NUTRITION

Carbohydrate Oxidation: Know the Limits
by Eric Goulet
 

Knowing that the body possesses a limited ability to process sugars that are ingested during exercise, it would consequently be inappropriate to consume an unlimited quantity of carbohydrate during physical work. Most studies that have reported improved performance through carbohydrate consumption during exercise have given subjects between 25 and 60 grams of carbohydrate per hour (3). These rates of supplementation were enough to provide the additional 45 to 60 grams of carbohydrate needed to maintain blood glucose oxidation late in exercise.

These observations agreed with the finding that the rate of intravenous glucose infusion required to restore and maintain blood glucose availability and oxidation late in exercise is about 1 gram/minute (6). It is likely that these rates will differ from individual to individual. In fact, there are athletes who are able to exercise for long periods of time without an apparent decrease in blood glucose level (10) whereas other athletes will rely on blood glucose to a greater extent and consequently will be more susceptible to develop fatigue at an early stage (4).

During the first hour of exercise, irrespective of the quantity of carbohydrate consumed, only about 20 grams of carbohydrate are oxidized by the working muscles (18). Consequently, consuming more than this quantity of carbohydrate may possibly increase muscle glycogen utilization by attenuating the fall in plasma insulin concentration and therefore retarding fat mobilization (11). Since the ingestion of carbohydrate during exercise has no effect on the rate of muscle glycogen utilization (16), athletes are strongly encouraged to begin a prolonged exercise with a respectable quantity of muscle glycogen, irrespective of whether or not they will consume carbohydrate during that exercise (11,14). The increase in glycogen content of a muscle at the start of exercise has no effect on the rates of plasma glucose oxidation. Finally, while the ingestion of carbohydrate during exercise seems to "spare" liver glycogen conversion to plasma glucose and thus preventing hypoglyceamia, it does not delay the fatigue which is associated with low muscle glycogen content (20 mmol/kg wet weight muscle) (11).

TIMING OF CARBOHYDRATE INGESTION
The goals of supplementing carbohydrate at a rate of 1 gram/minute to maintain blood glucose level during the later stages of exercise can be achieved by either ingesting carbohydrate throughout the exercise period or by delaying carbohydrate ingestion until 30 minutes prior to the time that usually coincides with fatigue (3,5,6). However, if an athlete waits to consume carbohydrate until too late in exercise (less than 30 minutes prior to the onset of fatigue), the rate at which the ingested carbohydrate enters the blood as glucose may not be fast enough to maintain blood glucose oxidation, thus hypoglyceamia may result and performance will suffer.

Although there is large variability between individuals, carbohydrate ingestion should normally happen at least 30 minutes before fatigue would usually occur if no carbohydrate were consumed during exercise and this action might postpone fatigue by about only 45 minutes (3). During competition such as an Ironman, athletes should strive to consume at the least 45 to 60 grams of carbohydrate per hour for blood glucose concentration to be maintained and exercise performance to be improved (3). It is theoretically possible that carbohydrate consumption at high rates during intermittent or low-intensity exercise may promote glycogen resynthesis in resting muscle fibers (12). As a result, I think that athletes participating in adventure racing should be very cautious about their carbohydrate consumption throughout a race.

TYPES OF CARBOHYDRATE
Studies that have directly measured the effects produced by the ingestion of glucose, sucrose and maltodextrins, either alone or in combination, have found little differences in their ability to maintain blood glucose concentration, and carbohydrate oxidation or to improve performance (2,14,23,24). Studies that depart from this conclusion have been observed, but the amounts as well as the types of carbohydrate were altered (3). Recently maltodextrins have become a popular form of carbohydrate in sport drinks. Probably the major reason for this lies in the fact that this type of sugar is not very sweet tasting. Accordingly, solutions containing >10% carbohydrate are more palatable for most people (3). Contrasting with the sugars mentioned previously, the ingestion of fructose during prolonged exercise might not improve performance as a result of its slower rate of absorption by the gut and subsequent conversion to glucose in the liver (24). Therefore, its ingestion will result in lowered blood glucose and insulin concentrations and consequently reduced rates of carbohydrate oxidation. Also, it may be worth noting that fructose ingestion may increase the incidence of gastrointestinal distress in certain athletes.

LIQUID VERSUS SOLID CARBOHYDRATE
Few studies have examined the effects of solid carbohydrate ingestion on energy metabolism and endurance exercise performance. Fielding et al. (13) and Hargreaves et al. (16) both compared the metabolic effects and endurance performance times of a solid carbohydrate candy bar, which contained some protein and fat, with the ingestion of a placebo solution during intermittent exercise. Both studies showed that solid carbohydrate ingestion every hour during 4 hours of exercise elevated blood glucose and delayed fatigue, thus increasing performance. Mason et al. (22) and Ivy et al. (19) further determined that solid and liquid carbohydrate feedings during 2 to 3 hours of exercise were equally effective in maintaining blood glucose and insulin responses (22).

More recently, Robergs et al. (25) have evaluated the blood glucose responses and exercise performances resulting from solid carbohydrate ingestion with added protein and fat during prolonged steady-state exercise. This was of interest since most carbohydrate bars on the market contain some protein and fat. Although the stomach emptied solid carbohydrate feedings containing fat and protein more slowly when compared with liquid carbohydrate ingestion, the results of the study indicated that blood glucose levels and exercise performances remained similar between both trials. Collectively these studies, in my opinion, do not provide a complete evaluation of solid carbohydrate ingestion due to differences in exercise protocols, types of carbohydrate used and additional amounts of fat and protein in solid bars. Nonetheless, in the light of these studies, it can probably be concluded that liquid and solid carbohydrate feedings during exercise are equally efficient in maintaining blood glucose level, carbohydrate oxidation and performance.

PRACTICAL IMPLICATIONS FOR ATHLETES
On the basis of the most recent findings highlighted in this brief article, the following recommendations are made to athletes participating in endurance and ultra-endurance training or competition:
1. Because the rate of muscle glycogen utilization in working muscles is unaffected by the ingestion of carbohydrate, athletes are therefore encourage to maximize their muscle glycogen store before prolonged exercise, irrespective of the decision to ingest or not carbohydrate during that exercise;

2. As only about 20 grams of consumed carbohydrate are oxidized in the first hour of exercise, athletes are consequently invited to ingest a carbohydrate solution containing 3-5 grams 100 ml -1 every 10 minutes for the first 60-75 minutes of exercise. Ingesting more than this quantity of carbohydrate may possibly retard fat mobilization, thus increasing reliance on muscle glycogen utilization;

3. After the first 60 to 75 minutes of exercise, athletes must increase their carbohydrate consumption to match the peak rates (~1 gram minute -1) of plasma glucose utilization. Table 1 lists the volume as well as the concentration of various carbohydrate solutions to consume to match this recommendation;

4. To assure a high rate of gastric emptying, athletes must continuously maintain their stomach fluid volume at approximately 300 ml to 500 ml. This may be achieved by consuming 400 ml at the start of exercise and by repeated drinking of 100-150 ml every 10 minutes. Moreover, athletes are encouraged to consume a carbohydrate solution concentrated at no more than 8g 100 ml -1 and to ingest no more than 1000 ml of fluid/hour;

5. In the light of the studies, both liquid and solid carbohydrate seem to be equally effective in maintaining blood glucose level and carbohydrate oxidation during exercise. Consequently, athletes may choose to use a combination of liquid and solid carbohydrate consumption during exercise for i) the replenishment of lost water and electrolytes from the body and ii) reasons of satiety, respectively;

6. Finally, while carbohydrate ingestion during exercise decreases the rates of liver glycogen transformation to blood glucose and therefore preventing or retarding hypoglyceamia, it can not prevent fatigue or exhaustion that corresponds with critically low level of muscle glycogen (20 mmol/kg -1 wet weight muscle)

Eric

Additional background information and references to this article now follow.

BACKGROUND INFORMATION
It was as early as the 1920s during the prestigious Boston Marathon that exercise scientists first demonstrated that carbohydrate ingestion during prolonged exercise (>90 min.) enhanced performance in runners by preventing the development of hypoglycaemia (15,21). This finding in the field was then confirmed by Dill et al. (12) in 1932 during researches conducted in the distinguished Harvard Fatigue Laboratory in Boston. In fact, they showed that when their dogs, Joe and Sally, were fed 20g/carbohydrate/hour, their treadmill running endurance times to fatigue were improved by about 15 hours. Unfortunately, these results were largely ignored by the athletic community up until the early 1980s.

Between the 1930s and the early 1960s, fundamental research conducted in exercise physiology focussed primarily on the importance of replacing lost water during exercise to prevent dehydration and hyperthermia rather than consuming carbohydrate to enhance performance. In the 1960s, which coincided with the appearance of the muscle biopsy, Scandinavian researchers showed that the depletion of muscle glycogen was closely related to the development of fatigue and exhaustion (1). Accordingly, their primary concerns during these years were to find ways to increase pre-start exercise muscle glycogen concentration to increase time to fatigue; from there came the development of the glycogen loading technique known today by every ultra-endurance athlete.

In the early 1970s, Costill and Saltin (7) in a great scientific paper reinforced the popular belief that water consumption alone rather than water and carbohydrate ingestion during exercise was more appropriated; it was shown that the ingestion of a carbohydrate solution retarded the rate of gastric emptying if compared to water. Therefore, athletes were greatly discouraged to consume even mildly concentrated (<5g/100 ml) carbohydrate solutions during physical work. Finally, it was not until the early 1980s where the commercial interests in the sale of carbohydrate containing products to athletes was studied by now worldly known companies that physiologist became, once again, interested in studying the effects of carbohydrate ingestion during exercise. Not surprisingly, these studies only confirmed what had already been established in the 1920s and 1930s by Levine et al., Gordon et al. and Dill et al.: carbohydrate consumption during exercise was in fact retarding fatigue. Concomitant with this finding came the urgency for exercise physiologists to search for the best types of carbohydrate to consume; thus consequently to find the best nutritional strategies to adopt by athletes when they were exercising for long period of time (>90 min.).

EARLY STUDIES
In the 1970s the general belief was that carbohydrate ingestion during prolonged exercise was of little benefit except for people who were susceptible to suffer from neuroglucopenia (nausea, lethargy, lightheadedness, muscular twitching) (8). This in fact was true, except for some studies that suggested otherwise (2,20). Brooke et al. (2) in 1975 reported that both blood glucose concentrations and the rates of carbohydrate oxidation were well maintained during 3 to 4 hours of cycling at 67% Vo2 Max when trained cyclists consumed 90 grams of carbohydrate in the form of maltodextrins or rice pudding plus sucrose every 20 minutes during the exercise. Oppositely, but not surprising, was that both blood glucose concentrations and rates of carbohydrate oxidation were considerably diminished when the cyclists ingested the low-energy drinks or nothing at all during other trials. Finally, they reported that the overall working times were 148 minutes when fasted, 180 minutes when fed the low-energy drinks, a more than respectable 200 minutes when fed rice pudding plus sucrose and 214 minutes when they ingested maltodextrins. Ivy et al. (20) in 1979 had trained cyclists attempting to maximize their average power production during 2 hours of exercise on a braked cycle ergometer. The cyclists were fed approximately 13 grams of maltodextrins immediately prior and then at every 15 minutes during the first 90 minutes of exercise whereas in another trial they only ingested at these times points a placebo solution.

The results showed that the average power produced in any trials of 2 hours was not significantly different. However, during the trial where cyclists were fed carbohydrate, their total power output during the last 30 minutes of exercise was greater than during the placebo trial. It was not mention if the difference was significant, however. These authors suggested but did not systematically prove that carbohydrate ingestion during prolonged exercise might enhance performance. Consequently, Coyle et al. (10) in 1983 tried to definitely answer the interrogations proposed by previous studies by conducting an impressive research. They had experience cyclists exercise at 74% Vo2 Max for as long as possible on two occasions. Fatigue or exhaustion was defined as the point where they were forced to reduce their exercise intensity by 10% Vo2 Max below their initial level; that is, from 74% Vo2 Max to 64% Vo2 Max.

In one trial, cyclists received 1g of maltodextrins/kg bodyweight after 20 minutes of exercise, and an additional 0.25g/kg bodyweight after 60, 90 and 120 minutes. In another trial they only received a placebo solution. Carbohydrate ingestion significantly increased time to fatigue by about 23 minutes for the entire group of 10 athletes. But what was interesting is that this effect was only perceptible in 7 athletes whose blood glucose declined below 3 mmol/l during their placebo trial. When these athletes were fed maltodextrins during the carbohydrate trial, their time to fatigue increased by an average 33 minutes. Of great interest also was that the 3 subjects whose blood glucose concentration did not significantly decline during the placebo trial were not able to increase in a significant way their time to fatigue when fed carbohydrate. Consequently, this was the first study to clearly demonstrate that carbohydrate ingestion during exercise might delay fatigue and improve performance by maintaining blood glucose oxidation at sufficiently high rates during the later stages of exercise. Since that study, much of the researches done on this subject have shown that carbohydrate ingestion during prolonged exercise enhances performance.

GASTRIC EMPTYING OF FLUIDS
Between the 1970s and early 1990s, it was thought that the rate of gastric emptying was the primary factor limiting the rate of carbohydrate delivery to the blood and working muscles (7). Considerable emphasis in these years was placed on studying factors influencing the rate of gastric emptying of solutions such as the osmolality, caloric content, exercise intensity and temperature (7). More recently, a number of studies have shown that gastric volume instead of osmolality, caloric content and temperature of a solution is of more importance for regulating the rate of gastric emptying (18). These studies have clearly demonstrated that when the ingested volume of a given solution was increased (i.e.: from 600 to 1700 ml/h -1), the rate of gastric emptying also increased. When different carbohydrate solutions with different osmolalities were repeatedly ingested in sufficient amounts, their rates of gastric emptying were similar. Also, carbohydrate solution containing up to 8g 100 ml -1 have little influence on the rate of gastric emptying, especially if a high gastric volume is maintained. That is, when a fluid volume corresponding to 350 ml and 600 ml is continuously maintained in the stomach by repeated drinking of liquid. Thus, it appears quite possible to replace 30 to 60 grams of carbohydrate per hour while still being able to ingest between 600 ml to 1000 ml of fluid. Also, gastric emptying does not seem to be altered up to exercise intensities corresponding to 65-80% Vo2 Max. Finally, while cool drinks are more palatable, temperature has little influence on the rate of gastric emptying. In truth, data suggest that the oxidation of ingested carbohydrate after 60 to 90 minutes of exercise at 70% Vo2 Max is not limited by the rate of gastric emptying as previously thought, but either by its rate of appearance in the systemic blood or by the rate of muscle glucose oxidation (18). Hawley et al. (18) in 1992 infused glucose intravenously to athletes, which could theoretically supply an unlimited rate of radioactive 14C labelled glucose into the blood. Not surprisingly to my eyes, this action did not increase the rate of blood glucose oxidation during 2 hours of exercise at 70% Vo2 Max if compared to the oral ingestion of the same type of carbohydrate. Thus suggesting that the oxidation of carbohydrate is not limited by its rate of appearance in the systemic blood. Instead, it is thought that the peak rates of ingested carbohydrate oxidation are probably limited by the rate at which physiological concentrations of plasma glucose can be oxidized by the exercising muscles (11).

REFERENCES
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