Tag Archives: glycogen

Glycogen: an efficient storage form of energy in aerobic conditions

What is the net energy yield for the oxidation of a glucose unit from glycogen in aerobic conditions?

Aerobic Conditions: Glycogen Structure
Fig. 1 – Glycogen Structure

In aerobic conditions, the oxidation of a free glucose to CO2 and H2O (glycolysis, Krebs cycle and oxidative phosphorylation) leads to the net production of about 30 molecules of ATP.

Glucose from the action of glycogen phosphorylase: glucose-1-phosphate release (about 90% of the removed units).

Glycogen synthesis from free glucose costs two ATP units for each molecule; a glucose-1-phosphate is released by the action of glycogen phosphorylase with recovering/saving one of the two previous ATP molecules.
Therefore in aerobic condition, the oxidation of glucose starting from glucose-6-phosphate and not from free glucose yields 31 ATP molecules and not 30 (one ATP instead of two is expended in the activation phase, 30 ATP are produced during Krebs cycle and oxidative phosphorylation: 31 ATP gained).
The net rate between cost and yield is 1/31 (an energy conservation of about 97%).
The overall reaction is:

glycogen(n glucose residues) + 31 ADP + 31 Pi → glycogen(n-1 glucose residues) + 31 ATP + 6 CO2 + 6 H2O

If we combine glycogen synthesis, glycogen breakdown and finally the oxidation of glucose to CO2 and H2O we obtain 30 molecules of ATP per stored glucose unit, that is the overall reaction is:

glucose + 29 ADP + 30 Pi → 29 ATP + 6 CO2 + 6 H2O

Glucose from the action of debranching enzyme: free glucose release (about 10% of the removed units).

The net yield in ATP between glycogen synthesis and breakdown is two ATP molecules expended because of free glucose is released.
In this case the oxidation of glucose starts from the not-prephosphorylated molecule so we obtain 30 ATP molecules.
The net rate between cost and yield is 2/30 (a energy conservation of about 93,3%).
Considering the oxidation of the glucose units from glycogen to CO2 and H2O we have an energy conservation of:

1-(((1/31)*0,9)+((2/30)*0,1))=0,9643

Conclusion

In aerobic conditions, there is the conservation of about 97% of energy into the glycogen molecule, an extremely efficient storage form of energy.

References

Arienti G. “Le basi molecolari della nutrizione”. Seconda edizione. Piccin, 2003

Cozzani I. and Dainese E. “Biochimica degli alimenti e della nutrizione”. Piccin Editore, 2006

Giampietro M. “L’alimentazione per l’esercizio fisico e lo sport”. Il Pensiero Scientifico Editore, 2005

Mahan LK, Escott-Stump S.: “Krause’s foods, nutrition, and diet therapy” 10th ed. 2000

Mariani Costantini A., Cannella C., Tomassi G. “Fondamenti di nutrizione umana”. 1th ed. Il Pensiero Scientifico Editore, 1999

Nelson D.L., M. M. Cox M.M. Lehninger. Principles of biochemistry. 4th Edition. W.H. Freeman and Company, 2004

Stipanuk M.H.. “Biochemical and physiological aspects of human nutrition” W.B. Saunders Company-An imprint of Elsevier Science, 2000

Prolonged exercise and carbohydrate ingestion

Prolonged Exercise: Open Water Swimming
Fig. 1 – Open Water Swimming

During prolonged exercise (>90 min), like marathon, Ironman, cross-country skiing, road cycling or open water swimming, the effects of supplementary carbohydrates on performance are mainly metabolic rather than central and include:

  • the provision of an additional muscle fuel source when glycogen stores become depleted;
  • muscle glycogen sparing;
  • the prevention of low blood glucose concentrations.

How many carbohydrates should an athlete take?

The optimal amount of ingested carbohydrate is that which results in the maximal rate of exogenous carbohydrate oxidation without causing gastrointestinal discomfort”. (Jeukendrup A.E., 2008).

Prolonged exercise: which carbohydrates should an athlete take?

Until 2004 it was believed that carbohydrates ingested during exercise (also prolonged exercise) could be oxidized at a rate no higher than 1 g/min, that is, 60 g/h, independent of the type of carbohydrate.
Exogenous carbohydrate oxidation is limited by their intestinal absorption and the ingestion of more than around 60 g/min of a single type of carbohydrate will not increase carbohydrate oxidation rate but it is likely to be associated with gastrointestinal discomfort (see later).
Why?
At intestinal level, the passage of glucose (and galactose) is mediated by a sodium dependent transporter called SGLT1. This transporter becomes saturated at a carbohydrate intake about 60 g/h and this (and/or glucose disposal by the liver that regulates its transport into the bloodstream) limits the oxidation rate to 1g/min or 60 g/h. For this reason, also when glucose is ingested at very high rate (>60 g/h), exogenous carbohydrate oxidation rates higher 1.0-1.1 g/min are not observed.

The rate of oxidation of ingested maltose, sucrose, maltodextrins and glucose polymer is fairly similar to that of ingested glucose.

Fructose uses a different sodium independent transporter called GLUT5. Compared with glucose, fructose has, like galactose, a lower oxidation rate, probably due to its lower rate of intestinal absorption and the need to be converted into glucose in the liver, again like galactose, before it can be oxidized.

Prolonged Exercise: Maltodextrin and Fructose: Oxidation of Ingested Carbohydrates
Fig. 1 – Oxidation of Ingested Carbohydrates

However, if the athlete ingests different types of carbohydrates, which use different intestinal transporters, exogenous carbohydrate oxidation rate can increase significantly.
It seems that the best mixture is maltodextrins and fructose.

Note: the high rates of carbohydrate ingestion may be associated with delayed gastric emptying and fluid absorption; this can be minimized by ingesting combinations of multiple transportable carbohydrates that enhance fluid delivery compared with a single carbohydrate. This also causes relatively little gastrointestinal distress.

Conclusion

The ingestion of different types of carbohydrates that use different intestinal transporters can:

References

Carbohydrate ingestion during exercise of relatively short duration and high intensity

Intermittent high intensity exercise and carbohydrate ingestion

High Intensity: During-Exercise Nutrition
Fig. 1- During-Exercise Nutrition

Carbohydrate ingestion during intermittent high intensity or prolonged (>90 min) sub-maximal exercise can:

  • increase exercise capacity;
  • improve exercise performance;
  • postpone fatigue.

The intake of very small amounts of carbohydrates or carbohydrate mouth rinsing (for example with a 6% maltodextrin solution) may improve exercise performance by 2-3% when the exercise is of relatively short duration (<1 h) and high intensity (>75% VO2max), that is, an exercise not limited by the availability of muscle glycogen stores, given adequate diet.
The underlying mechanisms for the ergogenic effect of carbohydrates during this type of activity are not metabolic but may reside in the central nervous system: it seems that carbohydrates are detected in the oral cavity by unidentified receptors, promoting an enhanced sense of well-being and improving pacing.
These effects are independent of taste or sweet and non-sweet of carbohydrates but are specific to carbohydrates.

It should be noted that performance effects with drink ingestion are similar to the mouth rinse; therefore athletes, when they don’t complain of gastrointestinal distress when ingesting too much fluid, may have an advantage taking the drink (in endurance sports, dehydration and carbohydrate depletion are the most likely contributors to fatigue).

Conclusion

It seems that during exercise of relatively short duration (<1 h) and high intensity (>75% VO2max) it is not necessary to ingest large amounts of carbohydrates: a carbohydrate mouth rinsing or the intake of very small amounts of carbohydrates may be sufficient to obtain a performance benefit.

References

Hydration before endurance sports

Dehydration and endurance sports

Pre-hydration
Fig. 1 – Pre-hydration

In endurance sports, like Ironman, open water swimming, road cycling, marathon, or cross-country skiing, the most likely contributors to fatigue are dehydration and carbohydrate (especially liver and muscle glycogen) depletion.

Pre-hydration

Due to sweat loss needed to dissipate the heat generated during exercise, dehydration can compromise exercise performance.
It is important to start exercising in a euhydrated state, with normal plasma electrolyte levels, and attempt to maintain this state during any activity.
When an adequate amount of beverages with meals are consumed and a protracted recovery period (8-12 hours) has elapsed since the last exercise, the athlete should be euhydrated.
However, if s/he has not had adequate time or fluids/electrolytes volume to re-establish euhydration, a pre-hydration program may be useful to correct any previously incurred fluid-electrolyte deficit prior to initiating the next exercise.

Pre-hydration program

If during exercise the nutritional target is to reduce sweat loss to less than 2–3% of body weight, prior to exercise the athlete should drink beverages at least 4 hours before the start of the activity, for example, about 5-7 mL/kg body weight.
But if the urine is still dark (highly concentrated) and/or is minimal, s/he should slowly drink more beverages, for example, another 3-5 mL/kg body weight, about 2 hours before the start of activity so that urine output normalizes before starting the event.

It is advisable to consume small amounts of sodium-containing foods or salted snacks and/or beverages with sodium that help to stimulate thirst and retain the consumed fluids.
Moreover, palatability of the ingested beverages is important to promote fluid consumption before, during, and after exercise. Fluid palatability is influenced by several factors, such as:

  • temperature, often between 15 and 21 °C;
  • sodium content;
  • flavoring.

And hyper-hydration?

Hyper-hydration, especially in the heat, could improve thermoregulation and exercise performance, therefore, it might be useful for those who lose body water at high rates, as during exercise in hot conditions or who have difficulty drinking sufficient amounts of fluid during exercise.
However there are several risks:

  • fluids that expand the intra- and extra-cellular spaces (e.g. glycerol solutions plus water) greatly increase the risk of having to void during exercise;
  • hyper-hydration may dilute and lower plasma sodium which increases the risk of dilutional hyponatraemia, if during exercise, fluids are replaced aggressively.

Finally, it must be noted that plasma expanders or hyper-hydrating agents are banned by the World Anti-Doping Agency (WADA).

Conclusion

“Pre-hydrating with beverages, if needed, should be initiated at least several hours before the exercise task to enable fluid absorption and allow urine output to return toward normal levels. Consuming beverages with sodium and/or salted snacks or small meals with beverages can help stimulate thirst and retain needed fluids” (Sawka et al., 2007).

References

Carbohydrate ingestion 60 min before exercise

Introductory statement

Carbohydrates
Fig. 1 – Carbohydrates

An high-carbohydrate diet in the days before exercise, as well as ingestion of meals high in carbohydrate 3-4 h before exercise, better if with low glycemic index, can have positive effects on athlete’s performance.

For many years it has been suggested that ingestion of carbohydrates 30-60 min before exercise may adversely affect performance because it could cause hypoglycemia (blood glucose < 3.5 mmol/l or < 63 mg/l), a major contributor to fatigue. In fact, a typical athlete’s mantra is: “Avoid carbohydrate in the hour before exercise”!
What is the reason of that?
Glucose ingestion may cause hyperglycaemia followed by hyperinsulinaemina that may result in:

  • a rapid decline in glycemia 15-30 minutes after the onset of exercise, called rebound or reactive hypoglycaemia, most likely the result of:

I. an increase in muscle glucose uptake (due to the mobilization of GLUT-4 transporters by the action of insulin but also from physical activity itself);
II. the reduction in liver glucose output;

  • in addition, higher availability of carbohydrates to the muscle stimulates glycolysis and this, in combination to insulin-induced inhibition of lipolysis in both adipose tissue and muscle, results in a reduction in fat oxidation (apparently long-chain fatty acids, not medium-chain fatty acids). This may lead to premature glycogen depletion and early onset of fatigue (glycogen would be almost the only available fuel for working muscle).
    This effect is temporary, approximately lasting only for the first 20 min of exercise so, it is likely that this little glycogen breakdown has no significant effect on exercise performance.

Therefore, at least in theory, carbohydrate ingestion 60 minutes before exercise could affect performance but only two studies (Foster et al. 1979, e Kovisto et al. 1981) have reported a reduced endurance capacity while the majority of studies have reported no change or an improvement in performance.
To clarify these results, a systematic series of studies was done in trained subjects. The conclusion of these studies was that:

  • There is no effect of pre-exercise carbohydrate feeding on performance, even though in some cases hypoglycaemia did develop”.
  • There was no relationship between low blood glucose concentrations and performance”. (Jeukendrup and Killer S.C. 2010)

Conclusion

Ingestion of meals rich in carbohydrates 3-4 h before exercise is important for the increase of liver and muscle glycogen stores, or for their resynthesis in previously depleted muscle and liver.
Carbohydrate ingestion 30-60 min before exercise may be important in topping-up liver glycogen stores which serve to maintain blood glucose concentrations during exercise.
Based on the currently available scientific evidences, there is no reason to avoid carbohydrates 60 min before the onset of exercise, because they don’t seem to have any detrimental effect on performance.

References

Carbohydrate loading before competition

Carbohydrate loading and endurance exercise

Carbohydrate loading: Alberto Sordi and Spaghetti
Fig. 1 – Alberto Sordi and Spaghetti

Carbohydrate loading is a good strategy to increase fuel stores in muscles before the start of the competition.

What does the muscle “eat” during endurance exercise?

Muscle “eats” carbohydrates, in the form of glycogen, stored in the muscles and liver, carbohydrates ingested during the exercise or just before that, and fat.

During endurance exercise, the most likely contributors to fatigue are dehydration and carbohydrate depletion, especially of muscle and liver glycogen.
To prevent the “crisis” due to the depletion of muscle and liver carbohydrates, it is essential having high glycogen stores before the start of the activity.

What does affect glycogen stores?

  • The diet in the days before the competition.
  • The level of training (well-trained athletes synthesize more glycogen and have potentially higher stores, because they have more efficient enzymes).
  • The activity in the day of the competition and the days before (if muscle doesn’t work it doesn’t lose glycogen). Therefore, it is better to do light trainings in the days before the competition, not to deplete glycogen stores, and to take care of nutrition.

The “Swedish origin” of carbohydrate loading

Very high muscle glycogen levels (the so-called glycogen supercompensation) can improve performance, i.e. time to complete a predetermined distance, by 2-3% in the events lasting more than 90 minutes, compared with low to normal glycogen, while benefits seem to be little or absent when the duration of the event is less than 90 min.
Well-trained athletes can achieve glycogen supercompensation without the depletion phase prior to carbohydrate loading, the old technique discovered by two Swedish researchers, Saltin and Hermansen, in 1960s.
The researchers discovered that muscle glycogen concentration could be doubled in the six days before the competition following this diet:

  • three days of low carb menu (a nutritional plan very poor in carbohydrates, i.e. without pasta, rice, bread, potatoes, legumes, fruits etc.);
  • three days of high carbohydrate diet, the so-called carbohydrate loading (a nutritional plan very rich in carbohydrates).

This diet causes a lot of problems: the first three days are very hard and there may be symptoms similar to depression due to low glucose delivery to brain, and the benefits are few.
Moreover, with the current training techniques, the type and amount of work done, we can indeed obtain high levels of glycogen: above 2.5 g/kg of body weight.

The “corrent” carbohydrate loading

Carbohydrate Loading
Fig. 2 – Carbohydrate Loading: 2500 kcal Diet

If we compete on Sunday, a possible training/nutritional plan to obtain supercompensation of glycogen stores can be the following:

  • Wednesday, namely four days before the competition, moderate training and then dinner without carbohydrates;
  • from Thursday on, namely the three days before the competition, hyperglucidic diet and light trainings.

The amount of dietary carbohydrates needed to recover glycogen stores or to promote glycogen loading depends on the duration and intensity of the training programme, and they span from 5 to 12 g/kg of body weight/d, depending on the athlete and his activity. With higher carbohydrate intake you can achieve higher glycogen stores but this does not always results in better performance; moreover, it should be noted that glycogen storage is associated with weight gain due to water retention (approximately 3 g per gram of glycogen), and this may not be desirable in some sports.

References

Endurance sports and nutrition

What are endurance sports?

Endurance Sports
Fig. 1 – Endurance Sports

In the last years endurance sports, defined in the PASSCLAIM document of the European Commission as those lasting 30 min or more, are increasing in popularity and competitions as half marathons, marathons, even ultramarathons, half Ironmans, or Ironman competitions attract more and more people.
They are competitions which can last hours, or days in the more extreme case of ultramarathons.
Athletes at all levels should take care of training and nutrition to optimize performance and to avoid potential health threats.
In endurance sports the most likely contributors to fatigue are dehydration and carbohydrate depletion (especially liver and muscle glycogen).

Dehydration and endurance sports

Dehydration is due to sweat losses needed to dissipate the heat that is generated during exercise. To prevent the onset of fatigue from this cause, the nutritional target is to reduce sweat losses to less than 2–3% of body weight; it is equally important to avoid drinking in excess of sweating rate, especially low sodium drinks, to prevent hyponatraemia (low serum sodium levels).

Glycogen depletion and endurance sports

Muscle glycogen and blood glucose are the most important substrates from which muscle obtains the energy needed for contraction.
Fatigue during prolonged exercise is often associated with reduced blood glucose levels and muscle glycogen depletion; therefore, it is essential starting exercise/competition with high pre-exercise muscle and liver glycogen concentrations, the last one for the maintaining of normal blood glucose levels.

Other problems which reduce performance and can be an health threat of the athlete, especially in long-distance races, are gastrointestinal problems, hyperthermia and hyponatraemia.
Hyponatraemia has occasionally been reported, especially among slower competitors with very high intakes of low sodium drinks.
Gastrointestinal problems occur frequently, especially in long-distance races; both genetic predisposition and the intake of highly concentrated carbohydrate solutions, hyperosmotic drinks, as well as the intake of fibre, fat, and protein seem to be important in their occurrence.

References

Glycogen: an efficient storage form of energy in anaerobic conditions

What is the net energy yield for the oxidation of a glucose unit from glycogen in anaerobic conditions?

In anaerobic conditions, the oxidation of a free glucose to lactate leads to the net production of two molecules of ATP.

Anaerobic Conditions: Glycolysis to Lactate
Fig. 1 – Glycolysis to Lactate

Glucose from the action of glycogen phosphorylase: glucose-1-phosphate release (about 90% of the removed units).

Glycogen synthesis from free glucose costs two ATP units for each molecule; a glucose-1-phosphate is released by the action of glycogen phosphorylase, with recovering/saving of one of the two previous ATP molecules.
Therefore the oxidation of glucose to lactate starting from glucose-6-phosphate and not from free glucose yields three ATP molecules and not two (one ATP is expended in the activation stage instead of two, 4 ATP are produced in the third stage: three ATP gained).
The net rate between cost and yield is 1/3 (an energy conservation of about 66,7%).
The overall reaction is:

glycogen(n glucose residues) + 3 ADP + 3 Pi → glycogen(n-1 glucose residues) + 2 lactate + 3 ATP

If we combine glycogen synthesis, glycogen breakdown and finally glycolysis to lactate we obtain only one ATP molecule per stored glucose unit, that is the overall sum is:

glucose + ADP + Pi → 2 lactate + ATP

Glucose from the action of debranching enzyme: free glucose release (about 10% of the removed units).

The net yield in ATP between glycogen synthesis and breakdown is two ATP molecules expended because of free glucose is released.
In this case the oxidation of glucose starts from the not-prephosphorylated molecule and it yields two ATP molecules.
Therefore the net yield in ATP is zero.
Considering the oxidation of the glucose units from glycogen to lactate we have an energy conservation of:

1-(((1/3)*0,9)+((2/2)*0,1))=0,60

Conclusion

In anaerobic conditions, there is the conservation of about 60% of energy into the glycogen molecule, a good storage form of energy.

References

Arienti G. “Le basi molecolari della nutrizione”. Seconda edizione. Piccin, 2003

Cozzani I. and Dainese E. “Biochimica degli alimenti e della nutrizione”. Piccin Editore, 2006

Giampietro M. “L’alimentazione per l’esercizio fisico e lo sport”. Il Pensiero Scientifico Editore, 2005

Mahan LK, Escott-Stump S.: “Krause’s foods, nutrition, and diet therapy” 10th ed. 2000

Mariani Costantini A., Cannella C., Tomassi G. “Fondamenti di nutrizione umana”. 1th ed. Il Pensiero Scientifico Editore, 1999

Nelson D.L., M. M. Cox M.M. Lehninger. Principles of biochemistry. 4th Edition. W.H. Freeman and Company, 2004

Stipanuk M.H.. “Biochemical and physiological aspects of human nutrition” W.B. Saunders Company-An imprint of Elsevier Science, 2000

Carbohydrate mouth rinse and endurance exercise performance

Carbohydrate mouth rinse and performance responses

The importance of carbohydrates as an energy source for exercise is well known: one of the first study to hypothesize and recognize their importance was the study of Krogh and Lindhardt at the beginning of the 20th century (1920); later, in the mid ‘60’s, Bergstrom and Hultman discovered the crucial role of muscle glycogen on endurance capacity.
Nowdays, the ergogenic effects of carbohydrate supplementation on endurance performance are well known; they are mediated by mechanisms such as:

  • a sparing effect on liver glycogen;
  • the maintenance of glycemia and rates of carbohydrate oxidation;
  • the stimulation of glycogen synthesis during low-intensity exercise ;
  • a possible stimulatory effect on the central nervous system.

However, their supplementation, immediately before and during exercise, has an improving effect also during exercise (running or cycling) of a shorter and more intense nature: >75% VO2max (maximal oxygen consumption) and ≤1 hour, during which euglycaemia is rarely challenged and adequate muscle glycogen store remains at the cessation of the exercise.

Hypothesis for carbohydrate mouth rinse

In the absence of a clear metabolic explanation it was speculated that ingesting carbohydrate solutions may have a ‘non-metabolic’ or ‘central effect’ on endurance performance. To explore this hypothesis many studies have investigated the performance responses of subjects when carbohydrate solutions (about 6% carbohydrate, often maltodextrins) are mouth rinsed during exercise, expectorating the solution before ingestion.
By functional magnetic resonance imaging and transcranial stimulation it was shown that carbohydrates in the mouth stimulate reward centers in the brain and increases corticomotor excitability, through oropharyngeal receptors which signal their presence to the brain.
Probably salivary amylase releases very few glucose units from maltodextrins which is probably what is needed in order to activate the purported carbohydrate receptors in the oropharynx (no glucose transporters in the oropharynx are known).
However, the performance response appears to be dependent upon the pre-exercise nutritional status of the subject: most part of the studies showing an improving effect on performance was conducted in a fasted states (3- to 15-h fasting).
Only one study has shown improvements of endurance capacity   in both fed and fasted states by carbohydrate mouth rinse, but in non-athletic subjects.

References

Beelen M., Berghuis J., Bonaparte B., Ballak S.B., Jeukendrup A.E., van Loon J. Carbohydrate mouth rinsing in the fed state: lack of enhancement of time-trial performance. Int J Sport Nutr Exerc Metab 2009;19(4):400-9 [Abstract]

Bergstrom J., Hultman E. A study of glycogen metabolism during exercise in man. Scand J Clin Invest 1967;19:218-28 [Abstract]

Bergstrom J., Hultman E. Muscle glycogen synthesis after exercise: an enhancing factor localized in muscle cells in man. Nature 1966;210:309-10 [Abstract]

Painelli V.S., Nicastro H., Lancha A. H.. Carbohydrate mouth rinse: does it improve endurance exercise performance? Nutrition Journal 2010;9:33 [Abstract]

Fares E.J., Kayser B. Carbohydrate mouth rinse effects on exercise capacity in pre- and postprandial States. J Nutr Metab 2011;385962. doi: 10.1155/2011/385962. Epub 2011 Jul 27 [Abstract]

Krogh A., Lindhard J. The relative value of fat and carbohydrate as sources of muscular energy. Biochem J 1920;14:290-363 [PDF]

Rollo I. Williams C. Effect of mouth-rinsing carbohydrate solutions on endurance performance. Sports Med. 2011;41(6):449-61 [Abstract]

Blood glucose levels and liver

Blood glucose levels and hepatic glycogen

One of the main functions of the liver is to participate in the maintaining of blood glucose levels within well defined range (in the healthy state before meals 60-100 mg/dL or 3.33-5.56 mmol/L). To do it the liver releases glucose into the bloodstream in:

  • fasting state;
  • between meals;
  • during physical activity.

Blood glucose levels and hepatic glucose-6-phosphatase

In the liver, glycogen is the storage form of glucose which is released from the molecule not as such, but in the phosphorylated form i.e. with charge, the glucose-1-phosphate (this process is called glycogenolysis). The phosphorylated molecule can’t freely diffuse from the cell, but in the liver it is present the enzyme glucose-6-phosphatase that hydrolyzes glucose-6-phosphate, produced from glucose-1-phosphate in the reaction catalyzed by phosphoglucomutase, to glucose (an irreversible dephosphorylation).

glycogen(n glucose residues) + Pi → glucose-1-phosphate + glycogen(n-1 glucose residues)

glucose-1-phosphate ↔ glucose-6-phosphate

glucose-6-phosphate + H2O → glucose + Pi

Then, glucose can diffuse from the hepatocyte, via a transporter into the plasma membrane called GLUT2, into the bloodstream to be delivered to extra-hepatic cells, in primis neurons and red blood cells for which it is the main, and for red blood cells the only energy source (neurons, with the exception of those in some brain areas that can use only glucose as energy source, can derive energy from another source, the ketone bodies, which becomes predominant during periods of prolonged fasting).

Note: the liver obtains most of the energy required from the oxidation of fatty acids, not from glucose.

Glucose-6-phosphatase is present also in the kidney and gut but not in the muscle and brain; therefore in these tissues glucose-6-phosphate can’t be released from the cell.
Glucose-6-phosphatase plays an important role also in gluconeogenesis.

Glucose-6-phosphatase is present into the membrane of endoplasmic reticulum and the hydrolysis of glucose-6-phosphate occurs into its lumen (therefore this reaction is separated from the process of glycolysis). The presence of a specific transporter, the glucose-6-phosphate translocase, is required to transport the phosphorylated molecule from citosol into the lumen of endoplasmic reticulum. Although a glucose transporter is present into the membrane of endoplasmic reticulum, most of the released glucose is not transported back into the cytosol of the cell but is secreted into the bloodstream. Finally, an ion transporter transports back into the cytosol the inorganic phosphate released into the endoplasmic reticulum.

References

Arienti G. “Le basi molecolari della nutrizione”. Seconda edizione. Piccin, 2003

Cozzani I. and Dainese E. “Biochimica degli alimenti e della nutrizione”. Piccin Editore, 2006

Giampietro M. “L’alimentazione per l’esercizio fisico e lo sport”. Il Pensiero Scientifico Editore, 2005

Mahan LK, Escott-Stump S.: “Krause’s foods, nutrition, and diet therapy” 10th ed. 2000

Mariani Costantini A., Cannella C., Tomassi G. “Fondamenti di nutrizione umana”. 1th ed. Il Pensiero Scientifico Editore, 1999

Nelson D.L., M. M. Cox M.M. Lehninger. Principles of biochemistry. 4th Edition. W.H. Freeman and Company, 2004

Stipanuk M.H.. “Biochemical and physiological aspects of human nutrition” W.B. Saunders Company-An imprint of Elsevier Science, 2000