Variation in the Uncoupling Proteins Genes in Different Sports

Elvira Bondareva

, Olga Parfenteva

and Valentine Son’kin

2,3 c

Institute and Museum of Anthropology, Moscow State University, Mokhovaya st, 11/1, Moscow, Russia

Moscow Center of Advanced Sports Technologies, Sovietskoi armii st, 6, Moscow, Russia

Russian State University of Physical Education, Sports, Youth, and Tourism, Sirenevyi blv. 4, Moscow, Russia

Keywords: UCP, “Thrifty” Genotype, Athletes, Selection, Mitochondria, Metabolism.

Abstract: Uncoupling protein (UCP) genes appear to be promising candidates for studying the effect of 'thrifty'

genotypes on various aspects of modern human life, starting from susceptibility to obesity and

cardiometabolic diseases and ending with sports talent. The study aims at studying directions of selection for

polymorphic systems of the genes UCP1, UCP2 and UCP3 among athletes engaged in various sports. The

study involved 268 people: 197 athletes (males: n=140; females: n=57) and 71 non-athletes as a control group

(males: n=38; females: n=33). Buccal epithelium was used as a sample of biological material. Genomic DNA

isolation and genotyping of samples for polymorphisms of UCP1 (rs1800592), UCP2 (rs660339), UCP3

(rs1800849) were performed at the premises of Lytech (Moscow). Differences in the distribution of genotype

frequencies of the UCP1 and UCP2 genes between the subgroups within the sample of athletes are statistically

significant (χ2 = 21.2 p = 0.006 and χ2 = 24.06 p = 0.002, respectively). Among the athletes representing

various kinds of sports, the subgroups of aerobic, mixed cyclic and team sports demonstrate the directional

selection of carriers of 'thrifty' genotypes of the studied genes. The subgroup of martial arts is characterized

by the opposite direction in selection.

1 INTRODUCTION

Studies in the field of molecular physiology (in

particular, knockout and overexpression of UCP

genes) revealed the main functions of uncoupling

proteins (UCP): heat production, metabolic

acceleration, and reduction of the rate of formation of

reactive oxygen species (ROS) due to the uncoupling

of oxidation and phosphorylation reactions in

mitochondria (Toda, Diano, 2014; Victorino et al.,

2015; Cardoso et al, 2014). UCPs provide controlled

“leakage” of protons from the intermembrane space

of mitochondria and dissipate chemical energy, stored

as a proton gradient, in the form of heat. However, a

high activity of UCPs is accompanied by a decrease

in the efficiency of the aerobic phase of energy

metabolism; i.e., the amount of synthesized ATP

decreases. Mutations that reduce the activity of UCP

allow more protons to pass through ATP synthase and

synthesize more ATP from a similar number of

substrates and oxygen, which, in turn, leads to the

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storage of unspent calories as fat. Thus, a decrease in

the activity of UCPs, on the one hand, leads to an

increase in predisposition to obesity and, on the other

hand, increases the efficiency of aerobic metabolism.

In the organisms of mammals, including humans, the

most common are three members of the family of

uncoupling proteins: UCP1, UCP2, and UCP3. The

first member of the family of uncoupling proteins,

UCP1 (thermogenin), is the key player that provides

heat production in nonshivering thermogenesis.

Thermogenin homologs, proteins UCP2 and UCP3,

have no such an explicit function and, according to

several studies, regulate the formation of reactive

oxygen species (ROS), protect cells from excess free

fatty acids and reduce their lipotoxicity, and affect the

efficiency of metabolism and its cost-effectiveness.

Mutations in the UCP genes, which are examples of

the “thrifty” genotype, are actively studied as

molecular-genetic markers of increased risk of

obesity, type 2 diabetes mellitus, and cardiometabolic

diseases (Flouris et al., 2017). Among all members of

Bondareva, E., Parfenteva, O. and Son’kin, V.

Variation in the Uncoupling Proteins Genes in Different Sports.

DOI: 10.5220/0008066800380046

In Proceedings of the 7th International Conference on Sport Sciences Research and Technology Support (icSPORTS 2019), pages 38-46

ISBN: 978-989-758-383-4

the family of uncoupling proteins, UCP2 is

characterized by the most widespread expression in

the human body (skeletal muscles, internal organs,

and adipose tissue), which is not limited, e.g., to

brown adipose tissue, such as in the case of UCP1.

The genes UCP2 and UCP3 are located side by side

on the same chromosome (Solanes et al., 1997),

which suggests a linkage group at least in the Russian

population. It is also assumed that UCP2 and UCP3

functionally overlap each other (Bouillaud et al.,

2016). The UCP2 gene polymorphism is associated

with excess weight gain, and it is also actively studied

as a factor of success in sports that require high

aerobic abilities of the athletes. The effect of the

UCP2 gene polymorphism on energy metabolism at

rest (Astrup et al., 1999) and during exercise

(Buemann et al., 2001) was studied.

The aim of the work is to study the directions of

selection for polymorphic systems of UCP genes in

various sports and to analyse the associations of

uncoupling protein genes with the indicators of

physical performance of athletes.

2 MATERIALS AND METHODS

The study involved 268 people. The surveyed sample

included 197 (males: n=140; females: n=57) athletes

involved in various sports and 71 non-athletes as a

control group (males: n=38; females: n=33). The

buccal smears were used as a sample of biological

material. Biological material was collected using

universal sterile disposable probes (Changzhou

Chuangjia Medical Appliance Co., Ltd, China). Next,

genomic DNA was isolated from the collected

samples, and for each sample genotyping of

polymorphic loci in the human genome was carried

out: UCP1 (rs1800592), UCP2 (rs660339), UCP3

(rs1800849). Genomic DNA isolation and

genotyping of samples were performed at the

premises of Lytech (Moscow).

The surveyed sample of athletes was divided into

three subgroups, according to the prevailing source of

energy supply for training and competition activities

(Table 1). In turn, the group of sports with

predominantly mixed energy supply was further

divided into three parts: the cyclic sports subgroup,

the martial arts subgroup, and the team sports

subgroup.

Statistical analysis of the data was carried out

using the STATISTICA 8.0 software (StatSoft,

USA). Differences in the distribution of genotypes of

the studied genes in the subgroups of the surveyed

sample were analyzed using a nonparametric criterion

Table 1: General characteristic of the studied sample.

Subgroup Sports Number

Control - 71

Athletes 189

Anaerobic

skating 500, 1000

1500

, sprint

Aerobic

biathlon 5, 7,5 and

10 km, rowing,

skating 5km and

all-around, ski race,

swimming, modern

entathlon

Mixed energy

supply

126

Team sports

football, basketball

Cyclic sports

Short track speed

skating, biathlon 3

Martial arts

sambo, boxing,

judo, taekwondo

(χ2). The total genotype score (TGS) was also

calculated (Williams and Folland, 2008). For this

purpose, we assigned a genotype score (GS) of 1 to 3

to each individual genotype (Table 2), where score 1

is for homozygous combination of the two original

alleles, 2 for heterozygous and 3 for homozygous

combination of “thrifty” alleles.

Table 2: Genotype scores for the UCP’s genes

polymorphism.

UCP1*AA UCP1*AG UCP1*GG

1 2 3

UCP2*CC UCP2*CT UCP2*TT

1 2 3

UCP3*CC UCP3*CT UCP3*TT

1 2 3

The scores for all three genes were then summed

up (Equation 1) and expressed as a percentage of the

maximum possible (∑GSmax=9) amount (Equation

2).

∑ GS (genotype score) = GSUCP1 +

GSUCP2 + GSUCP3

)

TGS% = (100/9) × ∑GS (2

)

∑GS9 = TGS100% = UCP1*GG +

UCP2*TT + UCP3*TT

)

Thus, the maximum sum of scores (Equation 3)

will correspond to the greatest predisposition to

excess weight gain, determined by the three

uncoupling protein systems. Or, in other words, to the

most effective/economical aerobic metabolism. The

Variation in the Uncoupling Proteins Genes in Different Sports

average TGS values in different subgroups of the

surveyed sample were compared using the median

test. All subjects who participated in the study were

informed of the research objectives and methods and

gave their written informed consent.

3 RESULTS AND DISCUSSION

Numerical distributions and frequencies of the

genotypes and alleles of the UCP1, UCP2, and UCP3

genes in the subgroups of the surveyed sample are

presented in Tables 3 to 7 and Figures 1 to 3. The

distribution of genotypes in the control group

conforms to the Hardy-Weinberg equilibrium for all

polymorphic systems studied.

Table 3: Numerical distributions and frequencies of the

UCP1, UCP2, and UCP3 genotypes and alleles in the

athletes’ and control groups.

Genotype

Athletes

N (%)

Control

N (%)

p-value

UCP1*AA

107

(54,3%)

42 (60%)

=7,07

p = 0,03

UCP1*AG 62 (31,5%) 26 (37,1%)

UCP1*GG 28 (14,2%) 2 (2,9%)

UCP2*CC 74 (37,6%) 22 (31,4%)

= 3,69

p = 0,14

UCP2*CT 86 (43,6%) 39 (55,7%)

UCP2*TT 37 (18,8%) 9 (12,9%)

UCP3*CC 93 (47,2%) 44 (62,9%)

= 5,3

p = 0,07

UCP3*CT 82 (41,6%) 21 (30%)

UCP3*TT 22 (11,2%) 5 (7,1%)

Table 4: UCP’s polymorphism genotype and allele

frequencies amongst all participants according to their

subgroup.

Allele Control Athletes

UCP1*A 78,6 70,1

UCP1*G

†

21,4 29,9

UCP2*C 59,3 59,4

UCP2*T

†

40,7 40,6

UCP3*C 77,9 68

UCP3*T

†

22,1 32

† - «thrifty» allele

Table 5: Numerical distributions of the UCP1 genotypes in

the studied subgroups.

Subgroup

Genotype

p-value

AA AG GG

Anaerobic 3 7 3

= 28,9

p = 0,001

Aerobic 26 12 12

team sports 22 14 5

cyclic sports 23 22 4

martial arts 28 4 4

Control 42 26 2

Table 6: Numerical distributions of the UCP genotypes in

the studied subgroups.

Subgroup

Genotype

p-value

CC CT TT

Anaerobic 3 7 3

χ2 = 28,7

p = 0,001

Aerobic 18 28 4

team sports 12 18 11

cyclic sports 15 20 14

martial arts 24 7 5

Control 22 39 9

Table 7: Numerical distributions of the UCP3 genotypes in

the studied subgroups.

Subgroup

Genotype

p-value

CC CT TT

Anaerobic 9 4 0

χ2 = 14,7

p = 0,14

Aerobic 24 20 6

team sports 14 20 7

cyclic sports 25 18 6

martial arts 17 17 2

Control 44 21 5

Table 8: Mean values of the TGS (%) in the studied

subgroups.

Subgroup TGS (%)

Control 51.3

Athletes 56.2

Aerobic 54.9

Anaerobic 56.3

Mixed energy supply 55.9

team sports 55.8

martial arts 52.6

cyclic sports 56.2

Gene mutations that reduce the activity of

uncoupling proteins in the past had allowed for a

more efficient calorie expenditure and fat

accumulation, which gave an advantage to carriers of

such genes in conditions of lack of food or during

long journeys. In the changed circumstances of the

modern world, the 'thrifty' genotype has become a

factor of increased risk of developing metabolic

diseases. However, the presence of 'thrifty' variants of

uncoupling protein genes in the human genome can

now provide an advantage to the carriers of the

original alleles when it comes to sports at an elite

level. Uncoupling proteins belong to a larger family

of mitochondrial anion carrier proteins (MACP).

Three out of five members of this family were found

to have associations with increased susceptibility to

icSPORTS 2019 - 7th International Conference on Sport Sciences Research and Technology Support

fat accumulation and with performance indicators,

that is, UCP1, UCP2 and UCP3. Physiological studies

performed on animal models - knockout and

overexpression of UCPs - suggest that the main

functions are heat production, metabolism

acceleration and reduction of the rate of reactive

oxygen species (ROS) formation (Toda and Diano,

2014; Victorino et al., 2015; Cardoso et al., 2014).

In general, the studied group of athletes

demonstrates significant differences in the

distribution of frequency occurrence of genotypes

from the control group of non-athletes for the UCP1

polymorphic system (Table 3). Carriers of the

UCP1*GG genotype are four times more frequent in

athletes than in the control group. The athletes also

have a higher frequency of 'thrifty' alleles of UCP1

and UCP3 (Table 4). A generalized sample of athletes

is characterized by an increased tendency to

accumulate fat due to the increased frequency of

occurrence of “thrifty” alleles for all studied gene

systems. This actually means that, in general, athletes

are characterized by lower mitochondrial uncoupling,

which provides a more efficient and economical

version of the aerobic phase of ATP synthesis.

Despite the general direction of selection for all

molecular genetic markers identified in the

generalized group of athletes, the sample is not

homogeneous. This group includes athletes who

practice sports with different energy supply

requirements and different motor activities (cyclic,

acyclic, and precise). The function of uncoupling

proteins allows us to reasonably assume that

directional selection of “thrifty” genotypes will be

observed most in those sports which require a high

level of development of the aerobic component of

physical performance. Therefore, an attempt was

further made to study the directions of genetic

selection in subgroups of athletes formed according

to the principle of the prevailing source of energy

supply for training and competitive activity (Table 1).

The sports which have extremely high requirements

on the aerobic qualities of an athlete are included in

the subgroup of “aerobic” sports. In contrast, we

singled out certain sports types and specializations

requiring mainly strength, speed and power qualities,

which formed a subgroup of 'anaerobic' sports. Those

sports types and/or specializations, which are

characterized by a combination of energy supply

sources, or predominantly glycolytic pathway of ATP

synthesis, entered the group with a mixed power

supply: cyclic, team and martial arts. Thus, the group

of athletes was divided into 5 subgroups (Table 1).

With a certain degree of assumption, we can say that

the aerobic subgroup represents long-distance

runners, the anaerobic one – sprinters, and the mixed

sports subgroup consists of middle-distance runners.

The analysis of the frequency of occurrence of

genotypes in the subgroups within the sample of

athletes revealed significant differences in the

distribution of genotypes for the polymorphic

systems of the UCP1 and UCP2 genes (Tables 3 to

7).

Differences in the distribution of genotype

frequencies of the uncoupling protein 1 gene between

the subgroups within the sample of athletes are

statistically significant (χ2 = 21.2 p = 0.006).

Compared to the control group, all subgroups within

the sample of athletes demonstrate an increase in the

number of carriers of the G allele of the UCP1 gene,

which is associated with an increased risk of obesity

and a decrease in uncoupling.

Figure 1: Frequencies (%) of genotypes of the UCP1 gene

in all subgroups within the surveyed sample.

This result is a logical one in terms of the

increased energy efficiency of aerobic metabolism

and the advantage of athletes carrying the 'thrifty

allele' over the carriers of the 'normal' level of

uncoupling. G allele carriers were reported to burn

200 Kcal/day less than the original A allele carriers

(Kogure et al., 1998), and this allele also leads to a

decrease in UCP1 mRNA in BAT adipocytes

(Esterbauer et al., 1998). All this suggests that the

allele -3826G UCP1 increases the efficiency of

energy metabolism in BAT (brown adipose tissue)

mitochondria. This genotype is most frequent in the

subgroup of aerobic sports requiring a very high level

of aerobic capacity. Further, directional selection of

GG*UCP1 carriers is observed due to the

simultaneous decrease in the frequencies of

occurrence of AA*UCP1 and AG*UCP1 (Fig.1). The

subgroup of anaerobic sports is also characterized by

a high frequency of occurrence of GG*UCP1;

however, it should rather be referred to the selection

100

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UCP1*AA UCP1*AG UCP1*GG

Variation in the Uncoupling Proteins Genes in Different Sports

of heterozygous genotype carriers. The high (as

compared to the control group) frequency of

occurrence of genotypes containing at least one G

allele found in the subgroup of anaerobic sports may

indicate the existence of mechanisms supporting a

high level of anaerobic performance. The molecular

physiological basis for such mechanisms is

uncoupling processes in BAT. The result may also be

related to the small number of subjects in the

subgroup of anaerobic sports.

The subgroup of martial arts demonstrates the

opposite direction of selection - AA*UCP1, i.e.

selection for the uncoupling of energy processes,

although this subgroup is characterized by an increase

in the number of carriers of the GG*UCP1 genotype.

The results of physiological studies on the effect of

BAT activity on athletes’ physical performance

suggest that a sufficient level of uncoupling of

oxidative phosphorylation processes in BAT

adipocytes, which is determined by the A allele of the

UCP1 gene, ensures rapid utilization of lactate

(Son’kin et al., 2014; Merla et al., 2010), which is

formed in large quantities by muscle activity typical

for martial arts. Thus, the presence of BAT

adipocytes with the original level of uncoupling in

athletes involved in martial arts allows them to both

effectively counter acidification of skeletal muscles

during the fight and quickly dispose of accumulated

lactate between fights, which ensures rapid recovery

of athletes. In other words, the selection of alleles

observed in the martial arts subgroup which

determine sufficient uncoupling of oxidative

phosphorylation is apparently connected with the

processes of recovery during and between fights. The

greater (as compared with the control group) number

of GG*UCP1 genotype carriers in combat athletes

may indicate the existence of a minor variant of

selection, i.e. selection of combatants with high

aerobic capabilities, who reach a high professional

level due to the fight strategy based not on the speed

and strength qualities, but on athlete's endurance.

Significant differences were found between the

subgroups within the sample of athletes for the

polymorphic system of the UCP2 gene (χ2 = 24.06 p

= 0.002) as well. The most pronounced differences in

the distribution of occurrence frequencies of

genotypes from the control group are characteristic of

the subgroup of martial arts athletes (Fig.2). This

group shows a pronounced selection of carriers of two

original alleles of the UCP2 gene – CC*UCP2 – due

to a decrease in the number of carriers of the

heterozygous genotype. At the same time, the

proportion of TT*UCP2 genotype carriers is identical

to that in the group of non-athletes. The direction of

selection for the UCP2 system is similar to UCP1

and, in terms of biochemistry, corresponds to the

normal level of uncoupling not only in BAT

adipocytes, but also in most internal organs.

Subgroups of team and cyclic sports which, according

to the type of energy supply, are in an intermediate

position between anaerobic and aerobic sports,

demonstrate the selection of carriers of the TT*UCP2

genotype, due to a decrease in the proportion of

heterozygotes. The result suggests that in such sports

as basketball, football, middle-distance running

advantage is gained by athletes with a more

economical type of energy supply mainly in internal

organ tissues. At the same time, the proportion of

CC*UCP genotype carriers is almost identical to the

control group. The anaerobic sports subgroup

demonstrates the selection of carriers of the UCP2 T

allele, due to a decrease in the proportion of

CC*UCP2 carriers. This UCP2 allele determines a

more efficient metabolism, which affects energy

expenditure during exercise (Beumann et al., 2001;

Kimm et al., 2002). Previously, the selection of

T*UCP2 (55Val) and T*UCP3 allele carriers was

reported in the group of Russian long-distance

runners (Ahmetov et al., 2009). The aerobic sports

subgroup, unlike all other sports, shows a decrease in

the proportion of TT*UCP2 carriers, with a

corresponding increase in the proportions of CT and

CC. It is likely that the presence of athletes with these

genotypes is associated with their functional

characteristics. It was reported that athletes with a

heterozygous genotype have the highest rates of

maximum oxygen consumption (Bondareva et al.,

2018). A high-fat diet stimulates UCP2 and UCP3

gene expression in athletes' muscles. And the most

significant increase in the concentration of UCP2 and

UCP3 mRNA was found in muscle fibers of type IIA

(fast oxidative-glycolytic fibers), which have a

greater metabolic plasticity in terms of choice of

oxidation sources (Schrauwen et al., 2001) and are

also characteristic of the skeletal muscles of athletes

in the subgroup of mixed energy supply.

We have found no statistically significant

differences in the distribution of genotype

frequencies of the UCP3 gene either between the

control group and the subgroups within the sample of

athletes, or between the subgroups within the sample

of athletes (χ2 = 9.08 p = 0.33). However, the results

would suggest some trends in the direction of

selection in different subgroups of athletes. Thus, the

selection of carriers of the UCP3 T allele – CT*UCP3

and TT*UCP3 genotypes – occurs in the subgroups

aerobic, team and cyclic sports, due to a cor-

icSPORTS 2019 - 7th International Conference on Sport Sciences Research and Technology Support

Figure 2: Frequencies (%) of genotypes of the UCP2 gene

in all subgroups within the surveyed sample.

responding decrease in the proportion of carriers of

the CC*UCP3 genotype. A similar result was

obtained when studying the distribution of UCP3

genotypes in subgroups of Italian sprinters and long-

distance runners compared to the control group (Sessa

et al., 2011). The anaerobic sports subgroup

demonstrates the selection of the CC*UCP3 carriers.

No carriers of the two mutant alleles were found,

possibly due to the advantage of carriers of a normal

level of uncoupling in skeletal muscles, or because of

the small number of subjects in the group. UCP3 is

expressed in skeletal muscles and myocardium,

which makes it a promising marker of physical

performance. The UCP3 T allele is associated with a

more economical calorie expenditure for ATP

synthesis at the aerobic stage of energy metabolism,

which makes it an 'aerobic capacity allele'. Earlier we

demonstrated the selection of T allele carriers in a

group of football players (Bondareva et al., 2016).

Therefore, the selection of UCP3 T allele carriers,

which is observed in all subgroups except the

anaerobic one, is logical. For successful training and

competitive activities of the athletes included in these

subgroups, the development of medium and high

aerobic capabilities is necessary. And advantage is

gained by the carriers of alleles allowing for the

development of higher and/or more effective

mechanisms for the aerobic synthesis of ATP, in the

skeletal muscles and heart. In the subgroup of martial

arts athletes, a directional selection of carriers of the

heterozygous genotype of CT*UCP3 was found, due

to a decrease in the proportion of both homozygous

genotypes. Martial arts athletes need to combine a

fairly high level of aerobic capabilities, extremely

high speed, strength and power qualities for rapid

attacks and throws, as well as high resistance to tissue

hypoxia and acidification of skeletal muscles.

Therefore, the heterozygous combination of alleles

makes it possible to find a compromise between

improving the energy efficiency of muscle

contraction and protection against oxidative stress.

Figure 3: Frequencies (%) of genotypes of the UCP3 gene

in all subgroups within the surveyed sample.

Regular exercise and weight training reduce the

level of UCP3 mRNA and protein in mitochondria,

which, in turn, is inversely proportional to BMD. All

of this contributes to increasing energy efficiency in

exercise (Schrauwen et al., 2005). UCP3 exports fatty

acid (FA) anions from the mitochondrial matrix

preventing accumulation of excess FA in the matrix

(Wang et al., 2003) and freeing CoA for continued β-

oxidation of FA, thus avoiding oxidative stress and

maintaining a high metabolic rate. One of the

consequences of the increased concentration of

uncoupling proteins 2 and 3 in cardiomyocytes, in

addition to the increased uncoupling of oxidative

phosphorylation, is the predominant use of FA as a

substrate for the oxidation and ATP synthesis

(Harmancey et al., 2013). Presumably, UCP3 is the

main regulator for the influx of energy substrates into

mitochondria, since its expression is controlled by

fatty acids, which is likely to play an important role

in the energy supply of sports with a predominantly

aerobic type of energy supply. Because during

prolonged loads, typical for aerobic and team sports,

fats are the main energy substrate. The observed

selection of carriers of CT*UCP3 and TT*UCP3

genotypes may be associated with increased energy

efficiency in skeletal muscle in combination with the

ability to maintain a sufficient rate of entry of FA into

the mitochondria of skeletal muscle. The results of

animal studies suggest that one of the main functions

of uncoupling proteins is to accelerate the metabolism

and protect the cell from ROS. ROS have a damaging

effect on both cellular and organism levels, since

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UCP2*CC UCP2*CT UCP2*TT

100

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UCP3*CC UCP3*CT UCP3*TT

Variation in the Uncoupling Proteins Genes in Different Sports

uncoupling proteins are present in the mitochondria

of all organs and tissues. Significant oxidative stress

is observed during intense training and competition in

martial arts athletes (Finaud et al., 2006). Cell

survival depends on the amount of ROS: low levels

of ROS trigger a cascade of reactions aimed at cell

survival (Tait and Green, 2012), while high levels of

ROS, which cannot be neutralized by means of cell

antioxidant protection, lead to cell damage and death

(Orrenius et al., 2007). The formation of ROS in

mitochondria is reduced due to the soft or strong

uncoupling of the electron transport chain of

mitochondria and ATP synthase by means of UCPs.

However, strong uncoupling leads to a lack of ATP in

the cell, which causes necrosis. It is presumed that

UCPs are able to “gently” uncouple oxidative

phosphorylation reactions without reducing the

amount of ATP (Sluse, 2012), but physiological

studies are still insufficient (Shabalina and

Nedergaard, 2011). Thus, a decrease in the activity of

uncoupling proteins, on the one hand, makes it

possible to increase the efficiency of the aerobic stage

of energy metabolism, but the concentration of ROS

increases simultaneously.

The above-mentioned directions of genetic

selection for gene systems of uncoupling proteins

were sometimes difficult to explain and differently

directed for some groups of sports. However, all

genetic factors act in the body and affect its

phenotypic characteristics simultaneously. Therefore,

we analysed the directions of selection in the formed

subgroups of the surveyed sample simultaneously for

all selected polymorphic systems. The results of

studies on the effect of several genes on various

phenotypic characteristics convincingly prove that

the tendency to obesity is proportional to the number

of risk genotypes/alleles present in the genome. The

higher the TGS, the greater the number of

alleles/genotypes that determine the accumulation of

excess weight, and in the case of uncoupling proteins,

this also means a more efficient ATP synthesis. Table

8 presents the average TGS calculated for the

subgroups of the surveyed sample. Comparative

analysis of average TGS revealed non-random

statistical differences (χ2 = 9.86 p=.02). The lowest

value (TGS=51.3%), which corresponds to the

smallest number of 'thrifty' alleles/genotypes, is

demonstrated by the test subjects included in the

control group. Non-athletes are predominantly

carriers of the original alleles of uncoupling proteins,

which determine the normal level of uncoupling of

oxidative phosphorylation in mitochondria, which

means a small risk of obesity. All subgroups within

the sample of athletes show higher TGS than the

control sample, which on the one hand is a sign of the

accumulation of uncoupling protein

alleles/genotypes, which increase the coupling of

oxidative phosphorylation reactions in mitochondria

and provides increased energy efficiency of the

aerobic phase of ATP synthesis, but simultaneously

increases the risk of obesity development. However,

due to the increased coupling of reactions of the

aerobic phase of metabolism, athletes have a reduced

potential in protecting against ROS and oxidative

stress. For different types of wrestling, a high

glycolytic load and a high level of oxidative stress

during competitions affect the selection of athletes

capable of quickly disposing of lactate and effectively

resisting peroxidation, both during and between

fights. These properties, which are partially

determined by uncoupling proteins, are leading and

provide high sports achievements in martial arts.

Therefore, for some systems of uncoupling proteins –

UCP1 and UCP2 – the selection of carriers of the

original alleles, rather than “thrifty” ones, occurs,

opposite to the other subgroups of athletes.

Previously, for another polymorphic system, the

selection of carriers of alleles of resistance to tissue

hypoxia among sambo athletes has already been

identified (Bondareva and Godina, 2016).

4 CONCLUSIONS

Selection of carriers of 'thrifty' genotypes among the

majority of subgroups of athletes allows to conclude

that for most sports types and specializations, a more

efficient metabolism is a factor that positively affects

the success in sports. It should be noted that to a lesser

extent the selection of carriers of 'thrifty' genotypes is

characteristic of martial arts athletes (TGS=52.6%).

ACKNOWLEDGEMENTS

The reported study was funded by RFBR according

to the research projects №№ 18-59-94015 and 17-26-

03004-OGN

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