingestion. J Appl Physiol (1985). Nutr. 2020;7:82.
1988;64(4):1480–5. 36. Hengist A, et al. The effects of glucose-fructose co-ingestion
14. Díaz-Lara J, et al. Delaying post-exercise carbohydrate intake on repeated performance during a day of intensified rugby
impairs next-day exercise capacity but not muscle glycogen union training in professional academy players. J Sports Sci.
or molecular responses. Acta Physiol (Oxf). 2024;240(10): 2021;39(10):1144–52.
e14215. 37. Naderi A, et al. Effects of two different doses of carbohydrate
15. Burke LM, et al. Carbohydrates for training and competition. J ingestion on taekwondo-related performance during a simulated
Sports Sci. 2011;29(Suppl 1):S17-27. tournament. J Int Soc Sports Nutr. 2021;18(1):40.
16. Gonzalez JT, Betts JA. Dietary sugars, exercise and hepatic car- 38. Ciok J, Dolna A. The role of glycemic index concept in carbohy-
bohydrate metabolism. Proc Nutr Soc. 2019;78(2):246–56. drate metabolism. Przegl Lek. 2006;63(5):287–91.
17. Murray B, Rosenbloom C. Fundamentals of glycogen metabo- 39. Wong SH, et al. Effect of glycemic index meals on recov-
lism for coaches and athletes. Nutr Rev. 2018;76(4):243–59. ery and subsequent endurance capacity. Int J Sports Med.
18. Podlogar T, Wallis GA. New horizons in carbohydrate 2009;30(12):898–905.
research and application for endurance athletes. Sports Med. 40. Erith S, et al. The effect of high carbohydrate meals with different
2022;52(Suppl 1):5–23. glycemic indices on recovery of performance during prolonged
19. Blom PC, et al. Effect of different post-exercise sugar diets on intermittent high-intensity shuttle running. Int J Sport Nutr Exerc
the rate of muscle glycogen synthesis. Med Sci Sports Exerc. Metab. 2006;16(4):393–404.
1987;19(5):491–6. 41. Stevenson E, et al. Influence of the glycaemic index of an even-
20. Van Den Bergh AJ, et al. Muscle glycogen recovery after exercise ing meal on substrate oxidation following breakfast and dur-
during glucose and fructose intake monitored by 13C-NMR. J ing exercise the next day in healthy women. Eur J Clin Nutr.
Appl Physiol (1985). 1996;81(4):1495–500. 2008;62(5):608–16.
21. Wallis GA, et al. Postexercise muscle glycogen synthesis with 42. Brown LJ, et al. High versus low glycemic index 3-h recovery
combined glucose and fructose ingestion. Med Sci Sports Exerc. diets following glycogen-depleting exercise has no effect on sub-
2008;40(10):1789–94. sequent 5-km cycling time trial performance. J Sci Med Sport.
22. Trommelen J, et al. Fructose coingestion does not accelerate 2013;16(5):450–4.
postexercise muscle glycogen repletion. Med Sci Sports Exerc. 43. Stevenson E, Williams C, Nute M. The influence of the glycae-
2016;48(5):907–12. mic index of breakfast and lunch on substrate utilisation dur-
23. Casey A, et al. Effect of carbohydrate ingestion on glycogen ing the postprandial periods and subsequent exercise. Br J Nutr.
resynthesis in human liver and skeletal muscle, measured by (13) 2005;93(6):885–93.
C MRS. Am J Physiol Endocrinol Metab. 2000;278(1):E65-75. 44. Jeukendrup AE, Gleeson M. Sport nutrition. 4th ed. Champaign:
24. Podlogar T, et al. Postexercise muscle glycogen synthesis with Human Kinetics; 2025.
glucose, galactose, and combined galactose-glucose ingestion. 45. Gonzalez JT, Wallis GA. Carb-conscious: the role of carbo-
Am J Physiol Endocrinol Metab. 2023;325(6):E672–81. hydrate intake in recovery from exercise. Curr Opin Clin Nutr
25. Fuchs CJ, et al. Sucrose ingestion after exhaustive exercise Metab Care. 2021;24(4):364–71.
accelerates liver, but not muscle glycogen repletion compared 46. Noakes TD. What is the evidence that dietary macronutri-
with glucose ingestion in trained athletes. J Appl Physiol. ent composition influences exercise performance? A narrative
2016;120(11):1328–34. review. Nutrients. 2022;14(4):862.
26. Gonzalez JT, et al. Liver glycogen metabolism during and after 47. Gonzalez JT, et al. Glucose plus fructose ingestion for post-
prolonged endurance-type exercise. Am J Physiol Endocrinol exercise recovery-greater than the sum of its parts? Nutrients.
Metab. 2016;311(3):E543–53. 2017;9(4):344.
27. Burke LM, Collier GR, Hargreaves M. Muscle glycogen storage 48. Coyle EF, et al. Muscle glycogen utilization during prolonged
after prolonged exercise: effect of the glycemic index of carbo- strenuous exercise when fed carbohydrate. J Appl Physiol (1985).
hydrate feedings. J Appl Physiol (1985). 1993;75(2):1019–23. 1986;61(1):165–72.
Nutritional Strategies to Improve Post-exercise Recovery and Subsequent Exercise Performance

49. Gonzalez JT, et al. Ingestion of glucose or sucrose prevents liver and duration in vivo in humans. Cell Rep Med. 2023;4(12):
but not muscle glycogen depletion during prolonged endurance- 101324.
type exercise in trained cyclists. Am J Physiol Endocrinol Metab. 69. Tarnopolsky M. Protein requirements for endurance athletes.
2015;309(12):E1032–9. Nutrition. 2004;20(7–8):662–8.
50. Jeukendrup AE, et al. Carbohydrate ingestion can completely 70. Kato H, et al. Protein requirements are elevated in endurance
suppress endogenous glucose production during exercise. Am J athletes after exercise as determined by the indicator amino acid
Physiol. 1999;276(4):E672–83. oxidation method. PLoS ONE. 2016;11(6): e0157406.
51. Moore DR, et al. Beyond muscle hypertrophy: why dietary 71. Bandegan A, et al. Indicator amino acid oxidation protein
protein is important for endurance athletes. Appl Physiol Nutr requirement estimate in endurance-trained men 24 h postexer-
Metab. 2014;39(9):987–97. cise exceeds both the EAR and current athlete guidelines. Am J
52. Doering TM, et al. Postexercise dietary protein strategies to Physiol Endocrinol Metab. 2019;316(5):E741-e748.
maximize skeletal muscle repair and remodeling in masters 72. Wooding DJ, et al. Increased protein requirements in female
endurance athletes: a review. Int J Sport Nutr Exerc Metab. athletes after variable-intensity exercise. Med Sci Sports Exerc.
2016;26(2):168–78. 2017;49(11):2297–304.
53. Pasiakos SM, Lieberman HR, McLellan TM. Effects of protein 73. Chapman S, et al. Does protein supplementation support adap-
supplements on muscle damage, soreness and recovery of muscle tations to arduous concurrent exercise training? A systematic
function and physical performance: a systematic review. Sports review and meta-analysis with military based applications. Nutri-
Med. 2014;44(5):655–70. ents. 2021;13(5):1416.
54. Tipton KD, Wolfe RR. Protein and amino acids for athletes. J 74. West S, et al. Nutritional considerations for the vegan athlete.
Sports Sci. 2004;22(1):65–79. Adv Nutr. 2023;14(4):774–95.
55. Areta JL, et al. Timing and distribution of protein ingestion dur- 75. Res PT, et al. Protein ingestion before sleep improves postexercise
ing prolonged recovery from resistance exercise alters myofibril- overnight recovery. Med Sci Sports Exerc. 2012;44(8):1560–9.
lar protein synthesis. J Physiol. 2013;591(9):2319–31. 76. Trommelen J, et al. Presleep dietary protein-derived amino
56. Churchward-Venne TA, Burd NA, Phillips SM. Nutritional regu- acids are incorporated in myofibrillar protein during postex-
lation of muscle protein synthesis with resistance exercise: strate- ercise overnight recovery. Am J Physiol Endocrinol Metab.
gies to enhance anabolism. Nutr Metab (Lond). 2012;9(1):40. 2018;314(5):E457-e467.
57. Macnaughton LS, et al. The response of muscle protein syn- 77. Valenzuela PL, et al. Pre-sleep protein supplementation in profes-
thesis following whole-body resistance exercise is greater fol- sional cyclists during a training camp: a three-arm randomized
lowing 40 g than 20 g of ingested whey protein. Physiol Rep. controlled trial. J Int Soc Sports Nutr. 2023;20(1):2166366.
2016;4(15):e12893. 78. Ormsbee MJ, et al. Pre-sleep protein supplementation after an
58. Moore DR, et al. Ingested protein dose response of muscle and acute bout of evening resistance exercise does not improve next
albumin protein synthesis after resistance exercise in young men. day performance or recovery in resistance trained men. J Int Soc
Am J Clin Nutr. 2009;89(1):161–8. Sports Nutr. 2022;19(1):164–78.
59. Aragon AA, Schoenfeld BJ. Nutrient timing revisited: is 79. Wilkinson K, et al. Association of postprandial postexercise
there a post-exercise anabolic window? J Int Soc Sports Nutr. muscle protein synthesis rates with dietary leucine: a systematic
2013;10(1):5. review. Physiol Rep. 2023;11(15): e15775.
60. Schoenfeld BJ, Aragon AA, Krieger JW. The effect of protein 80. Phillips SM. The impact of protein quality on the promotion of
timing on muscle strength and hypertrophy: a meta-analysis. J resistance exercise-induced changes in muscle mass. Nutr Metab
Int Soc Sports Nutr. 2013;10(1):53. (Lond). 2016;13:64.
61. Trommelen J, Holwerda AM, van Loon LJC. Protein intake dis- 81. Tang JE, et al. Ingestion of whey hydrolysate, casein, or soy
tribution: beneficial, detrimental, or inconsequential for muscle protein isolate: effects on mixed muscle protein synthesis at rest
anabolism? Response to Witard & Mettler. Int J Sport Nutr Exerc and following resistance exercise in young men. J Appl Physiol
Metab. 2024;34(5):325–8. (1985). 2009;107(3):987–92.
62. Sollie O, et al. Protein intake in the early recovery period after 82. Monteyne AJ, et al. Vegan and omnivorous high protein diets
exhaustive exercise improves performance the following day. J support comparable daily myofibrillar protein synthesis rates
Appl Physiol (1985). 2018;125(6):1731–42. and skeletal muscle hypertrophy in young adults. J Nutr.
63. Dahl MA, et al. Coingestion of protein and carbohydrate in 2023;153(6):1680–95.
the early recovery phase, compared with carbohydrate only, 83. van der Heijden I, et al. Alternative dietary protein sources to
improves endurance performance despite similar glycogen deg- support healthy and active skeletal muscle aging. Nutr Rev.
radation and AMPK phosphorylation. J Appl Physiol (1985). 2023;81(2):206–30.
2020;129(2):297–310. 84. Alghannam AF, Gonzalez JT, Betts JA. Restoration of muscle
64. West DW, et al. Rapid aminoacidemia enhances myofibrillar pro- glycogen and functional capacity: role of post-exercise carbohy-
tein synthesis and anabolic intramuscular signaling responses drate and protein co-ingestion. Nutrients. 2018;10(2):253.
after resistance exercise. Am J Clin Nutr. 2011;94(3):795–803. 85. Margolis LM, et al. Coingestion of carbohydrate and protein on
65. Thomson JS, Ali A, Rowlands DS. Leucine-protein supplemented muscle glycogen synthesis after exercise: a meta-analysis. Med
recovery feeding enhances subsequent cycling performance in Sci Sports Exerc. 2021;53(2):384–93.
well-trained men. Appl Physiol Nutr Metab. 2011;36(2):242–53. 86. Craven J, et al. The effect of consuming carbohydrate with and
66. Loenneke JP, et al. Per meal dose and frequency of protein con- without protein on the rate of muscle glycogen re-synthesis dur-
sumption is associated with lean mass and muscle performance. ing short-term post-exercise recovery: a systematic review and
Clin Nutr. 2016;35(6):1506–11. meta-analysis. Sports Med Open. 2021;7(1):9.
67. Moore DR, et al. Daytime pattern of post-exercise protein intake 87. McCartney D, Desbrow B, Irwin C. Post-exercise ingestion of
affects whole-body protein turnover in resistance-trained males. carbohydrate, protein and water: a systematic review and meta-
Nutr Metab (Lond). 2012;9(1):91. analysis for effects on subsequent athletic performance. Sports
68. Trommelen J, et al. The anabolic response to protein ingestion Med. 2017;48(2):379–408.
during recovery from exercise has no upper limit in magnitude
 A. Naderi et al.

88. Christensen PM, et al. Caffeine and bicarbonate for speed. development of a beverage hydration index. Am J Clin Nutr.
A meta-analysis of legal supplements potential for improv- 2016;103(3):717–23.
ing intense endurance exercise performance. Front Physiol. 110. Borra V, et al. Oral rehydration beverages for treating exercise-
2017;8:240. associated dehydration: a systematic review. Part I: carbohydrate-
89. Rustad PI, et al. Intake of protein plus carbohydrate during the electrolyte solutions. J Athl Train. 2025;60(1):34–54.
first two hours after exhaustive cycling improves performance 111. Ly NQ, Hamstra-Wright KL, Horswill CA. Post-exercise rehy-
the following day. PLoS ONE. 2016;11(4): e0153229. dration in athletes: effects of sodium and carbohydrate in com-
90. Goldstein E, et al. Carbohydrate-protein coingestion enhances mercial hydration beverages. Nutrients. 2023;15(22):4759.
cycling performance with minimal recovery time between bouts 112. Schleh MW, Dumke CL. Comparison of sports drink versus
of exhaustive intermittent exercise. J Exerc Nutr. 2022;5(2). oral rehydration solution during exercise in the heat. Wilderness
91. Sollie O, et al. Similar performance after intake of carbo- Environ Med. 2018;29(2):185–93.
hydrate plus whey protein and carbohydrate only in the early 113. Fan PW, Burns SF, Lee JKW. Efficacy of ingesting an oral rehy-
phase after non-exhaustive cycling. Scand J Med Sci Sports. dration solution after exercise on fluid balance and endurance
2023;33(7):1091–103. performance. Nutrients. 2020;12(12):3826.
92. Goldstein ER, et al. Carbohydrate-protein drink is effective for 114. Russo I, et al. Systematic literature review: the effect of dairy
restoring endurance capacity in masters class athletes after a two- milk on markers of recovery optimisation in response to endur-
hour recovery. J Int Soc Sports Nutr. 2023;20(1):2178858. ance exercise. Int J sports Sci. 2019:69–85.
93. Churchward-Venne TA, et al. Dose-response effects of dietary 115. De Brier N, et al. A systematic review on oral rehydration bev-
protein on muscle protein synthesis during recovery from endur- erages for treating exercise-associated dehydration, part II. The
ance exercise in young men: a double-blind randomized trial. Am effectiveness of alternatives to carbohydrate-electrolyte drinks.
J Clin Nutr. 2020;112(2):303–17. J Athl Train. 2025;60(1).
94. Hansen M, et al. Supplement with whey protein hydrolysate in 116. Lee JK, et al. Effects of ingesting a sports drink during exercise
contrast to carbohydrate supports mitochondrial adaptations in and recovery on subsequent endurance capacity. Eur J Sport Sci.
trained runners. J Int Soc Sports Nutr. 2020;17(1):46. 2011;11(2):77–86.
95. Huang WC, et al. Whey protein improves marathon-induced 117. McCartney D, et al. Fluid, energy, and nutrient recovery via
injury and exercise performance in elite track runners. Int J Med ad libitum intake of different commercial beverages and food in
Sci. 2017;14(7):648–54. female athletes. Appl Physiol Nutr Metab. 2019;44(1):37–46.
96. Leiper JB. Intestinal water absorption—implications for 118. McCartney D, et al. The effect of different post-exercise bev-
the formulation of rehydration solutions. Int J Sports Med. erages with food on ad libitum fluid recovery, nutrient provi-
1998;19(Suppl 2):S129–32. sion, and subsequent athletic performance. Physiol Behav.
97. Lorenzo I, Serra-Prat M, Yébenes JC. The role of water home- 2019;201:22–30.
ostasis in muscle function and frailty: a review. Nutrients. 119. Maughan RJ, Leiper JB, Shirreffs SM. Restoration of fluid bal-
2019;11(8):1857. ance after exercise-induced dehydration: effects of food and fluid
98. Maughan RJ, Shirreffs SM. Dehydration and rehydration in com- intake. Eur J Appl Physiol Occup Physiol. 1996;73(3–4):317–25.
petative sport. Scand J Med Sci Sports. 2010;20(Suppl 3):40–7. 120. Grgic J, et al. International Society of Sports Nutrition position
99. Baker LB. Sweating rate and sweat sodium concentration in ath- stand: sodium bicarbonate and exercise performance. J Int Soc
letes: a review of methodology and intra/interindividual vari- Sports Nutr. 2022;18(1):1.
ability. Sports Med. 2017;47(Suppl 1):111–28. 121. de Oliveira LF, et al. Extracellular buffering supplements
100. López-Torres O, et al. Hydration, hyperthermia, glycogen, and to improve exercise capacity and performance: a compre-
recovery: crucial factors in exercise performance—a systematic hensive systematic review and meta-analysis. Sports Med.
review and meta-analysis. Nutrients. 2023;15(20):4442. 2022;52(3):505–26.
101. Judge LW, et al. Hydration to maximize performance and recov- 122. Bouclin R, Charbonneau E, Renaud JM. N ­ a+ and K­ + effect on
ery: knowledge, attitudes, and behaviors among collegiate track contractility of frog sartorius muscle: implication for the mecha-
and field throwers. J Hum Kinet. 2021;79:111–22. nism of fatigue. Am J Physiol. 1995;268(6 Pt 1):C1528–36.
102. Singh R. Hydration strategies for exercise performance in hot 123. Jones RL, et al. Dose-response of sodium bicarbonate inges-
environment. Br J Sports Med. 2010;44(Suppl 1):i40.1-i40. tion highlights individuality in time course of blood analyte
103. Li C, et al. Predicting athletes’ pre-exercise fluid intake: a theo- responses. Int J Sport Nutr Exerc Metab. 2016;26(5):445–53.
retical integration approach. Nutrients. 2018;10(5):646. 124. Gurton WH, et al. Enhancing exercise performance and recov-
104. Maughan RJ. Fluid and electrolyte loss and replacement in ery through sodium bicarbonate supplementation: introduc-
exercise. J Sports Sci. 1991;9(Spec No):117–42. ing the ingestion recovery framework. Eur J Appl Physiol.
105. Maughan RJ, Shirreffs SM. Recovery from prolonged exer- 2024;124(11):3175–90.
cise: restoration of water and electrolyte balance. J Sports Sci. 125. Siegler JC, et al. The Hyperhydration potential of sodium
1997;15(3):297–303. bicarbonate and sodium citrate. Int J Sport Nutr Exerc Metab.
106. Shirreffs SM, Sawka MN. Fluid and electrolyte needs for train- 2022;32(2):74–81.
ing, competition, and recovery. J Sports Sci. 2011;29(Suppl 126. Siegler JC, et al. Pre-exercise alkalosis and acid-base recovery.
1):S39-46. Int J Sports Med. 2008;29(7):545–51.
107. McCartney D, Desbrow B, Irwin C. The effect of fluid intake 127. Pruscino CL, et al. Effects of sodium bicarbonate, caffeine, and
following dehydration on subsequent athletic and cognitive their combination on repeated 200-m freestyle performance. Int
performance: a systematic review and meta-analysis. Sports J Sport Nutr Exerc Metab. 2008;18(2):116–30.
Med Open. 2017;3(1):13. 128. Gough LA, et al. The influence of alkalosis on repeated high-
108. Jardine WT, et al. The effect of pre-exercise hyperhydration intensity exercise performance and acid–base balance recov-
on exercise performance, physiological outcomes and gas- ery in acute moderate hypoxic conditions. Eur J Appl Physiol.
trointestinal symptoms: a systematic review. Sports Med. 2018;118(12):2489–98.
2023;53(11):2111–34. 129. Gough LA, et al. Ingestion of sodium bicarbonate ­(NaHCO3) fol-
109. Maughan RJ, et al. A randomized trial to assess the poten- lowing a fatiguing bout of exercise accelerates postexercise acid-
tial of different beverages to affect hydration status: base balance recovery and improves subsequent high-intensity
Nutritional Strategies to Improve Post-exercise Recovery and Subsequent Exercise Performance

cycling time to exhaustion. Int J Sport Nutr Exerc Metab. 149. Low SY, Rennie MJ, Taylor PM. Modulation of glycogen syn-
2017;27(5):429–38. thesis in rat skeletal muscle by changes in cell volume. J Physiol.
130. Gough LA, et al. Post-exercise supplementation of sodium 1996;495(Pt 2):299–303.
bicarbonate improves acid base balance recovery and subse- 150. Roberts PA, et al. Creatine ingestion augments dietary carbohy-
quent high-intensity boxing specific performance. Front Nutr. drate mediated muscle glycogen supercompensation during the
2019;6:155. initial 24 h of recovery following prolonged exhaustive exercise
131. Gurton W, et al. Effects of post-exercise sodium bicarbonate in humans. Amino Acids. 2016;48(8):1831–42.
ingestion on acid-base balance recovery and time-to-exhaustion 151. Van Loon LJ, et al. Creatine supplementation increases glycogen
running performance: a randomised crossover trial in recreational storage but not GLUT-4 expression in human skeletal muscle.
athletes. Appl Physiol Nutr Metab. 2021;46(9):1111–8. Clin Sci (Lond). 2004;106(1):99–106.
132. Farias DEOL, et al. Is individualization of sodium bicarbonate 152. Nelson AG, et al. Muscle glycogen supercompensation is
ingestion based on time to peak necessary? Med Sci Sports enhanced by prior creatine supplementation. Med Sci Sports
Exerc. 2020;52(8):1801–8. Exerc. 2001;33(7):1096–100.
133. Peacock J, et al. Extracellular buffer choice influences acid- 153. Rico-Sanz J, et al. Creatine feeding does not enhance intramyo-
base responses and gastrointestinal symptoms. Res Sports Med. cellular glycogen concentration during carbohydrate loading:
2021;29(6):505–16. an in vivo study by 31P- and 13C-MRS. J Physiol Biochem.
134. Naderi A, et al. Timing, optimal dose and intake duration of 2008;64(3):189–96.
dietary supplements with evidence-based use in sports nutrition. 154. Tomcik KA, et al. Effects of creatine and carbohydrate load-
J Exerc Nutr Biochem. 2016;20(4):1–12. ing on cycling time trial performance. Med Sci Sports Exerc.
135. Gough LA, et al. The reproducibility of blood acid base 2018;50(1):141–50.
responses in male collegiate athletes following individualised 155. Greiwe JS, et al. Effects of endurance exercise training on mus-
doses of sodium bicarbonate: a randomised controlled crossover cle glycogen accumulation in humans. J Appl Physiol (1985).
study. Sports Med. 2017;47(10):2117–27. 1999;87(1):222–6.
136. Heibel AB, et al. Time to optimize supplementation: modify- 156. Forsberg AM, et al. Muscle composition in relation to age and
ing factors influencing the individual responses to extracellular sex. Clin Sci (Lond). 1991;81(2):249–56.
buffering agents. Front Nutr. 2018;5:35. 157. Grgic J, et al. Wake up and smell the coffee: caffeine supple-
137. Gough LA, et al. Post-exercise supplementation of sodium mentation and exercise performance-an umbrella review of 21
bicarbonate improves acid base balance recovery and subse- published meta-analyses. Br J Sports Med. 2020;54(11):681–8.
quent high-intensity boxing specific performance. Front Nutr. 158. Lopez-Seoane J, et al. Synergy of carbohydrate and caffeine
2019;6:155. ingestion on physical performance and metabolic responses to
138. Hilton NP, et al. Enteric-coated sodium bicarbonate attenu- exercise: a systematic review with meta-analysis. Crit Rev Food
ates gastrointestinal side-effects. Int J Sport Nutr Exerc Metab. Sci Nutr. 2024;64(10): 2941–59.
2020;30(1):62–8. 159. Chen HY, et al. Effects of caffeine and sex on muscle perfor-
139. Hilton NP, et al. A novel ingestion strategy for sodium bicarbo- mance and delayed-onset muscle soreness after exercise-induced
nate supplementation in a delayed-release form: a randomised muscle damage: a double-blind randomized trial. J Appl Physiol
crossover study in trained males. Sports Med Open. 2019;5(1):4. (1985). 2019;127(3):798–805.
140. Gough LA, Sparks SA. The effects of a carbohydrate hydrogel 160. Pedersen DJ, et al. High rates of muscle glycogen resynthesis
system for the delivery of bicarbonate mini-tablets on acid-base after exhaustive exercise when carbohydrate is coingested with
buffering and gastrointestinal symptoms in resting well-trained caffeine. J Appl Physiol (1985). 2008;105(1):7–13.
male cyclists. Sports Med Open. 2024;10(1):17. 161. Loureiro LMR, Reis CEG, da Costa THM. Effects of coffee com-
141. Gough LA, et al. The effects of sodium bicarbonate ingestion ponents on muscle glycogen recovery: a systematic review. Int J
on cycling performance and acid base balance recovery in acute Sport Nutr Exerc Metab. 2018;28(3):284–93.
normobaric hypoxia. J Sports Sci. 2019;37(13):1464–71. 162. Thong FSL, et al. Caffeine-induced impairment of insulin action
142. Terjung RL, et al. American College of Sports Medicine round- but not insulin signaling in human skeletal muscle is reduced by
table. The physiological and health effects of oral creatine sup- exercise. Diabetes. 2002;51(3):583–90.
plementation. Med Sci Sports Exerc. 2000;32(3):706–17. 163. Loureiro LMR, et al. Effect of milk on muscle glycogen recovery
143. Forbes SC, et al. Creatine supplementation and endurance perfor- and exercise performance: a systematic review. Strength Cond J.
mance: surges and sprints to win the race. J Int Soc Sports Nutr. 2021;43(4):43–52.
2023;20(1):2204071. 164. Battram DS, et al. Caffeine ingestion does not impede the
144. Robinson TM, et al. Role of submaximal exercise in promoting resynthesis of proglycogen and macroglycogen after prolonged
creatine and glycogen accumulation in human skeletal muscle. J exercise and carbohydrate supplementation in humans. J Appl
Appl Physiol (1985). 1999;87(2):598–604. Physiol. 2004;96(3):943–50.
145. Op ‘t Eijnde B, et al. Effect of oral creatine supplementation 165. Beelen M, et al. Impact of caffeine and protein on postex-
on human muscle GLUT4 protein content after immobilization. ercise muscle glycogen synthesis. Med Sci Sports Exerc.
Diabetes. 2001;50(1):18–23. 2012;44(4):692–700.
146. Wax B, et al. Creatine for exercise and sports performance, 166. Loureiro LMR, et al. Coffee increases post-exercise muscle
with recovery considerations for healthy populations. Nutrients. glycogen recovery in endurance athletes: a randomized clinical
2021;13(6):1915. trial. Nutrients. 2021;13(10):3335.
147. O’Reilly KP, et al. Eccentric exercise-induced muscle dam- 167. Taylor C, et al. The effect of adding caffeine to postexercise
age impairs muscle glycogen repletion. J Appl Physiol (1985). carbohydrate feeding on subsequent high-intensity interval-
1987;63(1):252–6. running capacity compared with carbohydrate alone. Int J
148. Sobolewski EJ, et al. The physiological effects of creatine Sport Nutr Exerc Metab. 2011;21(5):410–6.
supplementation on hydration: a review. Am J Lifestyle Med. 168. Andrade-Souza VA, et al. Effects of isolated or combined car-
2011;5(4):320–7. bohydrate and caffeine supplementation between 2 daily train-
ing sessions on soccer performance. Appl Physiol Nutr Metab.
2015;40(5):457–63.
 A. Naderi et al.

169. Barzegar H, et al. Caffeine coingested with carbohydrate on 189. Burke LM, et al. Effect of coingestion of fat and protein with
performance recovery in national-level paddlers: a randomized, carbohydrate feedings on muscle glycogen storage. J Appl
double-blind, crossover, placebo-controlled trial. J Sports Med Physiol (1985). 1995;78(6):2187–92.
Phys Fit. 2022;62(3):337–42. 190. Hammond KM, et al. Postexercise high-fat feeding suppresses
170. Negaresh R, et al. Effects of different dosages of caffeine p70S6K1 activity in human skeletal muscle. Med Sci Sports
administration on wrestling performance during a simulated Exerc. 2016;48(11):2108–17.
tournament. Eur J Sport Sci. 2019;19(4):499–507. 191. Bell PG, et al. Montmorency cherries reduce the oxidative
171. Impey SG, et al. Fuel for the work required: a theoretical stress and inflammatory responses to repeated days high-
framework for carbohydrate periodization and the glycogen intensity stochastic cycling. Nutrients. 2014;6(2):829–43.
threshold hypothesis. Sports Med. 2018;48(5):1031–48. 192. Clifford T, et al. Effects of beetroot juice on recovery of muscle
172. De Souza JG, et al. Risk or benefit? Side effects of caffeine function and performance between bouts of repeated sprint
supplementation in sport: a systematic review. Eur J Nutr. exercise. Nutrients. 2016;8(8):506.
2022;61(8):3823–34. 193. Machin DR, et al. Effects of differing dosages of pomegran-
173. Gardiner C, et al. The effect of caffeine on subsequent sleep: a ate juice supplementation after eccentric exercise. Physiol J.
systematic review and meta-analysis. Sleep Med Rev. 2023;69: 2014;2014(1):271959.
101764. 194. McLeay Y, et al. Effect of New Zealand blueberry consumption
174. Van Loon LJ. Use of intramuscular triacylglycerol as a sub- on recovery from eccentric exercise-induced muscle damage.
strate source during exercise in humans. J Appl Physiol (1985). J Int Soc Sports Nutr. 2012;9(1):19.
2004;97(4):1170–87. 195. Martínez-Sánchez A, et al. Biochemical, physiological,
175. Romijn JA, et al. Regulation of endogenous fat and carbohy- and performance response of a functional watermelon juice
drate metabolism in relation to exercise intensity and duration. enriched in l-citrulline during a half-marathon race. Food Nutr
Am J Physiol. 1993;265(3 Pt 1):E380–91. Res. 2017.
176. Kiens B, Richter EA. Utilization of skeletal muscle triacylg- 196. Kim J, So WY. Effects of acute grape seed extract supplementa-
lycerol during postexercise recovery in humans. Am J Physiol. tion on muscle damage after eccentric exercise: a randomized,
1998;275(2):E332–7. controlled clinical trial. J Exerc Sci Fit. 2019;17(2):77–9.
177. Vogt M, et al. Effects of dietary fat on muscle substrates, 197. Hutchison AT, et al. Black currant nectar reduces muscle damage
metabolism, and performance in athletes. Med Sci Sports and inflammation following a bout of high-intensity eccentric
Exerc. 2003;35(6):952–60. contractions. J Diet Supp. 2016;13(1):1–15.
178. Starling RD, et al. Effects of diet on muscle triglycer- 198. McBride JM, et al. Effect of resistance exercise on free radical
ide and endurance performance. J Appl Physiol (1985). production. Med Sci Sports Exerc. 1998;30(1):67–72.
1997;82(4):1185–9. 199. McFarlin BK, et al. Reduced inflammatory and muscle damage
179. Knudsen CB, et al. No net-utilization of intramuscular lipid biomarkers following oral supplementation with bioavailable
droplets during repeated high-intensity intermittent exercise. curcumin. BBA Clin. 2016;5:72–8.
Am J Physiol Endocrinol Metab. 2023;325(6): E700–10. 200. Morgan PT, et al. Flavanol-rich cacao mucilage juice enhances
180. Van Loon LJ, et al. Influence of prolonged endurance recovery of power but not strength from intensive exercise in
cycling and recovery diet on intramuscular triglyceride healthy, young men. Sports (Basel). 2018;6(4):159.
content in trained males. Am J Physiol Endocrinol Metab. 201. Barker T, et al. Supplemental vitamin D enhances the recovery
2003;285(4):E804–11. in peak isometric force shortly after intense exercise. Nutr Metab
181. Lundsgaard AM, Fritzen AM, Kiens B. The importance of (Lond). 2013;10(1):69.
fatty acids as nutrients during post-exercise recovery. Nutri- 202. Bongiovanni T, et al. Nutritional interventions for reducing
ents. 2020;12(2):280. the signs and symptoms of exercise-induced muscle damage
182. Owens DJ, et al. Exercise-induced muscle damage: what is it, and accelerate recovery in athletes: current knowledge, prac-
what causes it and what are the nutritional solutions? Eur J tical application and future perspectives. Eur J Appl Physiol.
Sport Sci. 2019;19(1):71–85. 2020;120(9):1965–96.
183. McGlory C, et al. Temporal changes in human skeletal 203. Bowtell J, Kelly V. Fruit-derived polyphenol supplementation for
muscle and blood lipid composition with fish oil sup- athlete recovery and performance. Sports Med. 2019;49(Suppl
plementation. Prostaglandins Leukot Essent Fatty Acids. 1):3–23.
2014;90(6):199–206. 204. Moopanar TR, Allen DG. Reactive oxygen species reduce myofi-
184. Tsao JP, et al. Oral conjugated linoleic acid supplementation brillar ­Ca2+ sensitivity in fatiguing mouse skeletal muscle at 37
enhanced glycogen resynthesis in exercised human skeletal C. J Physiol. 2005;564(1):189–99.
muscle. J Sports Sci. 2015;33(9):915–23. 205. Nikolaidis MG, et al. Does vitamin C and E supplementation
185. Kim Y, et al. Impact of conjugated linoleic acid (CLA) on impair the favorable adaptations of regular exercise? Oxid Med
skeletal muscle metabolism. Lipids. 2016;51(2):159–78. Cell Longev. 2012;2012:707941.
186. Ryder J, et al. Isomer-specific antidiabetic properties of con- 206. Somerville V, Bringans C, Braakhuis A. Polyphenols and per-
jugated linoleic acid: improved glucose tolerance, skeletal formance: a systematic review and meta-analysis. Sports Med.
muscle insulin action, and UCP-2 gene expression. Diabetes. 2017;47(8):1589–99.
2001;50(5):1149–57. 207. Tsao JP, et al. Short-term oral quercetin supplementation
187. Cunningham K, Read N. The effect of incorporating fat into improves post-exercise insulin sensitivity, antioxidant capacity
different components of a meal on gastric emptying and and enhances subsequent cycling time to exhaustion in healthy
postprandial blood glucose and insulin responses. Br J Nutr. adults: a pilot study. Front Nutr. 2022;9: 875319.
1989;61(2):285–90. 208. Tsai TW, et al. Effect of green tea extract supplementation on
188. Roy BD, Tarnopolsky MA. Influence of differing macronutri- glycogen replenishment in exercised human skeletal muscle. Br
ent intakes on muscle glycogen resynthesis after resistance J Nutr. 2017;117(10):1343–50.
exercise. J Appl Physiol (1985). 1998;84(3):890–6. 209. Pingitore A, et al. Exercise and oxidative stress: potential
effects of antioxidant dietary strategies in sports. Nutrition.
2015;31(7–8):916–22.
Nutritional Strategies to Improve Post-exercise Recovery and Subsequent Exercise Performance

210. Rothschild JA, Bishop DJ. Effects of dietary supple- 215. Egan B, Zierath JR. Exercise metabolism and the molecu-
ments on adaptations to endurance training. Sports Med. lar regulation of skeletal muscle adaptation. Cell Metab.
2020;50(1):25–53. 2013;17(2):162–84.
211. Paulsen G, et al. Vitamin C and E supplementation hampers cel- 216. Naderi A, et al. Fruit for sport. Trends Food Sci Technol.
lular adaptation to endurance training in humans: a double-blind, 2018;74:85–98.
randomised, controlled trial. J Physiol. 2014;592(8):1887–901. 217. Elliott-Sale KJ, et al. Methodological considerations for stud-
212. Ristow M, et al. Antioxidants prevent health-promoting effects ies in sport and exercise science with women as participants: a
of physical exercise in humans. Proc Natl Acad Sci USA. working guide for standards of practice for research on women.
2009;106(21):8665–70. Sports Med. 2021;51(5):843–61.
213. Gomez-Cabrera MC, et al. Oral administration of vitamin C 218. Burke LM. Practical issues in evidence-based use of performance
decreases muscle mitochondrial biogenesis and hampers train- supplements: supplement interactions, repeated use and individ-
ing-induced adaptations in endurance performance. Am J Clin ual responses. Sports Med. 2017;47(Suppl 1):79–100.
Nutr. 2008;87(1):142–9.
214. Schwarz NA, et al. (-)-Epicatechin supplementation inhibits
aerobic adaptations to cycling exercise in humans. Front Nutr.
2018;5:132.
Authors and Affiliations

Alireza Naderi1 · Jeffrey A. Rothschild2,3 · Heitor O. Santos4 · Amin Hamidvand5 · Majid S. Koozehchian6 ·
Abdolrahman Ghazzagh7 · Erfan Berjisian8 · Tim Podlogar9,10

4
* Alireza Naderi Postgraduate Program, Faculdade UNIGUAÇU, Cascavel,
[email protected] PR, Brazil
5
* Tim Podlogar Department of Biological Sciences in Sport, Faculty of Sport
[email protected] Sciences and Health, Shahid Beheshti University, Tehran,
Iran
Heitor O. Santos
6
[email protected] Department of Kinesiology, Jacksonville State University,
Jacksonville, AL 36265, USA
Majid S. Koozehchian
7
[email protected] Department of Sport Sciences, Faculty of Humanities,
Tarbiat Modares University, Tehran, Iran
1
Department of Sport Physiology, Faculty of Human 8
School of Medical and Health Sciences, Edith Cowan
Sciences, Borujerd Branch, Islamic Azad University,
University, Joondalup, Western Australia, Australia
Borujerd, Iran
9
2 School of Sport, Exercise and Rehabilitation Sciences,
Sports Performance Research Institute New Zealand,
College of Life and Environmental Sciences, University
Auckland University of Technology, Auckland, New Zealand
of Birmingham, Birmingham, UK
3
High Performance Sport New Zealand, Auckland, 10
Department of Public Health and Sport Sciences, Medical
New Zealand
School, University of Exeter, Exeter, UK