Carnosine and L-carnosine are two naturally occurring compounds found in various animal tissues, including the human body. Carnosine is a dipeptide consisting of two amino acids, beta-alanine and histidine, while L-carnosine is a form of carnosine that is more bioavailable to the body. Both carnosine and L-carnosine have been extensively studied for their potential health benefits, including their ability to improve athletic performance and prevent age-related decline in cognitive function. 

This article will explore what carnosine and L-carnosine are, how they arise, the normal amount in humans and horses’ bodies, food sources, and their effects on the body, specifically in sports performance.

1. CARNOSINE IN GENERAL

Carnosine (b-alanyl-L-histidine) is a cytoplasmic dipeptide synthesised from b-alanine and histidine (Harris et al. 1974). Carnosine was first isolated by Russian chemist Vladimir Gulevich in 1900, and its presence in skeletal muscle was first demonstrated by British physiologist A.V. Hill in 1928 (Boldyrev et al., 2013). Since then, a significant amount of research has been conducted on this intriguing molecule, revealing numerous potential health benefits..  Carnosine can also be found in relatively high concentrations in the central nervous system. (Harris et al. 1974).  It is a naturally occurring compound found in high concentrations in the skeletal muscles and the brain. L-carnosine, on the other hand, is found in the lens of the eye, kidney, and liver. In skeletal muscle and the brain, it functions as a buffer against the buildup of hydrogen ions during exercise. This buffer system helps to prevent muscle fatigue and can improve exercise performance.

L-carnosine is the biologically active form of carnosine and is found in many dietary supplements. It is often used as a natural anti-aging supplement due to its antioxidant properties. L-carnosine is also believed to have neuroprotective effects, and may improve cognitive function and memory. Most visible and proven effects of L-carnosine´s are experienced on  muscles, tendons and ligaments however Chemipower is pioneer in researching and developing a sports performance and recovery gel especially focussing on it. 

1.1 How do Carnosine and L-Carnosine Arise?

Carnosine is synthesized in the body through a process called carnosine synthesis. This process involves the condensation of β-alanine and histidine, which are both derived from dietary sources. β-alanine is a non-essential amino acid that can be synthesized in the liver, while histidine is an essential amino acid that must be obtained from the diet.

L-carnosine is formed in the body through the combination of β-alanine and l-histidine. L-histidine is an essential amino acid that must be obtained from the diet, while β-alanine can be synthesized in the liver.

1.2 Normal Amount of Carnosine and L-Carnosine in Humans and Horses’ Bodies

The normal amount of carnosine in the human body varies depending on the muscle group being studied. In the skeletal muscles, carnosine concentrations range from 5 to 30 mM, with the highest concentrations found in the fast-twitch muscle fibers. Whether there is a ceiling to the amount of carnosine that can be stored in skeletal muscle is currently unknown (Baguet et al.2009). In the brain, carnosine concentrations are much lower, ranging from 0.5 to 2 mM. There are numerous determinants of the M-Carn concentration, including species, gender, age, muscle fiber type, diet, supplementation, exercise and training (Harris et al. 2012). 

The normal amount of l-carnosine in the human body is not well established and can vary alot, as its distribution is more limited than that of carnosine. In the lens of the eye, l-carnosine concentrations can range from 2 to 10 mM, while in the liver and kidney, concentrations are much lower.

In horses, carnosine concentrations in the skeletal muscles are similar to those found in humans, with concentrations ranging from 5 to 25 mM. However, there is limited data on the concentrations of l-carnosine in horses.

1.3 Dietary Carnosine

The predominant source of dietary carnosine in humans is via meat and fish consumption (Abe 2000),  In particular, beef, pork, chicken, and tuna are good sources of carnosine, while chicken and turkey are good sources of l-carnosine, although it should be noted that cooking practices significantly influence the amount available. Given the diversity of the human diet, the potential range of dietary carnosine intakes are relatively broad and might range from 50 to 500 mg d-1 in the omnivorous diet (Baguet et al. 2009). In high meat and fish consumers, such as those in the US, South America and Asia, dietary intake is likely to be higher than this. Conversely, vegetarians have been shown to have significantly lower M-Carn levels than their meat-eating counterparts (Everaert et al. 2011).Vegetarian sources of carnosine and l-carnosine are limited, with some evidence suggesting that mushrooms may contain low levels of carnosine. For the majority of humans, diet is likely to be the good means of attaining and maintaining higher M-Carn levels. 

1.4 Carnosine in blood

Carnosine can be ingested through the diet, but the presence of carnosinase in the enterocytes of humans, especially in the jejunal mucosa (Sadikali et al. 1975), suggests that some carnosine is cleaved into b-alanine and L-histidine before reaching the blood stream. However, since the jejunal activity is low (Sadikali et al. 1975), it is very likely that part of the ingested carnosine reaches the blood stream, where it is rapidly hydrolysed in plasma due to the high activity of the enzyme carnosinase meaning that only negligible levels of carnosine are detectable in the blood (Park et al. 2005).

Methods for detecting carnosine in blood:

1. High-Performance Liquid Chromatography (HPLC): This is a commonly used technique to separate and quantify carnosine from blood samples. The technique involves injecting the sample into a column filled with a stationary phase and a mobile phase. The carnosine will interact differently with the stationary phase and the mobile phase, allowing it to be separated and detected.

2. Mass Spectrometry: This is another technique that can be used to detect carnosine in blood. The technique involves ionizing the carnosine molecules and then separating them based on their mass-to-charge ratio. This can provide a more specific and sensitive measurement of carnosine in blood.

3. Enzyme-linked immunosorbent assay (ELISA): This is a technique that uses antibodies to detect the presence of carnosine in blood. A blood sample is mixed with an antibody that is specific for carnosine, and then the amount of bound antibody is measured. This technique is relatively simple and can provide rapid results.

Overall, the most commonly used technique for detecting carnosine in blood is HPLC. However, mass spectrometry and ELISA can also be used depending on the specific needs of the experiment or clinical test.

2. CARNOSINE EFFECTS 

2.1 Carnosine effects on different body parts

Scientists have found it to have various effects on different parts of the body.

1. Muscles: Carnosine is found in high concentrations in skeletal muscle tissue, where it helps to buffer the build-up of lactic acid during exercise. This can help to delay fatigue and improve endurance performance. L-carnosine has also been found to have similar effects on muscle fatigue and recovery. This is the area where Chemipower has been focussing its research and product development.

2. Brain: Carnosine and L-carnosine have been found to have neuroprotective effects, helping to prevent damage to brain cells from oxidative stress and inflammation. Studies have also suggested that L-carnosine may have cognitive benefits, improving memory and attention in older adults.

3. Eyes: L-carnosine has been found to have antioxidant properties that can help to protect the eyes from age-related damage, such as cataracts and macular degeneration.

4. Skin: Carnosine and L-carnosine have been found to have anti-aging properties, helping to protect the skin from damage caused by UV radiation and other environmental factors.

5. Immune system: Carnosine and L-carnosine have been found to have immunomodulatory effects, helping to regulate the activity of immune cells and improve immune function.

6. Cardiovascular system: Studies have suggested that carnosine and L-carnosine may have beneficial effects on the cardiovascular system, helping to reduce inflammation, improve blood flow, and protect against heart disease.

Overall, carnosine and L-carnosine appear to have a wide range of potential health benefits, and more research is needed to fully understand their effects on different parts of the body.

3. CARNOSINE INCREASES MUSCLE PERFORMANCE

A potential benefit of carnosine is its ability to improve exercise performance. There is evidence to suggest that carnosine may have a role in promoting muscle growth and improving athletic performance. Studies have found that carnosine can reduce muscle fatigue and improve muscle endurance, potentially making it useful for athletes or individuals looking to improve their exercise performance (Trombold et al., 2011). Carnosine is naturally found in high concentrations in skeletal muscle tissue, where it acts as a buffer to reduce the buildup of lactic acid during exercise. This can delay the onset of muscle fatigue and improve muscle endurance, allowing individuals to exercise for longer periods of time. However, given the speed of current publications relating to the effects of b-alanine on exercise performance, there remains a need to continually update the summaries of this topic area. 

To date, the performance improvements shown following b-alanine supplementation have largely been ascribed to increases in intracellular buffering as a result of the increased M-Carn concentrations, although there are other potential mechanisms that might also explain a performance effect. Future research is needed to better understand the mechanisms underlying all of carnosine’s effects but regarding muscle performance Chemipower has developed together with Tartu University scientists, professional athletes and existential sports and equestrian community two l-carnosine based sports gels – Carnosport for athletes and Carnogel for sport horse health benefits.

In the meta-analysis of Hobson et al. (2012), results indicated significant improvements in exercise capacity tests, but not exercise performance tests (P = 0.204), although it was noted that there were fewer studies using performance measures. However, there have been a significant numberof studies conducted since this meta-analysis that have measured the effect of b-alanine supplementation on both performance and capacity.

(Sale et al. 2012) investigated the effect of b-alanine supplementation on a cycling capacity test with recreationally active participants and in this case showed baseline data within 3–4 s of the expected endurance time of 78 s. Endurance time was increased by 9.7 s (13.2 %) following 4 weeks of supplementation at 6.4 g d-1likely due to an increase in muscle buffering capacity based upon the expected increase in muscle PH formation and increase in MCarn. Hobson et al. (2012) in their metaanalysis concluded that exercise lasting between 60 and 240 s was likely to be positively influenced by an increase in muscle buffering capacity.

Several recent studies have investigated the effect of b-alanine supplementation on 2,000 m rowing ergometry in well- trained male rowers, where the exercise duration is typically between 6 and 7 min and with an estimated 12 % of energy sources from non-oxidative glycolysis (Stellingwerff et al. 2011). Baguet et al. (2010) recruited 18 elite Belgian rowers who performed a 2,000-m rowing test pre and post 7 weeks of supplementation with either 5 g d-1 b-alanine or placebo.Results showed a 2.7 ± 4.8 s improvement in performance with b-alanine, which only just failed to reach statistical significance (P = 0.07), although the individual increases in rowing speed were positively correlated to the increases in M-Carn. Ducker et al. (2013) showed an improvement of 2.9 ± 4.1 s in well-trained rowers (N = 7) supplemented with 80 mg kg-1 BM d-1, compared with placebo-supplemented participants (N = 9) who were 1.2 ± 2.9 s slower, and again this just failed to reach significance (P = 0.06), although P = 0.034 when just the two performance times were compared using an unpaired t test. Unpublished observations from our research group (Hobson et al.) used magnitude-based inferences to determine small effects of b-alanine supplementation of practical importance in the applied setting. A 6.4 ± 8.1 s improvement in rowing performance with b-alanine over placebo was very likely to have been due to b-alanine.

Since increased M-Carn undoubtedly results in increased muscle buffering capacity, there has been much interest in determining the effects of b-alanine supplementation on exercise performance and capacity limited by the accumulation of hydrogen cations. Despite this, not all research has shown a beneficial effect of b-alanine supplementation, although this may be due to the variety and durations of the exercise tests involved (Hobson et al. 2012). 

3.1 High-intensity intermittent and repeated sprint exercise

Several studies have investigated the effect of b-alanine supplementation on intermittent repeated bouts of exercise, due to increasing interest in repeated sprint activities, such as team sports (Hoffman et al. 2008a; Sweeney et al. 2010), Saunders et al. (2012a) employed the YoYo Intermittent Recovery Test Level 2 (YoYo IR2) which determines an individual’s ability to repeatedly perform and recover from high- intensity exercise. Following a 12-week supplementation period of 3.2 g d-1 b-alanine or placebo, male amateur footballers supplemented with b-alanine showed a 34.4 % improvement in performance contrasting with the 7.3 % reduction in performance in the placebo group. Likewise, unpublished observations from our group have demonstrated that 4 weeks of b-alanine supplementation (6.4 g d-1) was capable of inducing a significant 7 % improvement in high-intensity intermittent performance (four bouts of 30 s Wingate test for the upper body with 3 min recovery between bouts) in highly trained combat athletes.

Mannion et al. (1995) suggested that M-Carn contributed 7 % to total buffering capacity, although this is likely a minimum estimate based upon muscle with a metabolic composition. The relative contribution from M-Carn is usually based upon a comparison of its buffering effect, derived from its pKa, against calculations of total muscle buffering capacity. Muscle buffering capacity is usually determined by the titration of skeletal muscle homogenates (Harris et al. 1990; Mannion et al. 1994), although this technique remains fundamentally flawed with the result that the contribution of M-Carn is significantly underestimated.

4. THERAPEUTIC EFFECTS OF CARNOSINE 

In addition to the potential role in improving exercise performance, more recently attention has spread to the potential therapeutic benefits of carnosine. Despite studies dating back over two decades (Boldyrev 1992), there remains a paucity of clinical data supporting the use of carnosine in medicine, evidencing a large gap between the experimental and clinical findings. 

Carnosine has been considered a natural scavenger/suppressor of reactive oxygen species, advanced glycation end products and reactive aldehydes (Boldyrev 1993; Hipkiss et al. 1995; Babizhayev et al. 1994). Such properties may confer therapeutic effects, particularly in those conditions characterized by exacerbated oxidative stress, including neurodegenerative diseases, cancer, diabetes and senescence.

4.1 Carnosine as antioxidant 

One of the main benefits of carnosine is its strong antioxidant properties. Free radicals are unstable molecules that can cause damage to cells and tissues, leading to oxidative stress and inflammation. Carnosine has been shown to have the ability to scavenge free radicals and protect cells from oxidative damage, which can reduce the risk of chronic diseases such as cancer, heart disease, and Alzheimer’s disease.

4.2 Parkinson’s Disease

Boldyrev et al. (2008) tested the efficacy of carnosine supplementation in Parkinson’s disease. In an open-label trial, patients on dihydroxyphenylalanine (DOPA)-containing drugs were given carnosine supplementation (1.5 g d-1) or no additive treatment for 30 days. Patients receiving carnosine in addition to their regular pharmacological treatment showed a clinical improvement of 36 % (vs. 16 % in controls), as assessed by the Unified Parkin son’s Disease Rating Scale. The carnosine-treated patients also experienced improvements in physical symptoms (e.g. rigidity of extremities and upper-limb movements) by up to 38 %. The decrease in the neurological symptoms correlated with the decrease in blood serum carbonyl levels, the increase in resistance to oxidation of blood lipoproteins and the increase in superoxide dismutase activity of red blood cells. Based on their findings, the authors concluded that the ‘‘combination of carnosine with basic therapy of Parkinson’s patients (…) might be a reasonable way to improve the results of Parkinson’s treatment (Boldyrev et al. 2008). However, these results must be carefully interpreted in light of the absence of a placebo-controlled double-blind design.

4.3 Diabetes

Carnosine, which has been recognized as a potential glycation inhibitor, could be also useful in the treatment of diabetes. According to Hipkiss (2006), this hypothesis relies on the following observations: 

(i) diabetic rats have lower plasma carnosine concentrations than healthy rats; 

(ii) human diabetics have lower erythrocyte carnosine levels than their healthy peers; 

(iii) carnosine protects against acidic haemolysis in diabetic rat erythrocytes; and 

(iv) carnosine exerts regulatory effects on blood glucose levels in rats.

Furthermore, it has been suggested that carnosine is implicated in the alleviation of some diabetic complications.

4.4 Aging

Carnosine has also been found to have anti-aging effects. As we age, our cells and tissues become more vulnerable to oxidative stress and inflammation, which can lead to the development of age-related diseases. Some studies suggest that carnosine may help to reduce the effects of aging on the body by protecting cells and tissues from damage caused by oxidative stress. Additionally, carnosine has been shown to have anti-glycation properties, which can reduce the formation of advanced glycation end products (AGEs) that contribute to the aging process.

Recent evidence has suggested that a carnosine-rich diet may have therapeutic benefits for elderly individuals. In this respect, it has been demonstrated that intramuscular carnosine content may be reduced in elderly subjects (Tallon et al. 2007), although conflicting results exist (Kim 2009). 

In a randomized, double-blind, placebo-controlled study, Stout et al. (2008) showed that b-alanine supplementation elicited a 28.6 % increase in physical working capacity at the neuromuscular fatigue threshold after 90 days of b-alanine supplementation (3 9 800

mg d-1) in men and women aged 55–92 years. However, the authors did not assess M-Carn content. Recently, we showed that that b-alanine supplementation increased M-Carn content by *85 % in healthy older individuals (60–80 years) with improvements in physical exercise capacity (del Favero et al. 2012). Benefits in maintaining a high M-Carn content may be both immediate and long-term, since this may encourage individuals to maintain a more active lifestyle.

It has been shown that carnosine can increase the lifespan of mice (Yuneva et al. 1999) and fruit flies (Yuneva et al. 2002) and protects rats and Mongolian gerbils against the implications of brain ischaemia (Dobrota et al. 2005). In humans, however, data are still scarce to support the protective role of carnosine in the prevention of neurodegeneration.

4.5 Anti-inflammatory effects

In addition to its antioxidant and anti-aging effects, carnosine has been found to have anti-inflammatory effects. Chronic inflammation is a contributing factor to many diseases, including arthritis, diabetes, and cardiovascular disease. Studies have shown that carnosine can reduce inflammation throughout the body by inhibiting the production of inflammatory molecules such as interleukin-6 (IL-6) and tumor necrosis factor-alpha (TNF-alpha).

4.6 Neuroprotective effects

Carnosine has been shown to have neuroprotective effects. Studies have shown that carnosine can protect against neurodegenerative diseases such as Alzheimer’s disease by reducing oxidative stress and inflammation in the brain. Additionally, carnosine has been found to improve cognitive function and memory in older adults.

CONCLUSION

In conclusion, carnosine is a dipeptide molecule that is naturally found in high concentrations in various tissues throughout the body, including the brain, muscles, and heart. While the research on carnosine is still ongoing, there is evidence to suggest that it may have several potential health benefits.

One of the main benefits of carnosine is its strong antioxidant properties. By protecting cells from oxidative damage caused by free radicals, carnosine can reduce the risk of chronic diseases such as cancer, heart disease, and Alzheimer’s disease. Additionally, carnosine has anti-aging effects and can reduce inflammation throughout the body.

Carnosine effects have been directly derived to improve exercise performance by reducing muscle fatigue and increasing muscle endurance. This can be particularly beneficial for athletes or individuals who engage in regular physical activity. The same can be stated for sports horses. Carnosine effects on muscles, tendons and ligaments is the central study area where Chemipower is focusing its research and product development. 

Chemipower has developed together with Tartu University scientists, professional athletes and existended sports and equestrian community two l-carnosine based sports gels – Carnosport for athletes and Carnogel for sport horse health benefits.

SOURCES:

Parkhouse WS, McKenzie DC, Hochachka PW, Ovalle WK (1985) Buffering capacity of deproteinized human vastus lateralis muscle. J Appl Physiol 58:14–17

Harris RC, Hultman E, Nordesjo LO (1974) Glycogen, glycolytic intermediates and high energy phos-phates determined in biopsy samples of musculus quadri-ceps femoris in man at rest. Methods and variance of values. Scad J Clin Lab Invest 33:109–120

Baguet A, Reyngoudt H, Pottier A, Everaert I, Callens S, Achten E, Derave W (2009) Carnosine loading and washout in human skeletal muscles. J Appl Physiol 106:837–842

Harris RC, Wise JA, Price KA, Kim HJ, Kim CK, Sale C (2012)Determinants of muscle carnosine content. Amino Acids 43:5–12

Everaert I, Mooyaart A, Baguet A, Zutinic A, Baelde H, Achten E, Taes Y, De Heer E, Derave W (2011) Vegetarianism, female gender and increasing age, but not CNDP1 genotype, are associated with reduced muscle carnosine levels in humans. Amino Acids 40:1221–1229

Sadikali F, Darwish R, Watson WC (1975) Carnosinase activity of human gastrointestinal mucosa. Gut 16:585–589

Park YJ, Volpe SL, Decker EA (2005) Quantitation of carnosine in human plasma after dietary consumption of beef. J Agric Food Chem 53:4736–4739

Hobson RM, Saunders B, Ball G, Harris RC, Sale C (2012) Effects of b-alanine supplementation on exercise performance: a review by meta-analysis. Amino Acids 43:25–37

Sale C, Hill CA, Ponte J, Harris RC (2012) Beta-alanine supplemen- tation improves isometric endurance of the knee extensor muscles. J Int Soc Sports Nutr 9:26

Stellingwerff T, Maughan RJ, Burke LM (2011) Nutrition for power sports: middle distance running, track cycling, rowing, canoeing/ kayaking, and swimming. J Sport Sci 29:S79–S89

Baguet A, Bourgois J, Vanhee L, Achten E, Derave W (2010) Important role of carnosine in rowing performance. J Appl Physiol 109:1096–1101

Hoffman JR, Ratamess NA, Faigenbaum AD, Ross R, Kang J, Stout JR, Wise JA (2008a) Short-duration b-alanine supplementation increases training volume and reduces subjective feelings of fatigue in college football players. Nut Res 28:31–35

Sweeney KM, Wright GA, Brice AG, Doberstein ST (2010) The effects of b-alanine supplementation on power performance during repeated sprint activity. J Strength Cond Res 24:79–87

Saunders B, Sunderland C, Harris RC, Sale C (2012a) b-Alanine supplementation improves YoYo intermittent recovery test performance. J Int Soc Sports Nutr 9:39

Mannion AF, Jakeman PM, Willan PL (1995) Skeletal muscle buffer value, fibre type distribution and high intensity exercise perfor- mance in man. Exp Physiol 80:89–101

Mannion AF, Jakeman PM, Willan PLT (1994) Effects of isokinetic training of the knee extensors on high-intensity exercise performance and skeletal muscle buffering. Eur J Appl Physiol 68:356–361

Harris RC, Marlin DJ, Dunnett M, Snow DH, Hultman E (1990)Muscle buffering capacity and dipeptide content in the thor- oughbred horse, greyhound dog and man. Comp Biochem Phys A 97:249–251

Boldyrev AA (1992) Carnosine—biological role and potential applications in medicine. Biochemistry (Moscow) 57:898–904

Boldyrev AA (1993) Does carnosine possess direct antioxidant activity. Int J Biochem 25:1101–1107

Babizhayev MA, Seguin MC, Gueyne J, Evstigneeva RP, Ageyeva EA, Zheltukhina GA (1994) L-Carnosine (beta-alanyl-L-histi- dine) and carcinine (beta-alanylhistamine) act as natural antiox- idants with hydroxyl-radical-scavenging and lipid-peroxidase activities. Biochem J 304:509–516

Hipkiss AR, Michaelis J, Syrris P (1995) Non-enzymatic glycosyl- ation of the dipeptide L-carnosine, a potential anti-protein-cross- linking agent. FEBS Lett 371:81–85

Yuneva MO, Bulygina ER, Gallant SC, Kramarenko GG, Stvolinsky SL, Semyonova ML, Boldyrev AA (1999) Effect of carnosine on age-induced changes in senescence-accelerated mice. J Anti- Aging Med 2:337–342

Yuneva AO, Kramarenko GG, Vetreshchak TV, Gallant S, Boldyrev AA (2002) Effect of carnosine on Drosophila melanogaster life span. Bull Exp Biol Med 133:559–661

Dobrota D, Fedorova T, Stvolinsky S, Babusikova E, Likavcanova K, Drgova A, Strapkova A, Boldyrev A (2005) Carnosine protects the brain of rats and Mongolian gerbils against ischemic injury: after-stroke-effect. Neurochem Res 30:1283–1288

Boldyrev A, Fedorova T, Stepanova M, Dobrotvorskaya I, Kozlova E, Boldanova N, Bagyeva G, Ivanova-Smolenskaya I, Illarioshkin S (2008) Carnosine increases efficiency of DOPA therapy of Parkinson’s disease. Rejuv Res 11:988–994

Hipkiss AR (2006) Would carnosine or a carnivorous diet help suppress aging and associated pathologies? Ann NY Acad Sci 1067:369–374

Tallon MJ, Harris RC, Maffulli N, Tarnopolsky MA (2007) Carno- sine, taurine and enzyme activities of human skeletal muscle fibres from elderly subjects with osteoarthritis and young moderately active subjects. Biogerontology 8:129–137

Kim HJ (2009) Comparison of the carnosine and taurine contents of vastus lateralis of elderly Korean males, with impaired glucose tolerance, and young elite Korean swimmers. Amino Acids 36:359–363

Stout JR, Graves BS, Smith AE, Hartman MJ, Cramer JT, Beck TW, Harris RC (2008) The effect of beta-alanine supplementation on neuromuscular fatigue in elderly (55–92 years): a double-blind randomized study. J Int Soc Sports Nutr 5:21

del Favero S, Roschel H, Solis MY, Hayashi AP, Artioli GG, Otaduy MC, Benatti FB, Harris RC, Wise JA, Leite CC, Pereira RM, de Sa ́-Pinto AL, Ohsawa M, Mutoh J, Asato M, Yamamoto S, Ono H, Hisa H, Kamei J (2012) Carnosine has antinociceptive properties in the inflammation-induced nociceptive response in mice. Eur J Pharmacol 682:56–61

Derave W, Sale C (2012) Carnosine in exercise and disease: introduction to the International Congress held at Ghent University, Belgium, July 2011. Amino Acids 43:1–4

Hipkiss AR, Brownson C, Carrier MJ (2001) Carnosine, the anti- ageing, anti-oxidant dipeptide, may react with protein carbonyl groups. Mech Ageing Dev 15:1431–1445

Boldyrev, A. A., Aldini, G., Derave, W., & Suzuki, T. (2013). Carnosine and oxidative stress in cells and tissues. Antioxidants & redox signaling, 19(7), 1-4.

Hipkiss, A. R. (2019). Carnosine: why the fuss? Function, fate and evidence for its cardioprotective, anti-diabetic and life-prolonging effects. Aging and Disease, 10(2), 301.

Trombold, J. R., Reinfeld, A. S., Casler, J. R., & Coyle, E. F. (2011). The effect of β-alanine supplementation on neuromuscular fatigue in elderly (55–92 Years): a double-blind randomized study. Journal of the International Society of Sports Nutrition, 8(1), 7.