Fat metabolism and exercise -
WAT in mammalian body forms a few depots and can generally be classified into visceral and subcutaneous AT differing in terms of the composition of triacylglycerols-forming FA Garaulet et al.
Published data about the effects of exercise on FA composition in human AT are limited. An early studies revealed a decrease in oleic acid and increase in linoleic acid n-6 content in subcutaneous AT after chronic training Allard et al.
The decrease in the level of , the main monounsaturated FA MUFA , which was observed in both studies mentioned above, might be associated with reduced activity of stearoyl-CoA desaturase SCD1 in AT Nikolaidis and Mougios, Since metabolic disorders were shown to be associated with enhanced synthesis of and other MUFA by SCD1 Mika et al.
Published evidence suggests that physical training may contribute to a preferential mobilization of some FAs from AT. Already after 2 weeks of the training in senior oarsmen, the authors observed a significant decrease in total serum triacylglycerols and cholesterol, along with changes in the FA profile of AT: a decrease in palmitoleic acid and an increase in stearic acid content, comparing to previously untrained controls Danner et al.
A more recent study demonstrated that 6 months of increased physical activity contributed to a significant increase in n-6 in overweight elderly subjects, while no such effect was observed in untrained controls Sjögren et al.
Taken altogether, this sparse evidence from human suggests that chronic exercise may contribute to a decrease in content, with concomitant increase in n-6 and Since is the main FA found in triacylglycerols Ntambi and Miyazaki, ; Liu et al.
However, regular exercise training seems to reduce systemic inflammation Görgens et al. More data in this matter originate from rodent models, and based on this evidence we may compare the effect of exercise on FA composition in various WAT depots, as well as in BAT May et al.
Most of the animal studies demonstrated that chronic exercise contributed to a decrease in MUFA content, which is consistent with the observations made in humans Nikolaidis and Mougios, Regarding polyunsaturated FAs PUFAs , most animal studies showed an increase in their content, especially n-6 PUFAs; however, in some studies, the post-exercise levels of PUFAs were lower than prior to the exercise or remained unchanged.
The chronic exercise-induced changes in PUFA content in AT are depot-specific Bailey et al. Petridou et al. Among the MUFAs of visceral fat, chronic exercise contributed to a decrease in , but not in Petridou et al.
In recent study conducted by May et al. The study demonstrated that while the exercise contributed to a significant increase in MUFA level and a significant decrease in PUFA content in WAT phospholipids, an inverse phenomenon, i.
Moreover, a significant decrease in triacylglycerol content of SFAs, MUFAs and PUFAs in BAT and triacylglycerol content of PUFAs in WAT was observed May et al. However, it should be stressed that in that study, FA content was expressed in nmol per mg of protein, rather than as a percentage of total FAs as in previously mentioned experiments.
These findings suggest that post-exercise changes in FA composition of AT are not only depot- but also lipid molecule-specific. In both WAT and BAT, physical exercise contributed to a significant decrease in the total content of triacylglycerols, but with a concomitant increase in the level of triacylglycerols containing long-chain FAs 58—60 total carbons May et al.
The effects of exercise on FA composition in AT and other AT parameters in human and animal studies is summarized in Table 1. Table 1. The summary of effects of exercise on adipose tissue metabolism and adipokine secretion. The post-exercise decrease in triacylglycerols content in AT is with no doubt a consequence of enhanced lipolysis.
The process, initiated by adipose triglyceride lipase ATGL , is then continued by hormone-sensitive lipase HSL , upon phosphorylation thereof; eventually, the last FA chain is hydrolyzed by monoacylglycerol lipase MAGL Chen et al.
While the rate of lipolysis is decreased by obesity and high-fat diet, chronic exercise was shown to normalize the markers of this process, phosphorylated HSL and ATGL, in mice that have been previously maintained on a high-fat diet Chen et al. Surprisingly, however, Holland et al.
In contrast, chronic exercise was shown to stimulate the activity of lipolytic enzymes in the adipocytes of obese mice Woo and Kang, , and a recent study demonstrated that endurance exercise contributed to an increase in triacylglycerol lipase activity in human AT, especially during the first 10 min of the training Petridou et al.
An upregulation of HSL after chronic exercise was also mentioned in a review paper published by Steinberg Moreover, irisin, an adipokine released by working muscles, was shown to induce the expression of ATGL and HSL in 3T3L1 adipocytes Gao et al.
Physical activity may also modulate FA synthesis, desaturation and elongation. The reduction of MUFA content after chronic exercise reported by many authors might be a consequence of a decrease in FA desaturation by SCD1 Nikolaidis and Mougios, However, this conclusion is based on the desaturation indices calculated from SFA and MUFA contents in AT.
Thus, it cannot be excluded that those parameters were also influenced by preferential uptake and release of certain FAs in AT during exercise Halliwell et al. One study demonstrated that chronic exercise did not affect the expression of SCD1 in mice subcutaneous WAT, but contributed to lesser activity of this enzyme in BAT May et al.
Also in human subcutaneous AT, the expression of SCD1 gene remained unchanged after the chronic exercise Sjögren et al. Published data about the exercise-induced changes in the activity of other lipogenic enzymes are inconclusive.
According to May et al. Similarly, a 6-week exercise resulted in an increase in ACC protein level in visceral WAT of rats Holland et al. In contrast, Rocha-Rodrigues et al. The same study demonstrated a post-exercise increase in the expressions of enzymes involved in PUFA metabolism, FA desaturase 1 and elongase 5 Rocha-Rodrigues et al.
In line with those findings, May et al. Taken altogether, the abovementioned findings suggest that the effect of exercise on the expression of enzymes involved in lipid metabolism may vary depending on FA group and AT depot.
Muscle work during the exercise may activate a signaling cascade; specifically, myokines released from the muscle cells may trigger a release of adipokines, signaling molecules synthesized in the AT.
Aside from the production of adipokines, AT can also synthesize many myokines, among others IL-6, MCP1, TNFα, visfatin and myostatin, which are collectively referred to as adipomyokines Görgens et al.
Thus, plasma level of adipomyokines does not necessarily reflect solely the pool which is synthesized in the AT and acts on the muscles, and the origin of each molecule should be identified at a cellular level.
Adiponectin is an insulin-sensitizing hormone that enhances FA oxidation in the muscles and downregulates the synthesis of lipids and glucose in the liver Swierczynski and Sledzinski, The evidence from both human and animal studies analyzing the effects of exercise on serum adiponectin level is inconclusive; chronic exercise was either shown to increase the serum concentration or expression in AT of this adipokine or did not affect it at all Kato et al.
Available data imply that the release of adiponectin from human AT may depend on exercise intensity Görgens et al. Another adipokine, leptin, is synthesized primarily in the AT, regulates appetite and boosts peripheral metabolism Swierczynski and Sledzinski, Chronic exercise was shown to contribute to a decrease in serum leptin concentration, but this effect was associated with the reduction of AT mass Lehnig and Stanford, However, previous studies demonstrated that excess body weight was associated with leptin resistance Swierczynski and Sledzinski, , and chronic exercise might improve leptin sensitivity Kang et al.
Thus, the simultaneous reduction of serum leptin level and AT mass does not necessarily correspond to a decrease in the activity of this adipokine. Serum concentration of IL-6, acting as an anti-inflammatory myokine, was shown to increase substantially after acute exercise, and this effect was demonstrated to result from local synthesis of IL-6 by the working muscles.
However, the level of IL-6 after chronic exercise was decreased or remained unchanged Görgens et al. Also, the expression of IL-6 in the AT did not change or was reduced, depending on the type of chronic exercise Bruun et al. Generally, moderate chronic exercise seems to be associated with a decrease in the release of pro-inflammatory cytokines, such as TNF-α, leptin and MCP-1, from the AT and working muscles; this may contribute to attenuation of systemic inflammation Görgens et al.
A recent study demonstrated that chronically exercised rats showed enhanced expression of apelin, an adipomyokine that decreases insulin resistance Kazemi and Zahediasl, Apelin induced glucose uptake by AT, but at the same time decreases triglyceride amounts in mouse AT and lipid storage in 3T3-L1 preadipocytes Indrakusuma et al.
Also, serum concentration of resistin, an adipokine promoting insulin resistance, was shown to decrease in rats subjected to an chronic exercise Shirvani and Arabzadeh, Altogether, these findings suggest that chronic exercise may improve the profile of adipokines released from the AT, and thus, may be beneficial for health.
Aside from the metabolic processes discussed above lipolysis, FA uptake, FA synthesis , FAs are also oxidized in mitochondria, in a process referred to as β-oxidation. A number of previous studies demonstrated that chronic exercise enhanced mitochondrial activity in visceral and subcutaneous AT, in both rodents Stallknecht et al.
The process of mitochondrial β-oxidation in the AT is not as intensive as in the muscles but still can provide an extra pool of energy for adipocytes after the exercise. Furthermore, there is one AT depot that shows greater mitochondrial activity than visceral and subcutaneous WAT; this is BAT which contains numerous mitochondria whereby FAs undergo oxidization, becoming a source of energy for thermogenesis.
The main protein involved in thermogenesis is uncoupling protein 1 UCP1 , mediating proton leakage across the inner mitochondrial membrane into the mitochondrial matrix, and thus, playing a role in heat production Lehnig and Stanford, This phenotypic and functional switch includes an increase in mitochondrial respiration and enhancement of UCP1 protein expression, as well as the upregulation of other genes characteristic for BAT Wu et al.
In one study, ablation of beige adipocytes resulted in the development of obesity and insulin resistance in mice; this implies that these cells may play a role in the regulation of systemic energy metabolism Stanford and Goodyear, However, it needs to be stressed that BAT in adult humans resembles murine beige AT, rather than the BAT Wu et al.
According to one hypothesis, the process may be mediated by myokines and small molecules released from working muscles Lehnig and Stanford, , specifically by irisin Boström et al.
Physical exercise stimulates lipolysis, decreases FA uptake by the adipocytes, exerts an effect on FA composition within the AT and modulates the expression of enzymes involved in FA synthesis, elongation and desaturation.
As a result of all those metabolic processes, physically active persons can maintain adequate volume of AT. Chronic exercise influences the release of adipokines, which may attenuate systemic inflammation and prevent insulin resistance. Moreover, transplantation of AT from trained to untrained mice was shown to improve glucose tolerance Stanford et al.
Taken altogether, these findings imply that the exercise-induced changes in AT metabolism may exert a beneficial effect on global metabolic health. AM, FM, RB, VD, and TS studied the literature and wrote the manuscript.
All authors accepted the final version of manuscript. This study was supported by the Medical University of Gdańsk grant no. ST40 , National Science Centre of Poland grant no.
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
The handling Editor is currently co-organizing a Research Topic with one of the authors VD, and confirms the absence of any other collaboration. Allard, C. Changes in adipose tissue and increased serum cholesterol of coronary patients following training.
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Jeukendrup AE 1 ,. Saris WH ,. Wagenmakers AJ. Affiliations 1. Nutrition Research Center, Department of Human Biology, Maastricht University, The Netherlands.
Authors Jeukendrup AE 1. Share this article Share with email Share with twitter Share with linkedin Share with facebook. Abstract This is the first part in a series of three articles about fat metabolism during exercise. In this part the mobilization of fatty acids and their metabolism will be discussed as well as the possible limiting steps of fat oxidation.
It is known for a long time that fatty acids are an important fuel for contracting muscle. After lipolysis, fatty acids from adipose tissue have to be transported through the blood to the muscle.
Fatty acids derived from circulating TG may also be used as a fuel but are believed to be less important during exercise. In the muscle the IMTG stores may also provide fatty acids for oxidation after stimulation of hormone sensitive lipase.
In the muscle cell, fatty acids will be transported by carrier proteins FABP , and after activation, fatty acyl CoA have to cross the mitochondrial membrane through the carnitine palmytoyl transferase system, after which the acyl CoA will be degraded to acetyl CoA for oxidation.
The two steps that are most likely to limit fat oxidation are fatty acid mobilization from adipose tissue and transport of fatty acids into the mitochondria along with mitochondrial density and the muscles capacity to oxidize fatty acids. Longer-Term Effects of the Glycaemic Index on Substrate Metabolism and Performance in Endurance Athletes.
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