The Acute Administration of Carnosine and Beta-Alanine Does Not Improve Running Anaerobic Performance and has No Effect on the Metabolic Response to Exercise

doi:10.4236/ape.2013.34028

Paper Menu >>

Journal Menu >>

Advances in Physical Education

2013. Vol.3, No.4, 169-174

Published Online November 2013 in SciRes (http://www.scirp.org/journal/ape) http://dx.doi.org/10.4236/ape.2013.34028

Open Access 169

The Acute Administration of Carnosine and Beta-Alanine Does

Not Improve Running Anaerobic Performance and has No Effect

on the Metabolic Response to Exercise*

Pietro Luigi Invernizzi1, Stefano Benedini1,2#, Sandro Saronni3,

Giampiero Merati1, Andrea Bosio4

1Department of Biomedical Sciences for Health, Università degli Studi di Milano, Milano, Italy

2Research Centre of Metabolism, San Donato Milanese, Milan, Italy

3School of Exercise Sciences, Università degli Studi di Milano, Milan, Italy

4Human Performance Laboratory, MAPEI Research Centre, Olgiate Olona, Varese, Italy

Email: #[email protected]

Received July 26th, 2013; revised August 26th, 2013; accepted September 4th, 2013

Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium,

provided the original work is properly cited.

An increase in muscle carnosine content, following its chronic supplementation, has been shown to im-

prove anaerobic performance. In addition, carnosine can affect plasma glucose concentration and insulin

response. However, it is not clear whether the acute ingestion of carnosine can have the same effects. Aim

of this study was to investigate the acute effects of carnosine ingestion on anaerobic intermittent per-

formance and the responses of blood insulin, glucose, bicarbonate and lactate concentrations to exercise.

Twelve healthy, young, active participants (BMI 23.5 ± .6, age: 22 ± 2 years) underwent in two separate

occasions (double-blind, randomized, crossover design) the running-based anaerobic test (RAST), con-

sisting of 6 × 35-m sprints interspersed with 10 s rest after acute (4 hours before the test) ingestion of ei-

ther 1 g of L-carnosine and 1 g of β-alanine or placebo. None significant difference was found between

the acute ingestion of carnosine and the placebo conditions in terms of running performance (30.0 ± .8

and 29.8 ± .8, p = .302), perceptual response to exercise (RPE), blood lactate, insulin (23.8 ± 13.0 and

19.5 ± 9.0 μU·ml−1, p = .329), blood glucose (109 ± 23 and 104 ± 12 mg·dl−1, p = .969), and blood bicar-

bonates (16 ± 2 and 16 ± 2 mEq·l−1, p = .277). In conclusion, the acute ingestion of carnosine had no ef-

fect on performance, perceptual response to exercise, blood lactate concentration, insulin, glucose, and

bicarbonates responses to exercise compared to a placebo treatment. It is not clear whether these results

may be attributed to an insufficient dose of carnosine or to a lack of acute effect per sé.

Keywords: Carnosine; Performance; Administration; Metabolic Response; Running

Introduction

The dipeptide carnosine is found in various tissues, espe-

cially in skeletal muscle (Hobson et al., 2012; Harris et al.).

Carnosine is recognized to have, in addition to antioxidant

properties (Alhamdani et al., 2007), buffering effects on acido-

sis (Hobson et al., 2012), as well as stimulating actions on the

immune system and on various neurotransmitters (L-carnosine

lowers neural activities of sympathetic nerves and facilitates

those of parasympathetic nerves) (Nagai et al., 2012). Although

carnosine is synthesized endogenously, its tissue concentration

is influenced by diet (Baguet et al., 2010). Indeed, exogenous

supplementation of β-alanine (precursor and limiting factor in

the synthesis of carnosine) has been shown to increase muscle

carnosine concentration in vitro (Margolis et al., 1985) and in

animal models (Hill et al., 2007).

At physiological pH, carnosine exerts a strong buffering ac-

tion (stabilization of the degree of acidity) that is of fundamen-

tal importance during muscle activity, usually associated with

an acidification of the intracellular compartment (Smith, 1938).

Carnosine is found in muscles of both type I and type II, but

its concentration is highest in type II fibres (Baguet et al., 2010).

Carnosine levels seem to be correlated to exercise performance:

Suzuki et al (Suzuki et al., 2002) observed a positive correlation

between carnosine content in the vastus lateralis and the power

generated at the end of a 30 second cycle ergometer test (Win-

gate test) in untrained men. Hill et al. (Hill et al., 2007) demon-

strated that β-alanine administration to active men can produce

a significant increase of carnosine levels in skeletal muscle,

which was related to an improvement of exercise performance.

In particular, β-alanine was administered to untrained men for

10 weeks, producing not only a remarkable increase of car-

nosine content in the vastus lateralis, but also a significant pro-

longation of the time for exhaustion during a cycling test per-

formed at 110% of energy production and maximal heart rate,

where the endurance time was estimated to be about 2.5 min-

utes (Hill et al., 2007). The role played by carnosine, or by β-

*The authors declare no conflicts of interest that are directly relevant to

the content of this article.

#Corresponding author.

P. L. INVERNIZZI ET AL.

alanine, is of interest not only for untrained subjects: the two

compounds have a positive effect on strength in trained athletes,

especially sprinters, rowers and body-builder as well (Baguet et

al., 2010; Derave et al., 2010). Recent studies have described

other biological roles for carnosine as an antioxidant, antidia-

betic (reduction of glycosylation) (Derave et al., 2010; Hipkiss,

2009), or anti-aging (elongation of chromosomal telomeres)

agent (Shao et al., 2004). In particular, since carnosine content

in the diaphragm of streptozotocin (STZ)-diabetic rats was

lower than in wild type rats (Buse et al., 1980), it was hypothe-

sized that carnosine may be involved in the control of glucose

metabolism. In animal models peripheral administration of a

small amount of carnosine (.005 to 5 nmol/300 g body weight

[BW] by intraperitoneal injection [IP]), or administration of a

diet containing .001% carnosine, reduced 2 DG-hyperglycemia,

producing an increase in plasma insulin levels, a decrease in

plasma glucagon levels, and suppression of the activity of sym-

pathetic nerves innervating the adrenal glands and liver (Nagai

et al., 2003; Yamano et al., 2001). In addition, in animal models

of diabetes, the ability of carnosine to inhibit the adrenergic

system, with positive effects on glucose metabolism, was dem-

onstrated after both chronic and acute supplementation (Aldini

et al.).

Beta-alanine is usually administered orally for a period of 1 -

4 weeks (Jagim et al.), since chronic assumption seems to be

necessary to ensure carnosine increase inside the muscle fibres

(Derave et al., 2007). However, some evidences suggest that

few hours after the ingestion of carnosine the dipeptide is al-

ready available and ready to exert its function inside the muscle

fibers (Begum et al., 2005; Gardner et al., 1991). If this as-

sumption was valid, it would be interesting for athletes who

perform high intensity exercise because they might avoid the 3

- 4 weeks periods of β-alanine supplementation. The only study

on the acute effect of carnosine on performance failed to show

any improvement due to the ingestion of carnosine (Kraemer et

al., 1995). However, it is likely that the dose administered to

the participants was too low.

In a preliminary study (di Pierro et al., 2011) it has been

shown that the oral ingestion of l g of carnosine few hours be-

fore a heavy training session in volleyball players might have

decreased the blood lactate production and increased the spon-

taneous total amount of work.

Furthermore, other than the intracellular pH buffering capac-

ity, the increase in Ca2+ sensitivity of the contractile fibres

represents an alternative mechanism through which muscle

carnosine can improve muscle performance (Dutka & Lamb,

2004). This ergogenic effect of muscle carnosine might be use-

ful in those maximal efforts where the low levels of pH do not

represent the limits for the maintenance of the contractile prop-

erties of the muscles.

The aim of this study was to evaluate whether the acute in-

gestion of 1 g of L-carnosine and 1 g of β-alanine can improve

performance during a running anaerobic sprint test (RAST)

(Zagatto et al., 2009) compared to a placebo condition. Fur-

thermore, the response of blood insulin, glucose, bicarbonate

and lactate concentrations to the RAST, following the acute

ingestion of carnosine, was also investigated.

A recent meta analysis (Hobson et al., 2012) showed that the

supplementation of β-alanine can improve exercise perform-

ances with a duration comprised between 60 and 240 seconds.

However, some Authors (Suzuki et al., 2002) suggested that

towards the end of a 30-sec Wingate test, a higher content of

muscle carnosine may be important for the preservation of a

higher power output compared to a control condition. The use

of the RAST (due to its intermittent feature) could be a better

choice to investigate whether carnosine can increase exercise

performance towards the end of an effort lasting about 30 sec-

onds. We hypothesize that, at least, during the last 35 m-sprint

of the RAST, carnosine can have a beneficial effect on the final

time compared to a placebo condition.

Materials and Methods

Subjects

Participants were recruited in the Milan area in Italy, ac-

cording to the following inclusion criteria: males aged between

20 and 30 years old, no smokers, absence of chronic diseases or

chronic drug treatment. All subjects signed a written informed

consent prior to participation according to the Declaration of

Helsinki. All the procedures used complied with the Good

Clinical Practice (GCP) principles.

Twelve healthy male subjects were enrolled (age: 22 ± 2

years, body mass: 74.1 ± 7.1 Kg, stature: 178 ± 5 cm, body

mass index: 23.5 ± .6 kg/m2): all participants were on a stable

diet and had normal glucose tolerance (according to ADA).

They were physically active and used to exercise on average 4

± 2 times a week for 2.0 ± .6 hours and 2 ± 2 times a week for

1.6 ± .8 hours at vigorous and moderate intensities, respec-

tively. The participants recruited for this study were soccer

players, basketball players and track and fields athletes, accus-

tomed with repeated all out intermittent sprints. On a separate

day, prior to the beginning of the data collection, a familiariza-

tion session with the RAST used in the present study was done.

Study Desi gn

The study was conducted according to a double blind, ran-

domized, crossover, counterbalanced, placebo controlled design.

The randomization, the preparation of treatment and placebo

tablets were done by person not involved in the study. The

treatment and placebo conditions were revealed to the authors

of the present study only after the conclusion of the statistical

analysis.

All participants underwent two RAST on two different occa-

sions: on the first day, half of the participants (selected by a

computer-assisted simple randomization procedure) took 4

tablets containing 250 mg of L-carnosine + 250 mg of β-

alanine (DDM Carnosina, Omeopiacenza s.r.l., Pontenure—PC,

Italy) 4 hours before the test, whereas the other half of partici-

pants took 4 tablets of placebo (containing no active ingredients

but with the same appearance of L-carnosine and β-alanine

tablets), and vice versa on the second day of test (2 weeks

apart). Two weeks seems to be a more than sufficient washout

period when carnosine is acutely administered to human sub-

jects (Suzuki et al., 2006). The dose of carnosine administered

to the participants was similar to a previous study (di Pierro et

al., 2011) and in accordance to the prescriptions of the manu-

facturer.

The Running-Based Anaerobic Sprint Te st (RAST)

A standardized 10 min warm-up, consisting of 5 min running

at a moderate pace followed by 5 accelerations over a distance

of 40 m, interspersed with 1 min of passive recovery, was com-

pleted by each participants prior to the beginning of the RAST,

Open Access

170

P. L. INVERNIZZI ET AL.

followed by five min recovery.

Starting from a standing position, on a track and field 400 m

track, each participant sprinted all-out over a distance of 35 m

for six times (forth and back) interspersed by 10 seconds of

passive recovery. Performance time was electronically recorded

using photocells gates at a height of 1.3 meters and posi-

tioned .5 meters ahead of the start line. The 35 m were meas-

ured as the distance between the two photocells gates. During

all the sprints the athletes were verbally encouraged by the

researchers, in order to obtain the maximal performance. The

RAST has been shown to be a valid and reliable (for Total Ef-

fort Time ICC = .90) test for assessing both anaerobic power

and predicting short-distances performances in running (Za-

gatto et al., 2009).

The outcome measures were a) time to complete each section

of 35 m b) total time (the sum of the six partial times over the

35 m sections), c) ratings of perceived exertion (RPE) and mus-

cle pain (PAIN) immediately after the completion of the test

using the validated CR 10 Borg scale (Borg, 1998). Thirty min-

utes after the end of the RAST test RPE and PAIN were as-

sessed again. RPE collected at this time point was used to as-

sess the internal load through the session RPE-based method

(Foster et al., 2001; Impellizzeri et al., 2004). Finally, the de-

layed onset of muscle soreness (DOMS) was evaluated at 24

and 48 hours post RAST.

Blood Analysis

Blood samples for lactate evaluation was collected from an

ear lobe at 3, 5, 7, and 9 minutes after the completion of the test

and analyzed using a portable blood lactate enzymatic analyzer

(GM-7, Analox Instruments, Hammersmith, London, UK). Two

different blood samples were drawn from an antecubital vein

two hours prior to the beginning of the RAST test and 1 minute

after the completion of the test, respectively, for the analysis of

blood glucose, insulin and bicarbonates.

All blood samples (20 ml each) were collected in test tubes

containing EDTA and placed on ice until plasma or serum were

separated by centrifugation at 4˚C (within 1.5 hours from sam-

pling). All plasma and serum aliquots were frozen at −60˚C for

later analysis. All assessments were carried out in duplicate.

Plasma glucose was measured with a glucose analyzer (Beck-

man Instruments, Fullerton, CA). Free-insulin was assessed by

a highly specific two-site monoclonal antibody-based immu-

nosorbent assay (ELISA; Dako Diagnostics, Cambridgeshire,

UK).

Aliquots of blood were taken before centrifugation for meas-

urement of bicarbonate, enzymatically determined using a

commercial kit.

Statistical Analysis

The normality of data distribution was preliminary checked

by the Shapiro-Wilk’s test. Data are expressed as mean ± Stan-

dard Deviation (SD). Differences between the two treatments

(carnosine and placebo) in terms of performance (total time

given by the sum of each 35 m run), perceived exertion and

pain at the end of the test, and session RPE at 30 minutes after

the completion of the test, were tested using the Student paired

t-tests.

A series of two way (condition × time) fully repeated meas-

ures ANOVAs were used to test the differences in glycemia,

insulin and bicarbonates levels prior to and after the RAST test

(PRE and POST test, main factor time) in the two conditions

(treatment with carnosine or placebo, main factor condition).

A series of two way (condition × time) fully repeated meas-

ures ANOVAs were used to test the differences between the

two conditions (treatment with carnosine or placebo, main fac-

tor condition) either for the times to complete each bout of the

RAST and the time course of blood lactate after the RAST

(main factors time).

Differences in DOMS between the two conditions (treatment

with carnosine or placebo, main factor condition) at 24 and 48

hours after the RAST (main factor time) were tested by mean of

a two way (condition × time) fully repeated measures ANOVA.

Significant differences detected by ANOVAs were adjusted

using the Bonferroni method.

The level of significance was set a priori at α < .05.

Results

None of the participants reported any side effects due to the

administration of 1 g of Carnosine + 1 g of β-alanine.

No significant differences were found in performance and

RPE, pain, and session RPE immediately and 30 minutes after

the RAST respectively (Table 1).

After the RAST test blood insulin significantly increased

(main factor time, p < .001) both in the carnosine (pretest 6.0 ±

2.5 vs posttest 23.8 ± 13.0 μU·ml−1) and placebo condition

(pretest 5.2 ± 2.0 vs posttest 19.5 ± 9.0 μU·ml−1) but without

significant interaction (condition × time, p = .329). Similarly,

after the RAST blood glucose significantly increased (main

factor time, p < .001) both in the carnosine (pretest 79 ± 9 vs

posttest 109 ± 23 mg·dl−1) and placebo condition (pretest 74 ±

10 vs posttest 104 ± 12 mg·dl−1) but without significant interac-

tion (condition × time, p = .969).

Despite no significant interaction (condition × time, p = .277),

blood bicarbonates showed in both conditions (carnosine pre

test 30 ± 2 vs post test 16 ± 2 mEq·l−1 and placebo pre test 29 ±

2 vs post test 16 ± 2 mEq·l−1) a significant decrease (main fac-

tor time, p < .001).

During the RAST the time to complete each bout of 35 m

progressively increased in the two conditions (main factor time,

p < .001) but without significant interaction (condition × time,

p = .855) (Figure 1).

No significant interaction (condition × time, p = .509) was

detected for the blood lactate response monitored after the com-

pletion of the RAST (Figure 2).

Table 1.

The acute effects of carnosine administration, compared to a placebo

treatment, on the running-based anaerobic sprint performance (RAST),

rating of perceived exertion (RPE), pain, session RPE, blood glucose,

insulin and bicarbonate. Data are mean ± SD.

Carnosine Placebo p value

Performance (s) 30.0 ± .8 29.8 ± .8 .302

RPE (a. u. 0 - 10) 6.5 ± 1.2 7.0 ± 1.3 .195

Pain (a. u. 0 - 10) 4.0 ± 2.6 3.4 ± 2.1 .173

Session RPE (a. u.)107 ± 23 106 ± 28 .703

Blood glucose (mg/dl)109 ± 23 104 ± 12 .969

Insulin (μU/ml) 23.8 ± 13.0 19.5 ± 9.0 .329

Bicarbonate (mEq/l)16 ± 2 16 ± 2 1.0

Open Access 171

P. L. INVERNIZZI ET AL.

Figure 1.

The acute effects of carnosine administration, compared to a placebo

treatment, on the six sprint bouts of the RAST test; *Significant (p < .05)

main effects of time.

Figure 2.

Time course of blood lactate concentration in the two conditions (car-

nosine and placebo) following the running-based anaerobic sprint test

(RAST).

The RAST caused a very small and similar increase in

DOMS in the two conditions (condition × time, p = .755) with a

significant decrease towards zero (main factor time, p < .05) at

48 hours post test (carnosine 1.2 ± 1.3 vs .6 ± 1.0 at 24 and 48

hours respectively and placebo 1.0 ± 1.0 vs .4 ± .7 at 24 and 48

hours respectively).

Discussion

The ingestion of 1 g of carnosine and 1 g of β-alanine 4

hours prior to an all-out shuttle running test did not have any

effect on performance, perceived exertion, muscle soreness and

session RPE. Similarly, the response of insulin, glucose, bicar-

bonates and blood lactate concentration to exercise were not

affected by carnosine supplementation compared to placebo.

During the RAST the time to complete each bout of 35 m in-

creased similarly in the two conditions, with no difference in

total time. Contrary to the study by Suzuki et al. (Suzuki et al.,

2002) we failed to show a difference between the two condi-

tions towards the latter fractions of an intermittent all-out exer-

cise. The different type of exercises (that is cycling versus run-

ning) and the modality (that is continuous versus intermittent)

might represent some of the causes for the discrepancy between

the two studies. Moreover, we cannot exclude that the acute

ingestion of carnosine failed to increase the muscle content of

carnosine (see below in the Discussion). The occurrence of

fatigue is the most plausible explanation for the decrease of the

running speed. In particular, a decrease in muscle pH represents

one of the main factors responsible for the decline in muscle

contractility (Allen et al., 2008). The high level of blood lactate

observed at the end of the RAST might be an indirect marker of

increased acidosis in the muscle. Therefore, it can be hypothe-

sized that the dose of carnosine utilized in this study was not

sufficient to increase the muscle content of carnosine to a level

that was sufficient to exert a buffering effect on muscle cells

acidosis. Alternatively, it can be hypothesized that the RAST

did not determine such a decrease in muscle pH to require a

higher concentration of muscle carnosine content than normally

present. However, the blood lactate concentration and the level

of bicarbonates after the all-out shuttle running indirectly sug-

gest that the metabolic stress (and the hydrogen ions concentra-

tions) was high.

Noteworthy, in the present study we failed to detect any dif-

ference in blood bicarbonates concentration between conditions.

On the contrary, some authors (in spite of the administration of

a lower dose of carnosine) showed such a difference and hy-

pothesized an increased action exerted by the nonbicarbonat

buffering due to carnosine and anserine administration (Suzuki

et al., 2006).

The lack of difference in performance between the two con-

ditions neither support the increase in Ca2+ sensitivity of the

contractile fibres that represents an alternative mechanism

through which muscle carnosine can improve muscle perform-

ance (Dutka & Lamb, 2004). In theory, this ergogenic effect of

muscle carnosine should have been useful if the maximal effort

performed was not sufficiently high to elicit low levels of mus-

cle pH.

A further aspect that might have contributed to the lack of a

positive effect of acute carnosine ingestion on performance is

the duration of the exercise. According to a recent meta analy-

sis (Hobson et al., 2012) a higher content of muscle carnosine is

beneficial mostly for high intensity exercises lasting between

60 and 240 seconds. In the present study the average overall

duration of the RAST was approximately 35 seconds.

Blood glucose and insulin showed an increase after the

RAST compared to baseline that was similar in the two groups.

It is likely that the increase in blood glucose was mediated by

an increase in cathecolamines. However, as for performance, it

can be hypothesized that the amount of carnosine administered

was not sufficient to be of any effect on the adrenergic system.

Although in the present study a much higher dose of car-

nosine than in an earlier study (Kraemer et al., 1995) was used,

no enhancing effect on short high intensity running perform-

ance was found.

It cannot be excluded that the acute administration of car-

nosine is a vain strategy for the improvement of exercise. How-

ever, future studies should aim to evaluate the effects of inges-

tion of carnosine at higher dosages (e.g. twofold the dose used

in the present study) and to assess its efficacy using much

higher stressing exercise (e.g. all-out intermittent or continuous

anaerobic exercises performed for a longer time period). Fur-

thermore, it would be interesting to test the effect of acute in-

gestion of carnosine in individuals (like vegetarians) who might

be characterized by a lower level of muscle carnosine.

A possible limitation of this study could be the lack of

evaluation of the dietary habits of subjects. In fact the content

of muscle carnosine in a subject that follows a predominantly

vegetarian diet could be drastically lower than a person taking

meat in high amount.

Open Access

172

P. L. INVERNIZZI ET AL.

Conclusion

In conclusion, the acute ingestion of 1 g of L-carnosine and 1

g of β-alanine few hours prior to an intermittent all-out running

exercise does not have any effect on overall performance and

on the latter bouts of the exercise. Similarly, the responses of

blood insulin, glucose, bicarbonate and lactate concentrations to

the RAST were not affected by the acute ingestion of carnosine

compared to placebo.

Based on the present results, the acute oral administration of

carnosine prior to an anaerobic running exercise lasting around

30 seconds does not seem to be an effective strategy to improve

performance.

REFERENCES

Aldini, G., Orioli, M., Rossoni, G., Savi, F., Braidotti, P., Vistoli, G.,

Yeum, K. J., Negrisoli, G., & Carini, M. (2001). The carbonyl scav-

enger carnosine ameliorates dyslipidaemia and renal function in

Zucker obese rats. Journal of Cellular and Molecular Medicine, 15,

1339-1354. http://dx.doi.org/10.1111/j.1582-4934.2010.01101.x

Alhamdani, M. S., Al-Azzawie, H. F., & Abbas, F. K. (2007). De-

creased formation of advanced glycation end-products in peritoneal

fluid by carnosine and related peptides. Peritoneal Dialysis Interna-

tional, 27, 86-89.

Allen, D. G., Lamb, G. D., & Westerblad, H. (2008). Skeletal muscle

fatigue: Cellular mechanisms. Physiological Reviews, 88, 287-332.

http://dx.doi.org/10.1152/physrev.00015.2007

Baguet, A., Bourgois, J., Vanhee, L., Achten, E., & Derave, W. (2010).

Important role of muscle carnosine in rowing performance. Journal

of Applied Physiology, 109, 1096-1101.

http://dx.doi.org/10.1152/japplphysiol.00141.2010

Begum, G., Cunliffe, A., & Leveritt, M. (2005). Physiological role of

carnosine in contracting muscle. International Journal of Sport Nu-

trition and Exercise Metabolism, 15, 493-514.

Borg, G. (1998). Borg’s perceived exertion and pain scales. Cham-

paign, IL: Human Kinetics.

Buse, M. G., Weigand, D. A., Peeler, D., & Hedden, M. P. (1980). The

effect of diabetes and the redox potential on amino acid content and

release by isolated rat hemidiaphragms. Metabolism, 29, 605-616.

http://dx.doi.org/10.1016/0026-0495(80)90104-3

Derave, W., Everaert, I., Beeckman, S., & Baguet, A. (2010). Muscle

carnosine metabolism and beta-alanine supplementation in relation to

exercise and training. Sports Medicine , 40, 247-263.

http://dx.doi.org/10.2165/11530310-000000000-00000

Derave, W., Ozdemir, M. S., Harris, R. C., Pottier, A., Reyngoudt, H.,

Koppo, K., Wise, J. A., & Achten, E. (2007). Beta-alanine supple-

mentation augments muscle carnosine content and attenuates fatigue

during repeated isokinetic contraction bouts in trained sprinters.

Journal of Applied P hys i o l og y , 103, 1736-1743.

http://dx.doi.org/10.1152/japplphysiol.00397.2007

Di Pierro, F., Bertuccioli, A., Bressan, A., & Rapacioli, G. (2011).

Carnosine-based supplement. Nutrafoods, 1, 43-47.

http://dx.doi.org/10.1007/BF03223387

Dutka, T. L., & Lamb, G. D. (2004). Effect of carnosine on excitation-

contraction coupling in mechanically-skinned rat skeletal muscle.

Journal of Muscle Research and Cell Motility, 25, 203-213.

http://dx.doi.org/10.1023/B:JURE.0000038265.37022.c5

Foster, C., Florhaug, J. A., Franklin, J., Gottschall, L., Hrovatin, L. A.,

Parker, S., Doleshal, P., & Dodge, C. (2001). A new approach to

monitoring exercise training. The Journal of Strength & Condition-

ing Research, 15, 109-115.

Gardner, M. L., Illingworth, K. M., Kelleher, J., & Wood, D. (1991).

Intestinal absorption of the intact peptide carnosine in man, and

comparison with intestinal permeability to lactulose. The Journal of

Physiology, 439, 411-422.

Harris, R. C., Wise, J. A., Price, K. A., Kim, H. J., Kim, C. K., & Sale,

C. (2012). Determinants of muscle carnosine content. Amino Acids,

43, 5-12. http://dx.doi.org/10.1007/s00726-012-1233-y

Hill, C. A., Harris, R. C., Kim, H. J., Harris, B. D., Sale, C., Boobis, L.

H., Kim, C. K., & Wise, J. A. (2007). Influence of beta-alanine sup-

plementation on skeletal muscle carnosine concentrations and high

intensity cycling capacity. Amino Acids, 3 2, 225-233.

http://dx.doi.org/10.1007/s00726-006-0364-4

Hipkiss, A. R. (2009). Carnosine and its possible roles in nutrition and

health. Advances in Food and Nutrition Research, 57, 87-154.

http://dx.doi.org/10.1016/S1043-4526(09)57003-9

Hobson, R. M., Saunders, B., Ball, G., Harris, R. C., & Sale, C. (2012).

Effects of beta-alanine supplementation on exercise performance: A

meta-analysis. Amino Acids, 43, 25-37.

http://dx.doi.org/10.1007/s00726-011-1200-z

Impellizzeri, F. M., Rampinini, E., Coutts, A. J., Sassi, A., & Marcora,

S. M. (2004). Use of RPE-based training load in soccer. Medicine &

Science in Sports & Exercise, 36, 1042-1047.

http://dx.doi.org/10.1249/01.MSS.0000128199.23901.2F

Jagim, A. R., Wright, G. A., Brice, A. G., & Doberstein, S. T. (2013).

Effects of beta-alanine supplementation on sprint endurance. The

Journal of Strength & Conditioning Researc h , 27, 526-532.

http://dx.doi.org/10.1519/JSC.0b013e318256bedc

Kraemer, W. J., Gordon, S. E., Lynch, J. M., Pop, M. E., & Clark, K. L.

(1995). Effects of multibuffer supplementation on acid-base balance

and 2,3-diphosphoglycerate following repetitive anaerobic exercise.

Journal of the International Society of Sports Nutrition, 5, 300-314.

Margolis, F. L., Grillo, M., Kawano, T., & Farbman, A. I. (1985). Car-

nosine synthesis in olfactory tissue during ontogeny: Effect of ex-

ogenous beta-alanine. Journal of Neurochemistry, 44, 1459-1464.

http://dx.doi.org/10.1111/j.1471-4159.1985.tb08783.x

Nagai, K., Niijima, A., Yamano, T., Otani, H., Okumra, N., Tsuruoka,

N., Nakai, M., & Kiso, Y. (2003). Possible role of L-carnosine in the

regulation of blood glucose through controlling autonomic nerves.

Experimental Biolo g y and Medicine, 228, 1138-1145.

Nagai, K., Tanida, M., Niijima, A., Tsuruoka, N., Kiso, Y., Horii, Y.,

Shen, J., & Okumura, N. (2012). Role of L-carnosine in the control

of blood glucose, blood pressure, thermogenesis, and lipolysis by

autonomic nerves in rats: Involvement of the circadian clock and

histamine. Am ino Acids, 43, 97-109.

http://dx.doi.org/10.1007/s00726-012-1251-9

Shao, L., Li, Q. H., & Tan, Z. (2004). L-carnosine reduces telomere

damage and shortening rate in cultured normal fibroblasts. Bioche-

mical and Biophysical Research Communications, 324, 931-936.

http://dx.doi.org/10.1016/j.bbrc.2004.09.136

Smith, E. C. (1938). The buffering of muscle in rigor; protein, phos-

phate and carnosine. The Journal of Physiology, 9 2, 336-343.

Suzuki, Y., Ito, O., Mukai, N., Takahashi, H., & Taka-Matsu, K. (2002).

High level of skeletal muscle carnosine contributes to the latter half

of exercise performance during 30-s maximal cycle ergometer sprint-

ing. Japanese Journal of Physiology, 52, 199-205.

http://dx.doi.org/10.2170/jjphysiol.52.199

Suzuki, Y., Nakao, T., Maemura, H., Sato, M., Kama-Hara, K., Mori-

matsu, F., & Takamatsu, K. (2006). Carnosine and anserine ingestion

enhances contribution of nonbicarbonate buffering. Medicine & Sci-

ence in Sports & Exercise, 38, 334-338.

Yamano, T., Niijima, A., Iimori, S., Tsuruoka, N., Kiso, Y., & Nagai, K.

(2001). Effect of L-carnosine on the hyperglycemia caused by in-

tracranial injection of 2-deoxy-D-glucose in rats. Neuroscience Let-

ters, 313, 78-82. http://dx.doi.org/10.1016/S0304-3940(01)02231-5

Zagatto, A. M., Beck, W. R., & Gobatto, C. A. (2009). Validity of the

running anaerobic sprint test for assessing anaerobic power and pre-

dicting short-distance performances. The Journal of Strength & Con-

ditioning Research, 23, 1820-1827.

http://dx.doi.org/10.1519/JSC.0b013e3181b3df32

Open Access 173

P. L. INVERNIZZI ET AL.

Open Access

174

Abbreviations

BMI: Body Mass Index;

RAST: running-based anaerobic sprint test;

RPE: perceptual response to exercise;

DOMS: delayed onset of muscle soreness;

EDTA: Ethylenediaminetetraacetic acid;

ELISA: Enzyme-Linked Immuno Sorbent Assay