nutrients

Review
Dietary Patterns, Gut Microbiota and Sports Performance in
Athletes: A Narrative Review
Yonglin Chen 1 , Keer Yang 1 , Mingxin Xu 2 , Yishuo Zhang 1 , Xiquan Weng 1 , Jiaji Luo 1 , Yanshuo Li 1
and Yu-Heng Mao 1,3, *

1 School of Exercise and Health, Guangzhou Sport University, Guangzhou 510500, China;
[email protected] (Y.C.); [email protected] (K.Y.); [email protected] (Y.Z.);
[email protected] (X.W.); [email protected] (J.L.); [email protected] (Y.L.)
2 The Fifth College of Clinical Medicine, Guangzhou University of Chinese Medicine,
Guangzhou 510500, China; [email protected]
3 Guangdong Key Laboratory of Human Sports Performance Science, Guangzhou 510500, China
* Correspondence: [email protected]; Tel./Fax: +86-(020)-38024623

Abstract: The intestinal tract of humans harbors a dynamic and complex bacterial community known
as the gut microbiota, which plays a crucial role in regulating functions such as metabolism and
immunity in the human body. Numerous studies conducted in recent decades have also highlighted
the significant potential of the gut microbiota in promoting human health. It is widely recognized that
training and nutrition strategies are pivotal factors that allow athletes to achieve optimal performance.
Consequently, there has been an increasing focus on whether training and dietary patterns influence
sports performance through their impact on the gut microbiota. In this review, we aim to present the
concept and primary functions of the gut microbiota, explore the relationship between exercise and
the gut microbiota, and specifically examine the popular dietary patterns associated with athletes’
sports performance while considering their interaction with the gut microbiota. Finally, we discuss
the potential mechanisms by which dietary patterns affect sports performance from a nutritional
perspective, aiming to elucidate the intricate interplay among dietary patterns, the gut microbiota, and
sports performance. We have found that the precise application of specific dietary patterns (ketogenic
diet, plant-based diet, high-protein diet, Mediterranean diet, and high intake of carbohydrate) can
Citation: Chen, Y.; Yang, K.; Xu, M.;
improve vascular function and reduce the risk of illness in health promotion, etc., as well as promoting
Zhang, Y.; Weng, X.; Luo, J.; Li, Y.;
recovery and controlling weight with regard to improving sports performance, etc. In conclusion,
Mao, Y.-H. Dietary Patterns, Gut
Microbiota and Sports Performance in although it can be inferred that certain aspects of an athlete’s ability may benefit from specific dietary
Athletes: A Narrative Review. patterns mediated by the gut microbiota to some extent, further high-quality clinical studies are
Nutrients 2024, 16, 1634. https:// warranted to substantiate these claims and elucidate the underlying mechanisms.
doi.org/10.3390/nu16111634
Keywords: gut microbiome; dietary pattern; sports performance; athlete
Academic Editors: Tatsuhiro
Hisatsune and Xi Wang

Received: 24 April 2024

Revised: 17 May 2024 1. Introduction
Accepted: 22 May 2024
In recent decades, with the rapid development of competitive sports worldwide,
Published: 26 May 2024
there has been an increasing demand for greater sports performance. Factors such as
training strategies, dietary patterns and training environments have garnered significant
attention in improving sports performance. Among these factors, dietary patterns are
Copyright: © 2024 by the authors.
particularly crucial alongside training strategies. It is imperative for athletes to consume
Licensee MDPI, Basel, Switzerland. adequate nutrition to optimize their condition during training and facilitate proper recovery
This article is an open access article afterwards [1,2]. Different dietary patterns may yield varying effects on athletes’ sports
distributed under the terms and performance and be suitable for different athletic specialties [3–5]. However, there is a
conditions of the Creative Commons paucity of comprehensive reviews examining the potential mechanisms by which dietary
Attribution (CC BY) license (https:// patterns influence sports performance.
creativecommons.org/licenses/by/ The human intestinal tract harbors a dynamic and complex bacterial community
4.0/). known as the gut microbiota, which emerging evidence suggests has beneficial effects on

Nutrients 2024, 16, 1634. https://doi.org/10.3390/nu16111634 https://www.mdpi.com/journal/nutrients

Nutrients 2024, 16, 1634 2 of 23

human health, including strengthening the gastrointestinal barrier, improving immune

function, and regulating glucose and fat metabolism [6]. Consequently, there is growing
interest in investigating whether the gut microbiota acts as a mediator for various diseases
such as obesity, diabetes, cardiovascular diseases, and non-alcoholic fatty liver disease
(NAFLD) [7]. Furthermore, recent attention has focused on exploring the potential role of
the gut microbiota as a mediator between dietary patterns, especially for specific micronu-
trients such as dietary fiber and anthocyanins (ACNs) that are abundant in dietary patterns,
and sports performance in athletes [8–10]. However, most studies have primarily evaluated
the effects of supplements, like probiotics, on athletic performance rather than deeply
investigating the relationship between dietary patterns and sports performance through
their impact on the gut microbiota. This may be attributed to the complex interaction
among different nutrients within dietary patterns and a limited scientific understanding
of their specific influence on sports performance. Moreover, the application of dietary
patterns on animal models may pose challenges, while using human models may impede
the exploration of the potential mechanisms. Therefore, this review aims to summarize
recent studies examining how some primary dietary patterns affect sports performance
in athletes while also proposing some possible mechanisms involving nutrient-mediated
interactions with the gut microbiota to provide practitioners with insights into enhancing
sports performance through targeted dietary patterns.

2. The Overview of Gut Microbiota

2.1. Gut Microbiota
The human microbiota is defined as the microorganisms that exist in symbiosis with
the human body, encompassing approximately 1014 –1015 bacteria [11]. It comprises bacteria,
archaea, fungi, viruses, bacteriophages, and protozoa [12]. These microorganisms colonize
various regions of the human body from birth onwards and are predominantly concentrated
in the oral and nasal cavities, skin, the urogenital tract and the gastrointestinal tract [11].
Notably, within the gastrointestinal tract, recent studies have revealed a microbial cell count
comparable to that of host cells [13]. Among the vast array of bacterial cells constituting
the gut microbiota, which comprises around 2000 identified species [14–16], the microbial
concentration gradually increases along the gastrointestinal tract, with an abundance of
particular anaerobic taxa [17,18]. In the stomach, the acidic pH limits the existence of
bacteria, so it presents the lowest number of bacteria, which are primary represented by
Lactobacillus, Candida, Streptococcus, and Helicobacter pylori. However, in the colon, the
favorable pH creates a more suitable habitat for bacteria such as Bacteroides, Clostridium,
Bifidobacterium, and Enterobacteriaceae, and most of these species are obligate anaerobic
bacteria, which participate in the decomposition of polysaccharides and the production of
short-chain fatty acids (SCFAs) [19].
The gut microbial composition and diversity undergo changes with aging and are
influenced by various factors. For instance, the mode of delivery significantly impacts the
initial colonization of bacteria. It has been suggested that infants born through natural
delivery predominantly harbor Lactobacillus and Prevotella species in their gut microbiota,
while those born via Cesarean section tend to possess microbiota dominated by Streptococ-
cus, Propionobacterium, and Corynebacterium bacteria. In adulthood, the gut microbiota forms
a relatively stable community, but it might vary among individuals. This microbiota com-
munity is mainly represented by the Bacteroidota and bacillota phyla, as well as Escherichia
and Lactobacillus to a lesser extent, but Bifidobacterium species remains constant. Among
the elderly, Bifidobacterium species decrease in quantity, but Escherichia and Lactobacillus
tend to increase [20,21]. Apart from the delivery mode, numerous other factors can also
influence the diversity of the gut microbiota, including dietary habits, antibiotic usage, host
genetics, lifestyle choices, surgical interventions, substance abuse disorders, mental health
conditions, and physical exercise [6,7,19,22].
Nutrients 2024, 16, 1634 3 of 23

2.2. The Main Function of Gut Microbiota on Health

For a considerable duration, extensive research has focused on the perspective that
bacteria are pathogenic to humans, exemplified by Streptococcus pyogenes, Bordetella pertussis,
Corynebacterium diphtheriae, Clostridium tetani, Salmonella typhimurium, Vibrio cholera, and
numerous others [22–25]. However, the majority of the microbiota are non-pathogenic and
even crucial for human health. Substantial evidence now suggests that the gut microbiota
plays a pivotal role in human well-being. It participates in metabolic functions by process-
ing indigestible dietary residues and producing SCFAs, which contribute to host metabolic
homeostasis [26]. SCFAs subsequently influence mucosal or systemic circulation to impact
peripheral organs and tissues. Apart from SCFAs, numerous other microbial metabolites
also play crucial roles in various physiological functions. These include bile acids, which
promote lipid uptake and maintain gastrointestinal function; lipids such as Lipopolysac-
charide (LPS) and Peptidoglycan, which enhance immune system function and regulate
glucose homeostasis through the activation of the brain–enteric–liver axis; and choline,
which regulates lipid metabolism and glucose homeostasis [27–29]. The bacteria species
of the gut microbiota also participate in the synthesis of glycans, amino acids, vitamins
and other essential components of the human metabolism [14,30]. Furthermore, the gut
microbiota actively contributes to fortifying the gastrointestinal barrier by promoting the
proliferation and turnover of epithelial cells, thereby enhancing its physiological function.
Toll-like receptors (TLRs) play a key role in this process [30–33]. Within the small intestine’s
epithelium cells, Paneth cells recognize the enteric bacteria and subsequently initiate the
expression of diverse antimicrobial factors through TLR activation, effectively safeguarding
against pathogenic bacterial infiltration [31,33,34]. Additionally, the microbiota stimulates
immunoglobulin (IgA) secretion and the production of antimicrobial molecules that inhibit
the proliferation and colonization of pathogenic bacteria, thus facilitating the development
of gut-associated lymphatic tissue (GALT) and bolstering the host immune system [34,35].
The immune system detects pathogen-associated molecular patterns (PAMPs), which are
TLR ligands, enabling it to identify potentially pathogenic bacteria, and consequently
leading to increased cytokine levels and the enhanced activation of T cells against these
pathogens as a response. Although the gut microbiota has many benefits for the human
body, dysbiosis characterized by a quantitative and qualitative imbalance in the microbial
composition, along with reduced diversity among species, can give rise to various disorders,
including diabetes, cardiovascular diseases, inflammatory bowel diseases (IBD), NAFLD,
and obesity. Notably, the presence of Akkermansia muciniphila, which represents 3–5% of the
typical intestinal microbial members, is decreased in obese people, and Alistipes putredinis,
which belongs to the phylum Bacteroidota, seems to be represented in people with type
2 diabetes and obesity [19,36–39]. In this context, several studies have demonstrated that
dietary interventions as well as exercise interventions hold promise as effective strategies
for modifying the composition and diversity of the gut microbiota towards a more favorable
community structure [19,40–43].

3. The Relation between Gut Microbiota and Exercise

Exercise is widely acknowledged to have a positive impact on human health, and re-
cent studies have increasingly focused on its relationship with the gut microbiota (Figure 1).
In contract to sedentary subjects, athletes and physically active individuals exhibit a greater
diversity of fecal bacteria, an abundance of beneficial species [44–46], and a heightened
microbial metabolism, as evidenced by increased activity in the carbohydrate and amino
acid metabolic pathway [45–47]. Moreover, regular endurance exercise modulates the
composition of the gut microbiota and reduces the presence of inflammation-associated
proteobacteria [19].
Nutrients 2024, 16, 1634
Nutrients 2024, 16, 1634 4 of 23

Figure1.
Figure The interaction
1. The interactionbetween exercise
between and the gut
exercise andmicrobiota. Exercise can lead
the gut microbiota. to changes
Exercise caninlead to c
the gut microbiota [44,47–51]. Unhealthy lifestyles can lead to dysbiosis [36,39]. The administration
the gut microbiota [44,47–51]. Unhealthy lifestyles can lead to dysbiosis [36,39]. The adm
of probiotics can affect the condition of the gut microbiota, which can subsequently affect sport
of probiotics can affect the condition of the gut microbiota, which can subsequently affect
performance [6,52]. The upward arrows indicate a rise or improvement, the down arrows indicate
formance
a drop. [6,52]. The upward arrows indicate a rise or improvement, the down arrows
drop.
3.1. Gut Microbiota in Athletes
A growing body of research has demonstrated that exercise exerts a modulatory effect
3.1. Gut Microbiota in Athletes
on the gut microbiota, leading to a distinction in the microbial composition between athletes
A growing
or physically activebody of research
individuals has demonstrated
and sedentary counterparts. Asthat exercise
depicted in Tableexerts
1, therea modu
were
fect onsignificant
the gut differences
microbiota, in the major taxa
leading to ata various levels between
distinction the two population
in the microbial composition
groups. It is worth noting that the trend observed in the Bacteroidetes to Firmicutes ratio
athletes or physically active individuals and sedentary counterparts. As depicted
between the two groups across different studies was inconsistent [53,54], which may be
1,attributed
there were significant
to several differences
factors including in the
substantial major variance,
individual taxa at human
various levels
species, andbetween
population groups. It is worth noting that the trend observed in the Bacteroidet
enterotypes [55].
But generally,
micutes ratio between it is widely
the twoaccepted
groupsthat athletes
across exhibit
different an enrichment
studies was of health-
inconsisten
promoting species within their gut microbiota, such as a higher abundance of Akkermansia
which may
spp. and be attributed
Prevotella to several The
spp. [44,47–49,53–55]. factors
studyincluding
conducted substantial
by Clarke et al.individual
on male vari
man species, rugby
international and enterotypes [55]. investigated the dietary intake and physical
players from Ireland
activity
Butofgenerally,
these athletes,
it isrevealing
widelya accepted
higher α-diversity in the gutexhibit
that athletes microbiota
an compared
enrichment to of he
sedentary controls [44]. The study also included two sedentary control groups consisting of
moting species within their gut microbiota, such as a higher abundance of Ak
healthy non-professional athletes with different a body mass index (BMI), including a high
spp.
BMIand
(BMI Prevotella
> 28) and lowspp. [44,47–49,53–55].
BMI (BMI < 25). According to The
the study
findings,conducted byathletes
the professional Clarke et al
international rugby
exhibited greater players
diversity from
in their Ireland investigated
fecal microbiota compared to both thecontrol
dietary intake
groups. Theand ph
gut microbiota
tivity of theseofathletes,
elite athletes consistedaofhigher
revealing 22 phylaα-diversity
of bacteria, while onlygut
in the 11 and 9 phyla
microbiota com
were found in the low and high BMI groups, respectively. Notably, increased Akkermansia
sedentary controls [44]. The study also included two sedentary control groups c
muciniphila, associated with the lean phenotype, was observed in professional athletes
ofand
healthy
the low non-professional
BMI group compared athletes
with the with
highdifferent
BMI group. a body massmuciniphila,
Akkermansia index (BMI), in
high BMI (BMI > 28) and low BMI (BMI < 25). According to the findings,
associated with positive metabolic function, is a mucin-degrading bacterium that inhabits the pro
the nutrient-rich mucus layer of the gut [56]. Furthermore,
athletes exhibited greater diversity in their fecal microbiota compared this study suggested that the to bot
microbial metabolism levels differed between professional athletes and sedentary groups,
groups. The gut microbiota of elite athletes consisted of 22 phyla of bacteria, w
11 and 9 phyla were found in the low and high BMI groups, respectively. No
creased Akkermansia muciniphila, associated with the lean phenotype, was observe
fessional athletes and the low BMI group compared with the high BMI group. Ak
muciniphila, associated with positive metabolic function, is a mucin-degrading b
Nutrients 2024, 16, 1634 5 of 23

as indicated by the increased activity in the carbohydrate and amino acid metabolism
pathways in athletes. However, it is important to note that the differences in dietary
patterns, which refer to a higher total energy, macronutrient (especially protein), and fiber
intake in professional athletes compared with the control group, may also influence the gut
microbial composition [44].

Table 1. Comparison of the gut microbial composition between athletes/physically active population
and non-athletes/sedentary population.

Author,
Country Sample Size, Sex and Age Main Findings on Gut Microbial Composition
Year
Athletes/Physically Non-Athletes/Sedentary
Active Population Population
Bacteroidetes (52.53%) Bacteroidetes (62.81%)
Firmicutes (43.99%) Firmicutes (32.14%)
Xu et al., 2022 n = 66 (males = 36, females = 30), Prevotella (20.88%) Prevotella (26.81%)
China
[53] Age: 18–25 years Bacteroides (24.96%) Bacteroides (25.01%)
Faecalibacterium (6.86%) Faecalibacterium (10.57%)
Megamonas (11.67%) Megamonas (5.15%)
Enterotype:
Enterotype:
Humińska- Endurance group:
n = 52, males Control group:
Lisowska et al., Poland Bacteroides-driven (46.70%)
Age: 19–24 years Bacteroides-driven (40.90%)
2024 [55] Strength group:
Ruminococcus-driven (40.90%)
Prevotella-driven (50.00%)
n = 54 (males = 28, females = 26) Bacteroidetes (50.40%) Firmicutes (48.30%)
Hintikka et al., Age: Firmicutes (46.00%) Bacteroidetes (46.20%)
Finland
2022 [54] Athlete group: 27.1 ± 5.1 years Proteobacteria (2.30%) Proteobacteria (3.36%)
Control group: 27.4 ± 5.6 years Actinobacteria (0.79%) Actinobacteria (1.57%)

Additionally, a study revealed that competitive cyclists exhibited a decreased relative

abundance of Bacteroides spp. Furthermore, the relative abundance of Prevotella spp. was
found to be higher in cyclists who engaged in training for more than 11 h per week
compared with those who trained less frequently [45]. These findings provide evidence
supporting the notion that physical exercise can induce alterations in the composition of
the gut microbiota.

3.2. Impact of Exercise Interventions on Gut Microbiota

To further substantiate the impact of physical exercise intervention on the gut mi-
crobiota, several studies have been conducted to explore the causal relationship between
exercise and alterations in the gut microbial composition (Figure 1). One study demon-
strated that an endurance exercise intervention induced modifications in the gut microbial
composition of sedentary, non-trained Finnish women, while controlling for factors such
as dietary habits, weight, and body composition [50]. Notably, there were no significant
changes observed in the total energy intake or macronutrient and dietary fiber consumption
following training. Moreover, no discernible differences were found in the α-diversity
of the gut microbiota or the phylum-level relative abundance between pre-intervention
and post-intervention samples. However, endurance exercise did lead to an increase in
the relative abundance of members of the genera Verrucomicrobia and Akkermansia, while
reducing the levels of inflammation-associated Proteobacteria within the gut.
In addition to endurance exercise, resistance training also exerts an influence on the
gut microbiota. Smith et al. demonstrated that 10 weeks of resistance training can improve
the alpha diversity in younger untrained adults [57]. Another study conducted by Dupuit
et al. explored the impact of a combination of high-intensity interval training (HIIT) and
resistance training on the gut microbiota of postmenopausal women [58]; the authors
indicated that the training intervention did not significantly change the alpha diversity
Nutrients 2024, 16, 1634 6 of 23

and overall taxonomy of the fecal microbiota but modified the beta diversity, which is
inconsistent with the previous study, showing that more research about resistance training
is needed.
However, several other factors may also impact the effectiveness of exercise on the gut
microbiota. For instance, one study has indicated that BMI could potentially influence the
response of the gut microbiota to physical exercise. According to this particular study [59],
individuals with different body compositions (lean and obese) exhibit distinct baseline gut
microbiota profiles. However, after a 6-week aerobic exercise intervention, no significant
difference in the microbiota community composition was observed between lean and
obese subjects.

3.3. The Influence of Gut Microbiota on Sports Performance

Exercise exerts a significant impact on the composition of the gut microbiota, while it
is reciprocally influenced by the gut microbiota. Determining the precise effects of the gut
microbiota on sports performance in human clinical studies poses a challenge due to the
intricate interplay of nutritional, genetic and environmental factors [6]. However, germ-free
animal models provide a novel approach and have already been employed to elucidate the
impact of the gut microbiota on sports performance [60].
A cross-sectional study conducted by Hsu et al. investigated the swimming capacity
of specific pathogen-free (SPF), germ-free (GF), and Bacteroides fragilis gnotobiotic mice.
The results revealed that the swim-to-exhaustion time was the longest for SPF mice and the
shortest for GF mice, indicating a compromised sports performance in the absence of a gut
microbiota [60]. Although the effects of probiotics supplementation have been studied in
athletes and physically active populations, the small number of participants, the different
exercise intervention programs implemented, and the different training histories of the
participants may have influenced the outcomes [61]; therefore, the results remain contro-
versial. However, a review conducted by Marttinen et al. as summarized several benefits of
probiotics for the athlete. The authors demonstrated that the administration of probiotics
might reduce symptoms of gastrointestinal and upper respiratory tract illnesses, enhance
physical performance, improve post-exercise recovery, and improve mood-related out-
comes [6,62–65]. Therefore, there exists a significant association between the composition
of the gut microbiota and sports performance (Figure 1).

4. The Influence of Several Typical Dietary Patterns on the Gut Microbiota

Personal dietary habits play important roles in shaping the composition of the gut
microbiota in humans. Although further research is needed to fully understand the intri-
cate relationship between diet and the gut microbiota, numerous studies have highlighted
the significant impact of different types of dietary patterns on the composition of the gut
microbiota within 24 h [66,67]. The dietary patterns of individuals can be broadly catego-
rized into vegetarians, meat eaters and balanced eaters, each exhibiting a distinct profile
in the gut microbiota. Different types of dietary patterns elicit distinct alterations in the
proportions of Firmicutes, Bacteroidetes, Proteobacteria and Actinobacteria. Changes in
the gut microbiota induced by dietary interventions are observed within 24 h and return to
baseline levels within 48 h after discontinuation [66]. These changes encompass alterations
in carbohydrate and protein fermentation, intestinal inflammation, fat oxidation, as well as
an increase in amino acid availability, potentially promoting protein anabolism [46,68–71].
Furthermore, the quality, quantity and molecular characterization of carbohydrates, pro-
tein, and fat are key factors influencing both the composition and metabolism of the gut
microbiota. Unhealthy dietary patterns can stimulate the proliferation of detrimental gut
bacteria that pose risks to human health. However, a healthy dietary pattern has restorative
effects on beneficial gut bacteria [72]. The maintenance and modulation of beneficial gut
microbiota are vital for host health. In addition to general dietary patterns, probiotics
supplementation and wholefood supplementation are also common nutrition strategies.
Notably, probiotics are defined as living organisms with beneficial effects on health. Most
 probiotics supplementation and wholefood supplementation are also common nutrition
strategies. Notably, probiotics are defined as living organisms with beneficial effects on
health. Most probiotic supplementations contain high concentrations of Lactobacillus or
Nutrients 2024, 16, 1634
Bifidobacterium spp., which can support the immune system of the host, regulate7 gut of 23
per-
meability, and produce sanatory metabolites [73]. Unlike synthetic supplements, whole-
food supplements are based
probiotic on the core
supplementations ideahigh
contain of supplying
concentrationsthe body withornutrients
of Lactobacillus in their
Bifidobacterium
spp., which can support the immune system of the host, regulate
pure, unaltered state. This implies that these supplements are rich in a broad spectrum ofgut permeability, and
produce sanatory metabolites [73]. Unlike synthetic supplements, wholefood supplements
vitamins, minerals, antioxidants,
are based and
on the core idea other crucial
of supplying the bodynutrients that
with nutrients arepure,
in their inherently
unaltered found
state. in
the foods from which theythat
This implies arethese
sourced [74,75].
supplements areIn this
rich in asection, the influence
broad spectrum of minerals,
of vitamins, several typ-
ical dietary patterns on the gut microbiota will be discussed in detail (Figure 2).which
antioxidants, and other crucial nutrients that are inherently found in the foods from
they are sourced [74,75]. In this section, the influence of several typical dietary patterns on
the gut microbiota will be discussed in detail (Figure 2).

Figure 2. The effects of dietary patterns on the gut microbiota. Different dietary patterns will lead to
Figure 2. The effects of dietary patterns on the gut microbiota. Different dietary patterns will lead to
different changes in the abundance of the gut microbiota. Reference: ketogenic diet [52], plant-based
different changes diet
in the
[76],abundance of [48,77],
high-protein diet the gut microbiota.
Mediterranean Reference:
diet [78–80], highketogenic diet [52],[81].
intake of carbohydrates plant-based
The
diet [76], high-protein
upwarddiet [48,77],
arrows indicateMediterranean diet
a rise or improvement, the[78–80],
down arrows high intake
indicate of carbohydrates [81].
a drop.
The upward arrows indicate a rise or improvement, the down arrows indicate a drop.
4.1. Ketogenic Diet
The ketogenic diet (KD) is characterized by a high fat content, a low carbohydrate
4.1. Ketogenic Diet
intake, and an appropriate proportion of protein and other essential nutrients. There are
four main
The ketogenic diet types
(KD)ofis ketogenic diets: (1) the
characterized byclassical
a highKDfatwith a macronutrient
content, a low ratio of 4%
carbohydrate
carbohydrate, 90% fat and 6% protein, (2) medium-chain triglyceride with a macronu-
intake, and an appropriate proportion
trient ratio of 20% carbohydrate,of protein and other
10% long-chain essential
triglycerides nutrients.
fat, 60% There are
medium-chain
four main typestriglycerides
of ketogenic diets:
fat and (1) the(3)classical
10% protein, modified KD with
Atkins withaa macronutrient
macronutrient ratioratioof 10%of 4%
carbohydrate, 65% fat and 25% protein, (4) low-glycemic-index
carbohydrate, 90% fat and 6% protein, (2) medium-chain triglyceride with a macronutri- diet with a macronutrient
ratio of 10% carbohydrate, 60% fat and 30% protein [82]. It can be seen that the KD does
ent ratio of 20%notcarbohydrate,
have a fixed nutrient10%ratio,
long-chain
but a high triglycerides
proportion of fatfat,
and 60% medium-chain
a low proportion of car- tri-
glycerides fat and 10% protein,
bohydrates should be (3) modified
guaranteed. Atkins
The primary with a macronutrient
objective ratioisof
of this dietary pattern to 10%
shift car-
the glucose metabolism towards fat metabolism through
bohydrate, 65% fat and 25% protein, (4) low-glycemic-index diet with a macronutrientthe restriction of carbohydrate
intake. Consequently, the KD can effectively lower blood sugar levels and increase free
ratio of 10% carbohydrate, 60% fat
fatty acid and ketone and 30%
production, protein
thereby [82]. neuronal
influencing It can be seen that
excitability [83].the KD does
Notably,
not have a fixedthe
nutrient ratio, butbyathe
KD is characterized high proportion
production ofbodies
of ketone fat and a low proportion
(3-hydroxybutyrate, acetateof carbo-
and
acetoacetate). The elevation in ketones contributes to an increase
hydrates should be guaranteed. The primary objective of this dietary pattern is to shift thein anti-inflammatory and
antioxidant activity, immune regulation, intestinal mobility and barrier function, cellular
glucose metabolism
growthtowards fat metabolism
and differentiation, through
ionic absorption, theasrestriction
as well the prevention of of
carbohydrate
distal ulcers, in-
take. Consequently,
Crohn’sthe KD can
disease, effectively
and colon cancers. lower blood
Additionally, thesugar
KD waslevels
initiallyand increase
employed free fatty
for manag-
acid and ketone production, thereby influencing neuronal excitability [83]. Notably, the
KD is characterized by the production of ketone bodies (3-hydroxybutyrate, acetate and
acetoacetate). The elevation in ketones contributes to an increase in anti-inflammatory and
antioxidant activity, immune regulation, intestinal mobility and barrier function, cellular
Nutrients 2024, 16, 1634 8 of 23

ing refractory epilepsy and has progressively extended its application to encompass other
neurological disorders [84], such as Parkinson’s disease and Alzheimer’s diseases. With the
advancement of medical technology and sports science, there are several studies that have
demonstrated the potential of this dietary pattern in enhancing sports performance in some
ways [85–87]. Nevertheless, this diet still has some limitations; for example, the ability of
muscle to use glycogen for oxidation is impaired after long-term ketoadaptation, leading to
an inability to utilize the available glycogen, which provides a more effective energy source
when the oxygen supply becomes limiting. Therefore, the performance of higher-intensity
endurance exercise will be limited, which might increase the risk of injury for athletes [88].
A study conducted by Ang et al. in both mice and humans demonstrated that the
ketogenic diet resulted in decreased levels of Bifidobacterium, which was mediated by the
increased production of ketone bodies, especially beta-hydroxy butyrate. The decrease in
Bifidobacterium reduced the levels of intestinal and visceral fat pro-inflammatory Th17 cells,
which might be a potential mechanism contributing to the ketogenic diet’s ability to reduce
body fat because of the relationships between obesity and chronic low-grade inflammation.
Furthermore, the ketogenic diet also decreased Lactobacilli and increased Fusobacteria and
Escherichia [89].
Several studies have indicated that variations in the quantity and source of dietary
fat can exert distinct effects on the host, with some of these effects potentially mediated
by the gut microbiota. The consumption of saturated fat has been shown to increase
the abundance of bacteria expressing LPS, leading to elevated levels of LPS and a pro-
inflammatory state known as metabolic endotoxemia. Furthermore, excessive fat intake is
also associated with reduced levels of butyric acid and retinoic acid [90], both crucial for
maintaining gut homeostasis. Furthermore, the consumption of saturated fat can enhance
the relative abundance of Bilophila wadsworthia by facilitating the conjunction of taurine
with host LPS, which serves as a terminal electron acceptor and subsequently leads to the
production of hydrogen sulfide and secondary bile acids. This cascade may ultimately result
in intestinal barrier disruption and immune cell infiltration [91]. Hence, it implies that a KD
characterized by a high saturated fat intake could potentially elevate the inflammatory level
of the host. Conversely, polyunsaturated fat acid (w-3) can increase SCFAs and promote
gastrointestinal integrity and inflammation. Furthermore, polyunsaturated fat increases
the abundance of Bifidobacterium, Lactobacilli, and Akkermansia muciniphila, which are also
increased by exercise. Thus, polyunsaturated fat might contribute to health and sports
performance by mimicking the effects of exercise, but the dose remains controversial; more
research is needed to investigate this [92,93].
Notably, with the advancement of research on diet and nutrition, the classical KD has
undergone certain variations; for instance, the very-low-calorie KD (VLCKD) is charac-
terized by a caloric intake of below 800 kcal/day. One study revealed that the VLCKD
results in a more substantial weight reduction, rendering it an excellent option for weight
loss [94,95]. Regarding the gut microbiota, a review has summarized the effects of the VL-
CKD on the gut microbiota [96]; in this study, the authors demonstrated that the abundance
of Bacteroidetes, Firmicutes, Proteobacteria, and Verrucomicrobiota in people who under-
took the VLCKD increased and that the abundance of Firmicutes, Firmicutes/Bacteroidetes
ratio, Proteobacteria, and Actinobacteria decreased. These seemingly contradictory results
suggest that further research is warranted to explore the impact of the VLCKD on the
gut microbiota. Furthermore, the application of the VLCKD still remains controversial,
especially for athletes, because this diet is used more in obese patients. However, the
VLCKD could be used to control weight acutely in special sports during a special period
with strict medical supervision, such as in gymnastics [97–99].
In conclusion, the impacts of the KD on the gut microbiota remains inconclusive and
controversial, necessitating further studies to comprehensively understand its effects.
Nutrients 2024, 16, 1634 9 of 23

4.2. Plant-Based Diet

The plant-based diet is a dietary pattern primarily based on a diverse range of plants,
encompassing seeds, fruits, and plant tissues that provide energy for human consumption.
This includes cereals, tubers, legumes and their derivatives, as well as fruit and vegetable
products. The distinguishing features of the plant-based diet are its high carbohydrate
content, low energy density, low fat content, and absence of cholesterol, antibiotics or
hormones [100]. Long-term adherence to a plant-based diet not only reduces the risk of
many chronic diseases, but also contributes to the lower emission of greenhouse gases such
as carbon dioxide during food processing compared to other methods [101]. Consequently,
it plays an integral role in promoting human health and environmental preservation [102].
One study explored the microbial composition of 258 participants who adhered to
one of four dietary patterns: the Western diet group, flexitarian group, vegetarian group,
and vegan group [76]. Notably, the Western food group is characterized by a high intake
of energy, salt, saturated fat, simple or added sugar, and a low intake of fruits and vegeta-
bles [103]. The vegetarian group is characterized by omitting defined food groups such as
meat, sausage, fish, etc., and the vegan group is characterized by additionally omitting dairy
products and honey [104–106]. Flexitarians generally consume meat or sausage once or
twice per week [107]. The western diet group exhibited the lowest abundance of Bacteroides,
Lachnospiraceae_1, Butyricoccus, Lachnospiraceae UCG_004, and Haemophilus; whereas
the vegan group showed the highest abundance. For Dorea, the Ruminococcus torques group,
Eubacterium ruminantium group, Ruminococcaceae, Lachnospiraceae_2, Lactobacillus, and
Senegalimassilia, the lowest abundance was observed in the vegan group, while the highest
abundance was observed in the Western diet group [76]. Notably, a high abundance of
Lachnospiraceae in the vegan group indicates the extensive fermentation of plant-based
polysaccharides into SCFAs like butyrate, which is beneficial for human health. For ex-
ample, it serves as a crucial energy source for colonic epithelial cells, regulates intestinal
inflammation, and confers protection against colon cancer in humans [76,108,109].
Furthermore, the vegan diet might also contribute to improving performance and pro-
moting recovery in endurance sport by affecting body composition, blood flow, antioxidant
capacity, systemic inflammation, and glycogen storage [110].

4.3. High-Protein Diet

People who stick to a high-protein diet can take in higher-quality protein and provide
the body with amino acids. Protein is a macronutrient, as well as the main component of
skeletal muscle. The uptake and catabolism of specific proteins by the liver and skeletal
muscle are different, as is their ability to regulate the muscle protein synthetic response [111].
Amino acids can be metabolized into branched-chain fatty acids and SCFAs, ammonia,
sulfides, indole, and phenolic compounds via the gut microbiota [112]. Some of these (e.g.,
SCFAs and indole) may be beneficial for the health of the gut, while other metabolites (e.g.,
ammonia and p-cresol) may decrease gut epithelium integrity [113].
The high-protein diet is widely popular and frequently adopted by fitness enthusi-
asts and athletes, particularly for the latter who engage in intense exercise routines that
necessitate strict dietary practices to support optimal performance [114]. In contrast to
the general population, athletes often consume significantly higher amounts of protein;
however, if this excess protein remains unabsorbed, according to a study conducted by
Moreno-Pérez et al. [48], it can enter the colon and promote the growth and selection of
specific bacteria. In this study, a 10-week supplementation with protein, commonly used to
meet the elevated protein requirements among individuals undergoing training, resulted in
an increased abundance of Bacteroidetes while decreasing the taxa associated with overall
health, including Roseburia spp., Blautia spp., and Bifidobacterium longum, among runners.
Another study has compared the gut microbiota of bodybuilders consuming a high-protein
diet with sedentary controls [77], and found that excessive protein intake increased the
abundance of protein-fermenting bacteria such as Clostridium, Bacillus, Staphylococcus, and
other species belonging to the Proteobacteria family. Moreover, the high-protein diet might
Nutrients 2024, 16, 1634 10 of 23

lead to a reduction in carbohydrate-fermenting bacteria, such as Bacteroides, Lactobacillus,

Bifidobacterium, Prevotella, Ruminococcus, Roseburia, and Faecalibacterium. The fermentation of
incompletely digested protein in the colon might lead to the production of toxic metabolites
such as ammonia, biogenic amines, indole compounds, and phenols. However, there was
no significant difference in the abundance of selected bacteria (Bifidobacterium spp., Bac-
teroides spp., Faecalibacterium prausnitzii, Akkermansia Muciniphila) between the bodybuilder
group and control group; the possible reason for this is that both of the two groups met the
criteria for the recommended fiber intake, and the effect of high protein intake on the gut
microbiota might have been attenuated by the appropriate intake of carbohydrate and fiber.
Therefore, it is imperative to strictly control not only the types of protein consumed, but
also the quantity ingested by athletes.

4.4. Mediterranean Diet

The Mediterranean diet (MD) originates from the Mediterranean region, including
Greece, Spain, France, and Italy. It is based on the traditional dietary habits of the countries
bordering the Mediterranean Sea. This dietary habit is characterized by a high intake of
fruits, vegetables, cereals, olive oil, legumes and tree nuts, a moderate intake of seafood,
and a low intake of sugar sweetened foods, red and proceed meat, and carbonated bev-
erages [4]. However, there remains controversy surrounding its precise definition. In
a recent study [115], the authors attempted to establish a unified definition of the MD
by considering daily servings of key foods and their nutrient content: Vegetables: 3 to
9 servings; Fruit: 0.5 to 2 servings; Cereals: 1 to 13 servings; Olive oil: up to 8 servings.
Considering its energy intake and macronutrient composition, the MD can be classified
as a predominantly plant-based dietary pattern, encompassing vegetables, fruits, cereals,
and olive oil [116–118]. It is notable that the MD exhibits a relatively high fat content, with
monounsaturated fats comprising twice the amount of saturated fat. The primary source of
monounsaturated fats in the MD is olive oil, which is closely associated with the traditional
olive cultivation in the Mediterranean region. Additionally, the MD allows for the moderate
consumption of white and red meat. Extensive evidence supports that adherence to the MD
promotes longevity while reducing the metabolic risks associated with diabetes mellitus,
obesity, and other metabolic syndromes [119,120]. Moreover, it demonstrates a reduced
risk of malignancy and cardiovascular disease while enhancing cognitive function [116].
Numerous studies have consistently demonstrated that the gut microbiota plays
a crucial role as a potential mediator in the association between the MD and human
health. A study has indicated that nearly 60% of the overall composition of the gut
microbiota is responsive to dietary changes [121]. The MD can not only modulate the
diversity and composition of the gut microbiota, but also improve the generation of SCFAs
due to its high proportion of plant-based food [78]. Previous research has shown an
association between the MD and Prevotella [78,79], while another study suggests that the
MD contributes to reducing dysbiosis and increasing Bifidobacterium among patients with
metabolic syndrome [78,80]. However, it should be noted that not all studies support
the positive influence of the MD on the gut microbiota. Some investigations found no
significant difference in the gut microbiota composition between individuals adhering to
either the MD or Western diet interventions for 6 months. Therefore, further research is
warranted to comprehensively explore the impact of the MD on the gut microbiota [78,122].
Furthermore, according to a narrative review conducted by Griffiths et al., the application
of MD or individual foods and compounds in this dietary pattern might have potential
positive effects on oxidative stress, inflammation, injury, illness risk, and cognitive and
vascular function in competitive athletes [4].

4.5. High Intake of Carbohydrate

Limited research has been conducted on the high-carbohydrate diet, probably due
to the fact that a high intake of carbohydrate is not typically considered as an indepen-
dent dietary pattern but rather as a supplementary measure in other dietary patterns to
Nutrients 2024, 16, 1634 11 of 23

meet the energy requirements of athletes, given its role as a primary fuel source during
exercise [123]. It is recommended that athletes consume ample amounts of simple car-
bohydrates to maintain glucose homeostasis and limit their fiber intake prior to exercise
in order to minimize gastrointestinal discomfort. Non-digestible carbohydrates will be
discussed later. Adequate carbohydrate consumption is crucial for athletes. The ingestion
of simple carbohydrates before and during exercise (e.g., glucose, fructose, sucrose) can
alleviate fatigue, facilitate rehydration and the maintenance of optimal fluid balance, and
enhance sports performance [124–128]. For example, lactose may serve as an effective
fuel source before, during and after exercise, thereby enhancing sports performance and
aiding recovery while also potentially exerting beneficial effects on the gut microbiota,
such as increasing Bifidobacteria and Lactobacilli populations [81]. According to a study
conducted by Faits et al., which discusses the different effects of simple, refined, and unre-
fined carbohydrate-containing foods on the gut microbiota, after the consumption of an
unrefined carbohydrate diet, the abundance of Roseburia was higher and fecal secondary
bile acid concentrations were lower relative to the simple carbohydrate diet, whereas the
abundance of Anaerostipes was higher after the consumption of a simple carbohydrate diet
relative to the refined carbohydrate diet [129].
Notably, athletes in many sports often consume a high amount of fast-absorbed carbo-
hydrates to maximize glycogen storage. However, they also aim to avoid non-digestible
carbohydrates in order to prevent intestinal issues and other unfavorable syndromes that
can negatively impact sports performance, such as bloating and diarrhea [130]. While a
high intake of fast-absorbed carbohydrates can increase energy storage during training or
competition, a low intake of dietary fiber may lead to the reduced production of short-chain
fatty acids (SCFAs), altered intestinal transit times, and a loss of bacterial diversity [3], all
of which have negative implications for long-term health [131]. Therefore, it is important
for athletes to consume a certain amount of fiber to generate less gas after fermentation by
the gut microbiota in order to gain health benefits and avoid gastrointestinal issues.

5. Different Dietary Patterns and Sports Performance—Gut Microbiota as the Mediator

5.1. Gut Microbiota as the Mediator
With the advancement of competitive sports, whether it pertains to athlete-to-athlete
competition or the audience’s heightened expectations for sporting event enjoyment, both
lead to elevated demands on athletes’ capabilities. Numerous factors, such as exercise
intensity, dietary patterns, lifestyle choices and genetic inheritance, among others, can
influence the sports performance of athletes or physically active individuals. The gut
microbiota—an integral component of human beings since birth—has emerged as a promi-
nent area of research interest due to its intricate composition and structure. Several studies
have indicated disparities between the gut microbiota profiles of athletes and those of
normal individuals. Numerous investigations have attempted to establish whether the gut
microbiota is a mediator linking dietary patterns and sports performance. Here, we present
a concise overview of the current primary evidence pertaining to the aforementioned di-
etary patterns and the discuss probable mechanisms by which dietary patterns affect sports
performance (Table 2).

Table 2. The probable mechanism of dietary patterns affecting sports performance.

Author, Dietary
Substance Subjects Pathway Most Important Findings
Year Pattern
(Caesar et al., 2015 LPS/TLR4 Increases inflammatory
Ketogenic diet Saturated fat Male mice
[115]) pathway indices in WAT
Bacillus
Promote the Produces proteases which
(Minevich et al., High-protein coagulans
Males (n = 11) absorb and can increase amino
2015 [119]) diet GBI-30, 6086
utilize of protein acid absorption in humans
Protein
Nutrients 2024, 16, 1634 12 of 23

Table 2. Cont.

Author, Dietary
Substance Subjects Pathway Most Important Findings
Year Pattern
Higher abundance of
Lactobacilli
Higher ratio of Firmicutes
to Bacteroidetes
Decrease the
Lower butyrate
binding of
(Zhu et al., High-protein Lower SCFAs-producing
Animal protein Male rats (n = 32) CD14 and
2017 [121]) diet bacteria
LPS-binding
Lower LPS-binding
protein
protein
Lower transcription factor
CD14 receptor
Lower inflammation
Plant-based
(Jäger et al., diet/ C2C12 myotubes AMPK/PGC-1α Enhances fatty acid
Dietary fiber
2007 [81]) Mediterranean Female mice pathway oxidation of muscle
diet
Reduces oxidative stress
Plant-based
Promotes mitochondrial
(Yang et al., diet/ C2C12 myotubes AMPK signaling
Anthocyanins biogenesis
2023 [132]) Mediterranean Male mice (n = 60) pathway
Converse skeletal muscle
diet
fiber
This table shows the mechanisms of the effects of nutrients in different dietary patterns on the gut microbiota.
AMPK: adenosine 5-monophosphate-activated protein kinase; CD14: cluster of differentiation 14; LPS: lipopolysac-
charide; PGC-1α: proliferator-activated receptor gamma coactivator; SCFAs: short-chain fatty acids; TLR4: toll-like
receptors 4; WAT: white adipose tissue.

To date, the impact of the KD on sports performance remains controversial. As

mentioned earlier, the consumption of saturated fat increases the LPS level in the host,
which activates toll-like receptors 4 (TLR4) and cluster of differentiation 14 (CD14), leading
to obesity, increased inflammatory indices in white adipose tissue (WAT), and insulin
resistance [133]. Interestingly, this effect was observed only in subjects consuming saturated
fat. These findings suggest that athletes implementing a KD can increase their intake of
unsaturated fats to avoid inflammation and insulin resistance. Additionally, the VLCKD
may have a beneficial effect on obesity by regulating the gut microbiota and restoring
homeostasis [96]. The study by Gutierrez-Repiso et al., 2019, discussed the association
between the VLCKD and weight loss through the gut microbiota [134]. On one hand, the
authors reported that the abundance of Butyricimonas and Oscillospira increased at the genus
level. Notably, Oscillospira is positively associated with high-density lipoprotein, butyrate
and leanness, while Butyricimonas is positively associated with energy metabolism and
homeostasis between the microbiota and host. Both of these gut microbiota are beneficial for
weight loss. On the other hand, the proportion of Serratia and Citrobacter, whose abundance
has been positively correlated to obesity, decreased. Therefore, the VLCKD can positively
regulate the gut microbiota after obesity-relative dysbiosis. This dietary pattern enables
rapid short-term weight reduction, making it suitable for athletes who need to quickly
regain an optimal weight.
In terms of the high-protein diet, research has primarily focused on the impact of
protein. Evidence suggests that the gut microbiota contributes to the absorption and utiliza-
tion of protein, as well as the anabolism and functionality of skeletal muscle by providing
fuel and storage and modulating inflammation. For example, the co-administration of the
probiotic Bacillus coagulans (GBI-30,6086) with protein has been shown to reduce the inflam-
mation of epithelial cells, enhance nutrient absorption and stimulate protease production
for increased amino acid uptake in humans [135,136]. These effects have the potential to
mitigate muscle damage and facilitate muscle recovery, thereby promoting sports perfor-
Nutrients 2024, 16, 1634 13 of 23

mance [137]. In addition, animal studies have been conducted to investigate the effects of
different protein types on the gut microbiota, with a particular focus on comparing animal-
based proteins to plant-based proteins [138–141]. These studies have demonstrated that the
consumption of meat protein leads to a higher abundance of Lactobacilli and an increased
ratio of Firmicutes to Bacteroidetes, while also reducing levels of butyrate-producing bac-
teria (e.g., Bacteroides and Prevotella), LPS-binding protein, and transcription factor CD14
receptor when compared to non-meat protein intake. Furthermore, dairy proteins appear
to have an intermediate effect between meat and non-meat proteins. It is worth noting that
LPS-binding protein binds to CD14 in order to activate macrophages, which can subse-
quently produce inflammatory cytokines, leading to inflammation. Based on these findings,
it can be hypothesized that athletes may benefit from consuming more meat protein rather
than non-meat protein in order to mitigate muscle inflammation and maintain optimal
sports performance. However, the studies mentioned above have primarily focused on
rodents, with limited exploration of their effects on humans; one reason for this may be
that it is difficult to intervene individually with different types of proteins in humans,
and that other nutrients might interfere with the experimental results. A human study
investigating the impact of various protein types on gut the microbiota and incorporating
a high- or low-saturated fat component into the study design indicated that the intake of
saturated fat may cover up the effects of protein types [142]. Another study conducted by
Losasso et al. that compared the influence of vegan, vegetarian and omnivore-oriented
Westernized dietary styles on the gut microbiota indicated that vegans and vegetarians
show higher α-diversity than those who consume animal protein, the main operational
taxonomic units associated with the phylum Bacteroidetes, and the genus Prevotella, which
can improve glycogen storage, was more prevalent among individuals that consume more
fiber and vegetable protein. However, the subjects in this study also consumed different
nutrients, which may have influenced the results [143]. Consequently, it can be inferred
that enhancing the protein bioavailability and absorption, as well as muscle protein syn-
thesis, serves as an important mechanism through which the gut microbiota influences
muscle mass and function. This mechanism is likely regulated by SCFA production, thereby
affecting insulin sensitivity, inflammation, and insulin growth factor I (IGF-I) release to
maintain anabolic–catabolic balance. Furthermore, more studies elucidating the effects
of different protein types in humans that consider other dietary components beyond just
protein consumption are needed [144,145].
It is noteworthy that dietary fiber plays an essential role in both the plant-based diet
and MD, as it constitutes their main component. Dietary fiber is composed of complex car-
bohydrates, including fermentable (mainly soluble) and non- or poorly fermentable (mainly
insoluble) fibers, as well as oligosaccharide. Dietary fiber influences the composition of the
gut microbiota, contributing to the establishment and maintenance of a healthy and diverse
gut microbiota while improving intestinal immunity [146]. However, the insufficient intake
of dietary fiber may have adverse effects on human health. The dietary fiber in the afore-
mentioned dietary patterns includes “Microbiota-accessible carbohydrate (MACs)”, which
are complex carbohydrates found in fruits, vegetables, legumes, and whole grains [116].
A study conducted by Xu et al. has shown that a high intake of MACs promotes lipid
profile improvement, glycemic control, body weight reduction, and an inflammatory maker
decrease compared with low MAC intake [147]. Furthermore, MACs can influence the gut
microbiota and modulate the growth of species that produce SCFAs, which are the end
products of dietary fiber fermentation in the intestines. SCFAs play an essential role in
human metabolism. A study has indicated that SCFAs can directly activate Adenosine
5’-monophosphate (AMP)-activated protein kinase (AMPK) by increasing the AMP/ATP ra-
tio in skeletal muscle and liver or indirectly activate it via the Ffar2-leptin pathway [148–150].
The activation of AMPK triggers the expression of proliferator-activated receptor gamma
coactivator PGC-1α, which is known to regulate the transcriptional activity of key factors
including peroxisome proliferator-activated receptors PPARα, PPARδ, PPARγ, liver X
receptor (LXR), and farnesoid X receptor (FXR). These factors are crucial to regulate the
Nutrients 2024, 16, 1634 14 of 23

metabolism of cholesterol, lipid, and glucose. The fatty acid oxidation of muscle and liver is
ultimately enhanced, while de novo fatty acid synthesis in the liver is reduced [151,152]. In
addition, SCFAs have been demonstrated to enhance the protein expression of PGC-1α and
uncoupling protein-1 (UCP-1) in brown adipose tissue, subsequently promoting thermogen-
esis and fatty acid oxidation. These results suggest that the plant-based diet and MD, which
are rich in dietary fibers, could be considered for dietary planning among weight-conscious
athletes such as marathon runners. However, it is still crucial for endurance athletes to
maintain an adequate intake of simple carbohydrates. For instance, in an international
marathon competition that typically lasts for a minimum duration of approximately two
hours, athletes require sufficient glycogen reserves to optimize their sports performance.
Therefore, carbohydrate loading is commonly employed by endurance athletes as a strategy
to enhance glycogen concentrations prior to competitions. However, it is crucial to avoid
consuming carbohydrates that are indigestible and unabsorbable in the small intestine, such
as fiber and resistant starch [88]. Nevertheless, scientific evidence suggests that adopting a
high-carbohydrate, low-fiber dietary pattern can have detrimental effects on the gut micro-
biota and overall health. These effects include disruptions in intestinal transit times, the
loss of bacterial diversity, and reduced SCFA production [153–155]. Thus, athletes should
judiciously manage both the timing and quantity of their intake of simple carbohydrates
and nondigestible carbohydrates to optimize their sports performance while minimizing
gastrointestinal distress.
Notably, a clinical study conducted by Jang et al. in Korea revealed an inverse cor-
relation between total protein intake and the diversity of the gut microbiota, showing
that the athletes in resistance sport who have a high protein diet showed a decrease in
SCFs-producing commensal bacteria [49]. However, another study demonstrated a positive
correlation between a high protein intake and microbial diversity; the gut microbiota of
athletes consisted of 22 phyla of bacteria, while only 11 and 9 phyla were found in the
low and high BMI groups [44]. It is worth noting that Korean athletes did not meet the
recommended dietary fiber intake (≥25 g/day; median intake in bodybuilders 19 g/day,
endurance athletes 17 g/day) [49]. In contrast, Irish rugby players’ dietary fiber intake met
the recommendation level (median intake 39 g/day) [44]. Undigested dietary fiber serves
as an essential energy and carbon source of gut microbiota, contributing to its diversity
and acting as a substate for SCFA synthesis. Therefore, it can be inferred that combining
a high-protein diet with low-dietary-fiber diet may have detrimental effects on the gut
microbiota composition. This finding suggests that dietary fiber also plays an important
role in the high-protein diet. Further investigations are warranted to ascertain whether
alterations in SCFA levels serve as a pivotal mediator of the favorable physiological effects
associated with a high dietary fiber intake.
In addition to dietary fiber, anthocyanins (ACN) have recently attracted the atten-
tion of many researchers. Amongst some of the dietary patterns mentioned above, fruits
and vegetables are important components, particularly certain fruits that are abundant
in ACN, a subclass of polyphenols responsible for the red–blue–purple pigmentation ob-
served in fruits [156,157]. These bioactive compounds possess potent antioxidant and
anti-inflammatory properties that can effectively modulate the secondary cascade associ-
ated with exercise-induced muscle damage (EIMD) [10,158–161]. Delphinidin and cyanidin
are the most extensively investigated anthocyanins, which also encompass malvidin, pe-
onidin, petunidin and pelargonidin. These compounds exhibit favorable physiological
effects in humans [162]. The bioavailability of ACN in the human intestinal tract is lim-
ited, with only a fraction of the dietary intake being digested and absorbed in the small
intestine. However, this bioavailability can be enhanced through interactions with the gut
microbiota [163]. The sugar moieties of ACN undergo hydrolysis by bacterial enzymes in
the colon, leading to the transformation of aglycone forms into a variety of compounds,
including protocatechuic acid, vanillic acid and gallic acid [164]. According to a study [132],
cyanidin consistently converts into protocatechuic acid, which exhibits multiple protective
functions for muscle health, such as reducing oxidative stress, promoting mitochondrial
Nutrients 2024, 16, 1634 15 of 23

biogenesis, and converting skeletal muscle fibers from type II to type I. These effects on
oxidative stress reduction and mitochondrial biogenesis may have potential benefits for
athlete recovery. Notably, the conversion of skeletal muscle fiber emerges as a promising
research domain, deserving significant attention. In the past, the selection of athletes across
various sports has heavily relied on hereditary factors due to the perception that one’s
skeletal muscle type is genetically predetermined and difficult to change through training.
With advancements in our understanding of the skeletal muscle fiber conversion, as well as
potential nutritional strategies, the process of athlete selection may become more adaptable.
However, humans still cannot make genetic changes. This means that while a certain
genetic hereditary factor, such as alpha actinin-3 gene (ACTN3), plays a decisive role in
skeletal muscle fiber conversion, dietary patterns could be utilized as a helpful strategy to
improve it [165].
However, the specific bacterial taxa responsible for the transformation of anthocyanins
into protocatechuic or gallic acid remain unknown. The bacterial enzymes involved in ACN
hydrolysis may be present in several taxa of the genera, such as Bacteroides, Clostridium and
Eubacterium [26,163]. Furthermore, different microbiota compositions may be associated
with distinct pathways of ACN biotransformation, potentially leading to diverse effects
ranging from beneficial to unknown outcomes [166,167], implying that the interaction
between ACN and the gut microbiota could vary among individuals. Therefore, further
research is needed to investigate individual differences in ACN metabolism and its potential
health-promoting effects.

5.2. Practical Application

As mentioned earlier, different dietary patterns affect sports performance in different
ways. Athletes should choose the appropriate dietary pattern on the basis of their actual
situation during training. Athletes who need to control their weight strictly during com-
petition in heavy sports, athletics and gymnastics may consider a ketogenic diet, which
would enable them to lose weight in a short time, but this dietary pattern also has limita-
tions; it is not suitable for enhancing strength in weight lifters or high-intensity cyclists,
for example [168]. In terms of the plant-based diet, current evidence supports that this
diet does not have a significant impact on sports performance, but as mentioned before,
the special micronutrients in the plant-based diet have anti-inflammation and antioxidant
effects to a certain extent, and it would be friendly to vegan athletes [169]. For athletes who
seek to gain muscle mass and strength, such as bodybuilders, the high-protein diet is a
good choice, because it is necessary to generate more muscle protein and prevent lean mass
losses during the periods that restrict energy intake to promote fat loss [170]. Compared
with other dietary patterns, the Mediterranean diet may be more suitable for most athletes;
both aerobic and anaerobic athletes can select this dietary pattern, whose strengths are
that it is rich in foods that can support high energy demands and that it can provide the
antioxidants, essential vitamins and minerals that promote recovery [3]. In practice, these
dietary patterns are used alternately or in a certain period of time, because any special
dietary patterns used for a long time will cause adverse reactions [3,98].

6. Conclusions
In recent decades, it has been increasingly acknowledged that the gut microbiota plays
an important role in human health and sports performance. As mentioned earlier, the
impact of various dietary patterns on the gut microbiota and their subsequent effects on
sports performance may vary. Therefore, further evidence is required to substantiate the re-
lationship between different dietary patterns and their components with the gut microbiota
and sports performance. In addition, it should be noted that diet is inseparable from the
host; it is challenging to strictly disentangle exercise from daily diet during an experiment,
as the individual contributions of each participant are difficult to isolate and assess. To
date, there remains a dearth of research investigating the intricate interplay between diet,
exercise, and the gut microbiota. Additionally, the responses of the gut microbiota to diet
Nutrients 2024, 16, 1634 16 of 23

may vary among individuals, indicating that the formulation of diet regimens should
shift from standardized diet guidelines to flexible recommendations tailored to individual
preference and local customs, and the regular reassessment of these dietary regimens is
essential. Moreover, the significance of nutrients or compounds in diets that have tradition-
ally been regarded as non-nutritive cannot be disregarded, necessitating an exploration
into whether these nutrients exert their effects independently or synergistically. Future
research should focus on personalized nutrition strategies for different populations and the
combined effects of different nutrients. The aforementioned findings will contribute to a
comprehensive understanding of the intricate interplay among exercise, diet, and human
health, which has implications not only for athletes’ well-being but also for that of the
general population.
Key Points
• The interactions between exercise and the gut microbiota play a role in the sports
performance of athletes.
• The ketogenic diet, plant-based diet, high-protein diet, and Mediterranean diet may
improve sports performance from different aspects.
• The gut microbiota and its metabolites play an important role in the effects of dietary
patterns on sports performance.

Author Contributions: Conceptualization, Y.C. and K.Y.; methodology, Y.Z.; software, M.X.; valida-
tion, Y.C., K.Y. and Y.Z.; formal analysis, J.L.; investigation, Y.L.; resources, Y.-H.M.; data curation,
X.W.; writing—original draft preparation, Y.C.; writing—review and editing, Y.-H.M.; visualization,
M.X.; supervision, Y.-H.M.; project administration, Y.-H.M.; funding acquisition, Y.-H.M. All authors
have read and agreed to the published version of the manuscript.
Funding: This work was financially supported by the Guangdong Basic and Applied Basic Research
Foundation (Program No. 2023A1515010004), Special Funds in Key Areas of Guangdong Provincial
Department of Education (Program No. 2023ZDZX2035) and National Natural Science Foundation of
China (Program No. 82003434).
Acknowledgments: I thank Wang Minghan for their comments on the manuscript. I thank bioren-
der.com for their permission to quote material protected by copyright.
Conflicts of Interest: The authors declare that they have no known competing financial interests or
personal relationships that could have appeared to influence the work reported in this paper.

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