LEADING ARTICLE sports Med. 19 (4): 240-250.

1995
0112-1642/95/0004-0240/S05.5O/0

© Adis International Lm~ed. An rights reserved.

Aerobic Glycolytic and Aerobic Lipolytic

Power Systems
A New Paradigm with Implications for
Endurance and Ultraendurance Events
John A. Hawleyl and Will G. Hopkins2
1 Department of Physiology, University of Cape Town Medical School, Cape Town, South Africa
2 Department of Physiology and School of Physical Education, University of Otago, Dunedin,
New Zealand

In the traditional view of exercise metabolism, training regimens for high-intensity endurance
the energy for muscular contraction is produced by pursuits should continue to be aimed at enhancing
3 independent systems: the phosphagen, anaerobic the aerobic glycolytic system, but for ultraendur-
(or oxygen-independent) glycolytic, and aerobic ance events such regimens should be aimed at en-
(or oxygen-dependent) systems.£I-4) The phospha- hancement of the aerobic lipolytic system.
gen system uses intracellular stores of adenosine
triphosphate (ATP) and creatine phosphate (CP) to 1. Power Systems
provide the power needed for maximal bouts of
strength or speed that last for a few seconds. [5-9) For In most texts the term energy system is used to
high-intensity exercise of up to I minute in dura- refer to a physiological entity that produces ATP to
tion, energy is provided mostly by the anaerobic sustain cellular processes, including muscular con-
glycolytic system, which breaks down muscle traction. What criteria must be satisfied for the 3
glycogen without consuming oxygenp,IO-13) Ex- existing energy systems to be considered distinct
from each other? We are unaware of any previous
ercise lasting longer than I minute is powered
formal attempt to address this question, so we have
mostly by the aerobic system, which uses oxygen
devised our own criteria by examining the differ-
in the breakdown of carbohydrate (glycogen, glu-
ences between the 3 systems. We suggest that these
cose), lipids (triglycerides, fatty acids and ketones)
systems are considered distinct because: (1) they
and, to a very small extent, amino acidsp,3,14-16)
have substantially different metabolic pathways;
We believe that this traditional view of only 3 (2) their relative contributions to the energy re-
energy systems is too simplistic. Instead, we pro- quired for exercise depend on the intensity and
pose that the oxidation of carbohydrates and lipids duration of the exercise; and (3) their contributions
should be regarded as the basis of 2 functionally can be modified selectively by training or other
distinct aerobic power systems: the aerobic glyco- interventions. Debate amongst exercise scientists
lytic system (which oxidises carbohydrate for on whether these criteria are necessary and suffi-
high-intensity endurance events) and the aerobic cient to define distinct energy systems would be
lipolytic system (which oxidises lipid to provide appropriate. In the meantime, we suggest that the
most of the energy for longer, less intense endur- systems defined by these particular criteria be re-
ance and ultraendurance activities). A practical im- ferred to as something other than energy systems,
plication of this new paradigm is that dietary and to avoid confusion with the existing nomenclature.
Aerobic Glycolytic and Lipolytic Systems 241

From the point of view of athletes and sports CoA. A separate set of enzymes controls the con-
scientists, the main function of an energy system is version of intramuscular and extramuscular carbo-
to produce mechanical power rather than the en- hydrate to pyruvate and then to acetyl CoA. There-
ergy required to sustain metabolic processes. after, the pathways for oxidation of the acetyl
Therefore, we will use the term power system to moieties from either lipid or carbohydrate are the
describe a system that satisfies our criteria. We will same.
show that our putative aerobic glycolytic and aero- Each of the pathways has unique elements, but
bic lipolytic systems, along with the existing phos- some also have common components. Thus, the
phagen and anaerobic glycolytic systems, satisfy anaerobic and aerobic glycolytic systems share the
these criteria. pathway for the breakdown of glycogen to py-
ruvate and, as discussed above, the aerobic glyco-
1.1 Distinct Power Systems Have lytic and lipolytic systems share the pathways for
Different Metabolic Pathways oxidative phosphorylation in the mitochondria.
We note that under the existing paradigm of only
Different metabolic pathways are an essential 3 systems, the anaerobic glycolytic and aerobic
feature of the 3 existing energy systems. Indeed, systems share the pathway for the breakdown of
the names of the systems represent descriptive glycogen to pyruvate. Overlap of pathways is,
summaries of the pathways each uses to produce therefore, not a sufficient reason to reject the new
ATP. Thus, the phosphagen system is named for the scheme.
2 high-energy phosphate compounds (ATP and Although the pathways shown in figure I ap-
CP) that are used to store energy for this system; pear to be independent, the figure conceals a myr-
the anaerobic glycolytic system derives its name iad of complex interactions between the pathways
from the pathway that can produce ATP from the mediated by enzymes, metabolites, cofactors and
breakdown of carbohydrate without the use of oxy- intracellular messengers. Thus, at the biochemical
gen; and the aerobic system is named for the fact level, all of the pathways are interdependent. The
that it generates ATP from the oxidative breakdown crucial issue is whether the macroscopic function
of fuels. of these pathways under different conditions pro-
The pathways in the phosphagen and anaerobic vides sufficient grounds for the power systems to
glycolytic systems, along with the pathways for the be considered distinct. We address this issue in sec-
metabolism of carbohydrate and lipid in the aero- tions 1.2 and 1.3.
bic system, are summarised in figure 1. For the
purposes of this article, the small contribution that 1.2 Relative Contributions of Distinct
the aerobic breakdown of amino acids makes to the Power Systems Differ with Exercise
metabolic demands of exercise has been ignored. Intensity and Duration
The pathways depicted in figure 1 are widely
accepted. However, a new feature is the terminol- The contributions of the power systems to the
ogy for the 2 aerobic pathways, which we have total energy requirements of exercise have been
devised to highlight the differences between the studied most frequently during exercise in which
substrates metabolised in these and in the other individuals perform either the maximal amount of
pathways. The differences between the pathways work in a predetermined time or a given amount of
for the breakdown and oxidation of lipids and work in the shortest possible timeJIO·12,17.22] We
carbohydrates are not insubstantial: a set of cyto- will use the term maximal to define both kinds of
plasmic and membrane-bound enzymes is respon- exercise, even though for exercise lasting more
sible for the ~-oxidation of lipids to fatty-acyl than 1 minute, maximal effort may not be attained
co-enzyme A (CoA), the transport of fatty-acyl until the end of the exercise period. Most compet-
CoA into mitochondria, and its conversion to acetyl itive athletic events require such maximal exercise.

© Adis International Umited. All rights reserved. Sports Med. 19 (4) 1995
242 Hawley & Hopkins

Phosphagen Anaerobic
glycolytic

Glycogen
Creatine
phosphate
~~ATP
l~m Pyruvate

Creatine

Lactate

Aerobic Aerobic
glycolytic lipolytic

Glycogen Lipid

Glucose,
~c:::::> ATP
Pyruvate Lipid
~
Fatty acyl-CoA
lactate

Fig. 1. The 4 metabolic pathways for the production of adenosine triphosphate in muscle cells. Abbreviations: ATP =adenosine
=
triphosphate; CoA co-enzyme A.

Figure 2 shows the contributions of the power exercise,[19-21 ,29) whereas lipid makes substantial
systems to the total work performed during maxi- contributions to the energy requirements of more
mal exercise lasting::::l second to::::l day. The fig- prolonged activities.[2,3,15,29-38) On the basis of the
ure is similar to those depicted in many textbooks few relevant published studies,[29,35,37,38) we esti-
on exercise physiology,[I,23-26) but we have modi- mate that the total contributions made by aerobic
fied and updated the time courses to take into ac- glycolysis and aerobic lipolysis are equal after 3 to
count recent data showing that the crossover points 5 hours of maximal exercise, although this duration
for the phosphagen, anaerobic glycolytic and aero- is modified by training, diet and caloric supple-
bic glycolytic contributions occur at ::::6 seconds mentation during the exercise (see section 1.3).
and ::::1 minute in athletes.l 5,6,1 1,27,28) More impor- When the exercise is lower in intensity than that
tantly, we have separated the contributions derived necessary to achieve maximal work or work in
from the aerobic breakdown of carbohydrates and minimal time, the relative contributions of the
lipids to illustrate how these too are affected by the power systems change somewhat from those
duration of exercise. Carbohydrate provides most shown in figure 2. The phosphagen system is still
of the energy for short-term maximal endurance the major source of ATP for activities lasting 1 or

© Adis International Um~ed. All rights reserved. Sports Med. 19(4) 1995
Aerobic Glycolytic and Lipolytic Systems 243

2 seconds, so its contribution for such short activ- is maintained at this intensity, the energy derived
ities is unaffected by intensity. For slightly longer from lipid oxidation gradually increases, with a
activities, the contribution of the anaerobic glyco- corresponding decline in carbohydrate oxida-
lytic system will not only be delayed by the slower tion.[29,31,33-35,37) The greater contribution from
rate of change of concentrations of regulatory me- lipid oxidation to total energy metabolism with in-
tabolites, but will also be truncated by the more creasing exercise duration is most likely due to a
rapid rate of activation of the aerobic system at combination of factors, including a gradual deple-
lower exercise intensities,l39) tion of endogenous carbohydrate reserves[2,3,31)
For sub maximal endurance activities, utilisa- and changes in the hormonal milieu,l52) During
tion of lipid relative to carbohydrate in the aerobic prolonged exercise, plasma glucose and insulin
pathways increases as the intensity of endurance
levels faW 53 ,54) and the levels of circulating glu-
exercise decreases, a well known effect that has cagon,[52-54) catecholamines[53,55-57) and growth
recently been termed the crossover concept. [40) The
hormone[55-57) rise; these responses promote the
exercise intensity at which lipid and carbohydrate
mobilisation of fatty acids and hepatic glucose pro-
oxidation are equal defines the crossover: above it,
duction.[52)
the main fuel is carbohydrates[15,41-43); below it, the
main fuel is lipids,l15,44,45) The shift from lipid to What must have seemed an undifferentiated pat-
carbohydrate at high power outputs is regulated tern of utilisation of carbohydrate and lipid in ex-
mainly by changes in concentrations of metabo- ercise of moderate intensities and durations may
lites within the active muscle fibres,[46-48) and by have led earlier authors to place aerobic glycolysis
changes in muscle fibre activity from predomi- and aerobic lipolysis under the one rubric of the
nantly type I oxidative fibres at low intensity to aerobic system. In reality, there are clear-cut ef-
increasing recruitment of type II glycolytic motor fects of exercise intensity and duration: the lower
units at higher intensities.[49-51) the intensity or the longer the duration, the greater
During relatively short-term exercise, the cross- the dependence on lipid relative to carbohydrate.
over occurs at 60 to 70% of maximum oxygen up- Our putative aerobic glycolytic and lipolytic
take CV0 2max ) in trained athletes,l38,40) If exercise power systems therefore satisfy the requirement

100
#: Phosphagen
Q)
c:
0
"0
~
0
~

~
.8
c:
:s
0

.0
.;::
E
0
u
0
0.1 10 100 1000
Duration of exercise (min)

Fig. 2. The contribution of the 4 power systems to the maximal total work performed during exercise of a given duration. Adaptation
to a high-fat diet shifts the aerobic lipolytic curve and the corresponding right-hand segment of the aerobic glycolytic curve to the left;
adaptation to a high-carbohydrate (high-CHO) diet shifts these curves to the right. The dashed sections marked with a ? indicate
present uncertainty about the relative contributions.

© Adis International Limited. All rights reserved. Sports Med. 19 (4) 1995
244 Hawley & Hopkins

that their contributions to exercise change with ex- pathway also engage either the phosphagen or aero-
ercise duration and intensity. bic glycolytic power systems (see figure 2). Never-
theless, it appears that very short, intense workouts
1.3 Contributions of Distinct Power Systems that train only the phosphagen and anaerobic glyco-
can be Modified Selectively lytic systems do not result in enhanced maximal
aerobic performance. [S,60,66)
If 2 power systems were tightly coupled by Certain acute physiological interventions also
underlying biochemical or physiological mecha- provide evidence of the independence of the 3 tra-
nisms, any intervention that modified the power ditional systems. A treatment aimed at improving
provided by 1 system would produce a similar oxygen transport that appears to enhance only the
change in the power of the other. We would hesitate aerobic system is blood doping.[67-69) Breathing
to regard such coupled systems asfunctionally dis- pure oxygen also enhances aerobic endurance[70)
tinct, regardless of differences in their pathways but not short-term exercise that makes use of the
and in their contributions to energy metabolism phosphagen and anaerobic glycolytic systemsPl)
during different kinds of exercise. Experimental Conversely, the aerobic system can be inhibited se-
evidence of the lack of coupling between power lectively by conditions that compromise oxygen
systems therefore seems to be essential if the sys- supply, such as hypobaric hypoxaemia.£72,73) Fi-
tems are to be regarded as distinct or independent.
nally, bicarbonate ingestion enhances performance
There would appear to be 2 kinds of evidence: ei-
of sprint and short-term endurance activities that
ther it must be possible to change the maximal
depend on anaerobic and aerobic glycolysis,£74-76)
power of 1 system without affecting the maximal
but feats of strength that rely only on power from
power of the other systems; or for exercise powered
the phosphagen system are unaffected. Bicarbon-
by 2 systems, it must be possible to change the
ate appears to work by providing a buffer against
relative proportions of energy provided by each
the fatigue-inducing hydrogen ions released during
system. We will cite first some well known exam-
anaerobic glycolysisP7J
ples of such selective modifications of the 3 tradi-
tional energy systems, then provide evidence of Evidence of the independence of the aerobic
similar effects for the putative aerobic glycolytic lipolytic and glycolytic systems comes from inves-
and lipolytic systems. tigations of the effects of both acute and chronic
Studies of the specificity of training provide 1 dietary manipulations, which result in changes in
approach for testing whether power systems are the proportions of carbohydrate and lipid utilised
distinct. In such studies, the main focus is usually during exercise that uses both substrates. When
the practical question of whether training bouts of plasma free-fatty-acid (FFA) levels are elevated
a specific duration and intensity affect perfor- before exercise, either by a fatty meal and heparin
mance in exercise tests or competitive events of a injection or by caffeine ingestion, there is an in-
different duration and intensity. Strength or power creased utilisation oflipid and concomitant sparing
training of an intensity sufficient to engage pre- of muscle glycogen during exercise, especially at the
dominantly the phosphagen system enhances max- high intensities typical of endurance eventspS-Sl)
imal performance of this system but not of the On the other hand, when FFA availability is re-
aerobic system.[S,17,5S.60) Similarly, prolonged en- duced by administration of nicotinic acid, muscle
durance workouts enhance aerobic performance, glycogen degradation during exercise is increased
but not strength.£61-65) Thus, these 2 systems are and exercise capacity is markedly reduced.£s2-S5)
distinct, at least with respect to training. Training Chronic adaptation to high-fat diets (>55%
studies have yet to provide unequivocal evidence of total energy), which results in a decline in pre-
of the independence of the anaerobic glycolytic exercise muscle glycogen content,[S6-S7) also re-
system, in part because activities that engage this sults in an increased utilisation of lipid during ex-

© Adis Intemational Limited, All rights reserved, Sports Med. 19 (4) 1995
Aerobic Glycolytic and Lipolytic Systems 245

ercise.l86-90] Conversely, adaptation to a diet high For athletes, the most important implication is
in carbohydrate (>65% of total energy) elevates that endurance and ultraendurance events require
muscle glycogen content[91,92] and increases the different dietary and training regimens. Research
utilisation of carbohydrate during exercise.l90,93-95] is urgently needed to better define such regimens.
The resulting effects on endurance performance In the meantime, athletes can apply the new para-
are largely a function of training status and preced- digm by invoking the specificity principle, which
ing period of dietary adaptation: untrained and states that training must stress the physiological
moderately trained individuals show enhanced systems critical for optimal performance in a given
endurance after high-carbohydrate diets,[91-93] event.[100] Thus, training for endurance events
whereas highly endurance-trained individuals should consist mainly of maximal steady-state
may, under certain conditions, benefit from a high- workouts or high intensity repetition training with
fat diet. [86-88,96] Regardless of the effects on perfor- short (<1 minute) rest intervals. Athletes should
mance, the fact that substrate utilisation during en- also be adapted to a high-carbohydrate diet, and
durance exercise can be modified substantially by any supplementation during and after an event
diet is confirmatory evidence of functional in- should be high in carbohydrate. Conversely, train-
dependence of aerobic glycolysis and lipolysis. ing for ultraendurance events should consist of
prolonged, continuous, moderate-intensity work-
outs; adaptation to a high-fat diet may be beneficial
2. Training and Dietary
for performance, and supplements consumed dur-
Recommendations for Athletes
ing the event should contain some easily digested
fats. [96,98] High-fat diets are associated with increased
Research over the past 25 years has emphasised
risk of various diseases, and although chronic endur-
the role of carbohydrates in endurance exercise,
ance training attenuates the risk,[101, 1021 athletes
and athletes generally appear to have accepted the
should nevertheless limit their exposure to dietary
advice that optimal performance will be obtained
fat. Optimal performance in ultraendurance events
from a high-carbohydrate diet combined with
may still be obtained if an athlete trains for most
carbohydrate supplementation during an event.
Only recently have sports scientists begun to of the year on a high-carbohydrate diet, then adapts
appreciate that lipids can make an important con- to a high-fat diet for 1 to 2 weeks prior to a major
event. [96]
tribution to performance, even in the shorter en-
durance events where they are not the main It is difficult to recommend training and dietary
fueI.l86-88,96- 991 For the longer endurance events, regimes for events that make substantial demands
lipids must play an even more important role. [37.38] on both the aerobic glycolytic and the aerobic lipo-
Our new paradigm reflects and should augment the lytic power systems. Clearly, both systems need to
developing interest in the role of lipid utilisation be trained, possibly by workouts that alternate be-
during prolonged exercise. However, by asserting tween moderate and high intensities in combina-
that there are 2 functionally separate aerobic tion with a mixed diet. However, a better strategy
power systems, we infer that there are 2 qualita- might emphasise maximal steady-state workouts
tively different kinds of prolonged, maximal event: combined with a period of adaptation to a moder-
those lasting less than about 4 hours, performed at ate- to high-fat diet during training,[86-88] followed
greater than 70% of "V02max and powered by aero- by a high-carbohydrate diet to increase glycogen
bic glycolysis; and the longer, lower-intensity levels during the final day or two before the event.
events powered by aerobic lipolysis. We will de- Results of a recent study support the latter recom-
fine these respectively as endurance and ultra- mendation.l96] The optimal energy supplementa-
endurance events. This concept has important im- tion strategy for such endurance events is also un-
plications for athletes and sports scientists. clear. For example, how does an athlete's diet

© Adls International Umited. All rights reserved. Sports Med. 19 (4) 1995
246 Hawley & Hopkins

immediately preceding the event affect fuel utilisa- onstration that prolonged, low- to moderate-inten-
tion during the event? Should supplementation be- sity training enhances lipid utilisation and improves
gin at the start of the race, or part way through? ultraendurance performance is now required.
What is the optimal mix of carbohydrate and fat? Our paradigm also predicts that endurance train-
What is the optimal type of fat? Sports scientists ing of an intensity sufficiently high to engage only
will need to answer these practical questions by aerobic glycolysis should increase the power of the
undertaking appropriately controlled experimen- aerobic glycolytic system while leaving the aerobic
tal trials. lipolytic system unaffected. At first sight, this pre-
diction may seem counterintuitive, because high-
3. Implications for Sports Scientists intensity endurance training will, inter alia, in-
crease the density of muscle mitochondria, which
3.1 Specificity of Training should then be available for increased oxidation of
lipid at moderate exercise intensities. However, the
The new paradigm raises several issues. The rate of lipid oxidation at these intensities may be
most important of these is the extent to which the limited not by the density of mitochondria, but
specificity principle operates in the training of the by the availability ofFFA to the mitochondria,1105]
aerobic power systems. To investigate this issue, it by the ~-oxidation process in the cytoplasm, or by
will be necessary to assay changes in the maximal the enzymatic capacity of mitochondria to use
power of the aerobic glycolytic and aerobic lipo- FFA,II06.I07] none of which may be increased sub-
lytic systems following training aimed specifically stantially by high-intensity endurance training.
at one or other of the systems. The maximal power The effects of endurance training on lipid util-
of the aerobic glycolytic system is relatively isation during exercise have already been ad-
straightforward to assay. It can be measured indi- dressed in several published studies.II08-HO] A
rectly as V0 2max , because at this intensity of exer- problem with some of these studiesl108.109] is that
cise very little fat will be oxidised.l40] It is also well the training consisted of a combination of high- and
represented by the maximal steady-state power or low-intensity workouts, which would have pro-
speed at the lactate threshold; such exercise is still vided training stimuli to both the aerobic glycolytic
of sufficient intensity to be powered predominantly and the lipolytic systems. A further problem is that
by carbohydrates. Assaying the maximal power of lipid utilisation was investigated only at the same
the aerobic lipolytic system will be a little more absolute intensity of submaximal exercise after
challenging: researchers will need to find the max- training. The observed increase in lipid utilisation
imum rate of lipid oxidation that occurs as exercise could, therefore, represent a change in the maximal
is gradually increased in intensity. The maximum power of the lipolytic system, or it could be a con-
rate could be derived simply from measurements sequence simply of the fact that the same absolute
of the respiratory exchange ratio, although tracer exercise intensity represents a lower relative inten-
techniques would also be helpful in estimating the sity following training, which favours utilisation
contributions of intramuscular and extramuscular of lipid.l40]
lipids.
Our paradigm predicts that long workouts (> 1 3.2 The Limits of Aerobic Lipolysis
hour) at low to moderate intensities (==50% of
V02max) should increase the maximal power of the Defining more precisely the factors that limit
aerobic lipolytic system but not of the aerobic the power of the aerobic lipolytic system is itself
glycolytic system. Studies showing little or no ef- an area for further research. Endurance training en-
fect of such training on V0 2max [103,104] are con- hances fat oxidation and spares intramuscular
sistent with this prediction, but the effect on lipid glycogen stores by producing changes in the mix
utilisation in these studies is unclear. A clear dem- of fuel utilised for submaximal exercise that uses

© Adls Intemational Umned. All rlgh1s reserved. Sports Med. 19 (4) 1995
Aerobic Glycolytic and Lipolytic Systems 247

both carbohydrate and fat. [2,111 ,112) A theoretical hydrate supplementation will also limit the break-
basis for this shift in substrate utilisation has been down of muscle protein during maximal ultra-
discussed elsewhere:[46,113) the shift is associated endurance exercise.[127) The optimal composition
with an increased capillary density in muscle,[1l4) and ingestion regimen for the supplement are fur-
enhancement of skeletal muscle oxidative enzyme ther topics for research.
concentration,[1l5) and an increased oxidation of
both intramuscular triglycerides[l09,llO,1l6) and 4. Conclusions
plasma FFA.l ll7 ,1l8)It remains to be determined to
what extent the maximal power of the aerobic lipo- We have defmed criteria for determining whether
systems that provide power for exercise are dis-
lytic system can be modified independently of that
tinct, and have presented evidence that the gly-
of the aerobic glycolytic system.
colytic and lipolytic components of the aerobic
The power of the aerobic lipolytic system could,
system satisfy such criteria. The evidence is per-
theoretically, be increased by the ingestion of eas-
suasive but not yet conclusive: studies of the spec-
ily absorbed lipids, such as medium-chain (C8-IO )
ificity of training and of the effects of different
triglycerides (MCTs). Because MCTs are more
dietary manipulations on exercise metabolism are
water soluble than long-chain (C I6-22) triglycer-
now required to confirm the existence of 4 func-
ides, they are absorbed more rapidly from the
tionally distinct systems rather than 3.
gut.l 1l9) They also diffuse into muscle rapidly be-
It is clear, nevertheless, that the body meets the
cause they do not require the transport mechanisms
metabolic demands of endurance and ultraendur-
that limit the rate of uptake of long-chain fatty
ance exercise in fundamentally different ways:
acids. They can be oxidised rapidly during long
carbohydrates are the optimal fuel for brief work
term exercise,[120-122) and may enhance endurance
bouts, but lipids can be the main fuel to sustain
by either sparing the intramuscular lipid and carbo- prolonged, submaximal exercise. The implications
hydrate stores,[90,96,99) or by augmenting the rate of for athletes are considerable, because the existence
utilisation of extramuscular lipid when the intra- of 2 aerobic power systems implies that different
muscular pool becomes depleted. training and dietary regimens are necessary to
Further research also needs to be undertaken on optimise performance in endurance and ultra-
the energy sources for maximal exercise lasting endurance events. By moving to 4 distinct power sys-
longer than 5 hours. The available evidence indi- tems, we will accord lipid the autonomy and attention
cates that lipids will make the main contribution to it deserves as the main fuel for sustaining the longest
energy requirements,[49,123,124) but there may be bouts of exercise that humans can endure.
obligatory utilisation of a certain amount of carbo-
hydrate to maintain citric acid cycle intermediates Acknowledgements
at a level needed to support the oxidative capacity
of muscle.l l25 ) For example, when the concen- The authors wish to thank Dr E. Vicki Lambert for
providing an unpublished manuscript during the preparation
tration of oxaloacetate in mitochondria is low,
of this article, and our colleagues for helpful suggestions.
entry of acetyl groups to form citrate is reduced, The insightful comments of two anonymous reviewers are
which in tum markedly impairs fatty acid utili sa- also acknowledged.
tion,l2,97, 125) Some carbohydrates will therefore
need to be consumed during the event, because References
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