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1 Research Institute for Sport and Exercise Sciences, Liverpool John Moores University, Liverpool L3 2ET; 2 University Department of Anaesthesia, University Hospitals of South Manchester, Withington Hospital, Manchester M20 2LR; and 3 Oxford Lipid Metabolism Group, Radcliffe Infirmary, Oxford OX2 6HE, United Kingdom
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ABSTRACT |
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The physiological and metabolic demands of hill walking have not been studied systematically in the field despite the potentially deleterious physiological consequences of activity sustained over an entire day. On separate occasions, 13 subjects completed a self-paced hill walk over 12 km, consisting of a range of gradients and terrain typical of a mountainous walk. During the hill walk, continuous measurements of rectal (Tre) and skin (Tsk) temperatures and of respiratory gas exchange were made to calculate the total energy expenditure. Blood samples, for the analysis of metabolites and hormones, were taken before breakfast and lunch and immediately after the hill walk. During the first 5 km of the walk (100- to 902-m elevation), Tre increased (36.9 ± 0.2 to 38.5 ± 0.4°C) with a subsequent decrease in mean Tsk from this time point. Tre decreased by ~1.0°C during a 30-min stop for lunch, and it continued to decrease a further 0.5°C after walking recommenced. The total energy intake from both breakfast and lunch [5.6 ± 0.7 (SE) MJ] was lower than the energy expended [14.5 ± 0.5 (SE) MJ; P < 0.001] during the 12-km hill walk. Despite the difference in energy intake and expenditure, blood glucose concentration was maintained. The major source of energy was an enhanced fat oxidation, probably from adipose tissue lipolysis reflected in high plasma nonesterified fatty acid concentrations. The major observations were the varying thermoregulatory responses and the negative energy balance incurred during the hill walk. It is concluded that recreational hill walking can constitute a significant metabolic and thermoregulatory strain on participants.
energy balance; thermal regulation; field study
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INTRODUCTION |
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THE PROLONGED DURATION OF a typical hill walk places exceptional demands on the participants. The specific demands of hill walking incorporate activity of varying intensity and duration, both of which are influenced by factors such as the physical fitness of the participant, dietary intake, backpack weight, environmental weather conditions, and severity of the terrain. Besides, hill walkers can be caught unexpectedly and unprepared when rain and wind accompany outdoor activities in cool weather. Decreased thermal insulation of wet clothing presents a serious challenge to body temperature regulation, which can be compounded by fatigue associated with prolonged exercise such as hill walking (33, 36, 38, 42).
The problem of wet-cold hypothermia is recognized by search-and-rescue organizations worldwide. Nevertheless, their ability to design educational material concerning this hazard is hindered by lack of knowledge of the physiological and psychological responses (42). The information that is available derives from the pioneering work of Pugh (36-38), supplemented by anecdotal descriptions of exposure incidents (23, 25, 36, 41).
Pugh (37) proposed that maintaining an oxygen uptake
(
O2) of 2-2.5 l/min or
50-60% maximum oxygen uptake
(
O2 max) would offset heat loss
and combat the debilitating effects of the cold, wet, and windy
environment. Although these observations were based on only three
subjects, recent work by Weller et al. (45) supported
Pugh's postulate. Both experiments were based in an environmental
chamber, and subjects exercised on a cycle ergometer and treadmill,
respectively. Pugh (37) and Weller et al.
(45) showed that, when exercise metabolism is reduced, the
increase in shivering may be insufficient to prevent a decrease in deep
body temperature. Weller et al. (45) reported that rectal temperature (Tre) and the metabolic responses to an initial
120-min phase of exercise at ~60% of peak
O2
(
O2 peak) were not influenced by the
cold stress of a wet and windy environment. However, during a
subsequent 240-min phase of exercise at 30%
O2 peak, Tre was lowered by
0.6°C, whereas the following were elevated:
O2 (25%), the proportion of
carbohydrate (CHO) oxidized, and the venous concentrations of lactate,
glucose, norepinephrine, and epinephrine.
The studies reviewed have been limited to simulated conditions.
Consequently the influence of a "typical" day's hill walk on a
range of physiological and metabolic variables, in the field, has not
been established. Given such limitations, the aim of the present study
was to investigate selected responses to a typical hill walking event
to gauge the overall physiological and metabolic strain. The field
conditions would likely impose additional stresses not encountered
during simulated conditions, such as stoppage of activity for fluid and
food intake (transiently altering the balance between heat production
and heat dissipation), in conjunction with varying terrain and weather
conditions. Furthermore, we aimed to quantify both the energy cost of
such activities and relevant responses that are important in the safety
of hill walkers, such as the potential thermal stress, impaired
psychomotor performance, and the ability to maintain glycemia. This
type of study may be important in adding to the mostly anecdotal
information regarding exposure and recreational activities. Because of
the continuously changing intensity of activity, it is unlikely that
individuals will be able to operate at or above 50-60%
O2 max, the
O2 "cutoff" point described by Pugh
(37) for combating heat loss during a hill walk. Although
Pugh's postulate may have an element of truth, it would depend on
factors such as favorable ambient temperature, the clothing worn,
terrain, and physiological capabilities of the participants. The first
hypothesis, therefore, is that the
O2
cutoff is not a realistic component of hill walking. Second, it was
hypothesized that a seemingly "normal" hill walk in possible
adverse, but not uncommon, conditions leads to a significant physiological, psychomotor, and metabolic stress on the body.
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METHODS |
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Subjects
Thirteen subjects (11 men and 2 women) participated in this study, which was reviewed and approved by the Human Ethics Committee of Liverpool John Moores University. The subjects gave written consent to participate in the study after they had been fully informed of the nature, purpose, and possible risks associated with the study. The physical characteristics of the subjects are shown in Table 1. The majority of the subjects were active and experienced hill walkers. Experiments were conducted from January through March. Body density and percentage of body fat (%fat) were estimated from skinfold thicknesses over the biceps, triceps, and subscapular and suprailiac areas (14). Fitness level was established by using a continuous incremental treadmill running test to exhaustion (3). A plateau in the
O2-to-work relationship was reached in
only four subjects; therefore, the highest aerobic power was expressed
as
O2 peak and not as
O2 max.
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Protocol and Procedures
On separate occasions, subjects completed a 12-km (8 mile) hill walk. The course varied in elevation from 100 to 902 m above sea level and consisted of a range of gradients and terrain typical of a mountainous hill walk. A caravan was used as a temporary field laboratory and for living accommodation, and was located at the start and end of the hill walk. Subjects woke each morning between 0500 and 0530 and completed the preliminary experiments before the hill walk (Fig. 1). Self-paced walking began each day between 0700 and 0800. Before the walk and on its completion, subjects weighed themselves nude. Subjects were permitted fluid and food ad libitum. They selected their own food and fluids for the walk, which were preweighed before the walk. The energy gained from CHO, fat, and protein was subsequently determined by using standardized food tables (30). After initial weighing, the participants inserted a rectal temperature probe to a depth of 10 cm beyond the anal sphincter. Skin temperature (Tsk) was assessed by the placement of temperature thermistors on the chest, forearm, thigh, and shin. Thermistors and the rectal probe were connected to a data logger (Squirrel meter 1000, Grant Instruments, Cambridge, UK) that recorded data every 6 min. On the walk, a rest period of ~1-3 min was allowed every time thermal measurements were made, and 30 min were allowed for lunch (Fig. 1). During the hill walk, respiratory gas-exchange measures were obtained with a portable telemetry system (Metamax, Cortex Biophsik, Borsdorf, Germany). All subjects carried a lightweight waterproof backpack that contained the Metamax system and thermal logger to allow continuous recording of respiratory gas exchange and Tre and Tsk, respectively. The loaded pack weighed 9.5 kg, which is consistent with a hill-walking scenario.
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Measurements and Analysis
Temperature and heart rate.
Tre and Tsk were monitored continuously and
recorded every 6-min with the data logging system. Heart rate (HR) was
recorded by means of short-range radiotelemetry (PE3000 Sports Tester, Polar Electro, Kempele, Finland) every minute and were subsequently averaged over 5-min blocks. Mean Tsk
(



x) · 

Environmental measurements.
Environmental air, dry, and wet bulb temperatures and velocity were
recorded with a digital sling psychrometer thermohygrometer and a
kestrel vane anemometer, respectively. Wind chill index was calculated
from the air temperature and velocity by the equation of Nishi and
Gagge (32): Ko = (33
Ta )(10V0.5
V + 10.45), where Ko is the cooling power
of the environment (kcal · m
2 · h
1),
Ta is the ambient temperature (°C), and V
represents the air velocity (m/s).
Thermal balance.
All thermal balance data (rates) are expressed in units of watts per
square meter. The rate of heat gained or lost (heat debt) from the body
mass (±S) was computed from the equation of Burton (9): ±S = 

1,
where 


1 · °C
1), and SA is
the body surface area (m2). The thermal balance was
calculated and averaged every 2 km of the walk.
Indirect calorimetry.
Continuous assessment of respiratory gas exchange was performed by the
use of a portable telemetry system. Signals from the Metamax system
were logged and subsequently retrieved at the end of the hill walk.
This procedure allowed for continuous monitoring of
O2, ventilation, and respiratory
exchange ratio. The percent contributions of the CHO and fat oxidation
were estimated from nonprotein (NP)
O2
and respiratory exchange ratio (RER) data with the use of the following
formulas: %CHO = [(RERNP
0.707)/(1
0.707)] and %fat = 100
%CHO. It was assumed that
protein oxidation contributed 12.5% of energy expenditure at rest and
that exercise did not alter this relative rate of protein utilization
(45). The respiratory quotients of CHO, fat, and protein
were taken as 1.00, 0.707, and 0.81, respectively. Oxidation rates
(g/min) were estimated for CHO, fat, and protein, assuming 0.829, 2.019, and 0.966 liters of oxygen were consumed per gram of substrate oxidized (28), respectively. During the hill walk, CHO and
fat oxidization rates were calculated and averaged over 10-min blocks at predetermined points in accordance with the other measurements. Total oxidation rates were then averaged for the whole hill walk. Energy expenditure was calculated from the averaged
O2 and CO2 production from
the whole walk by using the formulas of Elia and Livesey
(15). Before use, the Metamax system was calibrated using
both calibrated gas and ambient air. The volume transducer was
calibrated using a 3-liter syringe. To decrease any error, the system
was recalibrated during the hill walk when the subjects stopped for
lunch. Because of technical problems during three of the walks, the
respiratory gas exchange data were based on 10 subjects.
Psychomotor measurements. The Profile of Mood State (POMS) was measured using 65 ratings each on a 5-point rating scale. The scales are factored into six mood scores: depression/dejection, tension/anger, anger/hostility, confusion/bewilderment, fatigue/inertia, and vigor/activity (31). In addition, the subjects were asked to rate their overall ratings of perceived exertion (RPE) on a 6-20 scale (8) from the start of the walk until the lunch stop, and from the lunch stop until the completion of the walk. Reaction time (cognitive function) tests (Hick's law) (1-, 2-, 4-, and 8-choice reaction time for a finger response) were assessed on a laptop computer before, during, and after the walk. Subjects were fully familiarized with the use of the equipment. Grip strength (motor function) was assessed by means of a handgrip dynamometer (Takei, Narragansett, Japan).
Blood and urine sampling and analysis. Blood samples were obtained from subjects in a semireclined position before, during, and immediately on completion of the walk as illustrated in Fig. 1. The venous blood samples (5 ml) were drawn from a superficial forearm vein with minimum stasis, then immediately placed in a vacuum flask containing ice. From the blood samples, serum was separated rapidly at ~4°C and frozen for later determination of serum nonesterified fatty acids (NEFA) and triacylglycerol (TAG) concentrations by enzymatic methods. In addition, a portion of the whole blood was immediately deproteinized with perchloric acid (7% wt/vol) in preparation for whole blood glycerol, lactate, 3-hydroxybutyrate (3-OHB), and glucose determination by enzymatic methods. All enzymatic methods were adapted to a Monarch centrifugal analyzer (Instrumentation Laboratory, Warrington, UK). Serum insulin concentrations were determined with a double-antibody radioimmunoassay (Pharmacia and Upjohn, Milton Keynes, UK). Serum cortisol concentrations were determined by using a solid-phase radioimmunoassay (Diagnostic Products, Llanberis, Wales, UK). Some of the uncoagulated blood was also used for the measurement of hemoglobin and packed cell volume (conventional microhematocrit method). Plasma volume changes were calculated from changes in hemoglobin and packed cell volume relative to initial resting values as described by Dill and Costill (12).
Urine was collected during the rest day before the hill walk, in which subjects performed no exercise, and during the hill walk at the following times: 0800-13:00, 1301-1800, and 1801-2000. From these collections, a 5-ml mixed sample was removed. Urine epinephrine, norepinephrine, creatinine, and dopamine concentrations were then analyzed by using high-performance liquid chromatography with electrochemical detection (in-house method, Dept. of Clinical Chemistry, The Royal University Hospital, Liverpool, UK). Index of dehydration was determined in triplicate using urine osmolality determined by freezing-point depression (model 3300 Advanced Microosmometer, Vitech Scientific, West Sussex, UK). For the urine osmolality, a 5-ml sample was produced after the first void of the day, and then from the first sample after the walk.Statistical analysis.
Variables are presented as means ± SE. Data were initially tested
for normality, before being analyzed by repeated-measures ANOVA. The
ANOVA results were corrected by the Huynh-Feldt
-adjusted degrees of
freedom when the violation to sphericity was minimal (>0.75), and the
Greenhouse-Geisser correction was used when sphericity was violated
(<0.75) and significant condition and condition-time interactions were
identified (16). Post hoc tests (honestly significantly
different) were performed to isolate any significant differences.
Student's paired t-tests ascertained between-condition differences when a variable was measured once. Statistical significance was set at P
0.05 for all statistical tests.
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RESULTS |
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Exercise Duration
All subjects completed the 12-km hill walk. The mean (range) duration for the hill walk was 348 (245) min. The differences in the time to complete the walk were due mainly to variations in weather conditions and terrain. Both cold, wet, and windy weather and deep snow underfoot led to an increased time to complete the hill walk.Energy Balance
Energy intake and energy expenditure during the hill walk are given in Table 2. Total energy intake was lower than total energy expended during the hill walk in all subjects. CHO, fat, and protein comprised 65, 25, and 10%, respectively of all food consumed, in contrast to 47, 42, and 11%, respectively, of fuels oxidized. The energy intake from both CHO and fat was lower than the amount oxidized (P < 0.001), leading to the lower energy intake relative to expenditure (P < 0.001). The relatively high energy expenditure of 14.5 ± 0.5 MJ reflects the high energetic cost of hill walking, even when pursued over a relatively short duration. The negative energy balance is also shown by a decrease (72 ± 2 to 70 ± 2 kg; P <0.05) in nude body mass as a consequence of the walk.
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Thermoregulatory and Environmental Data
The mean (range) of the recorded environmental data for air velocity, air temperature, and wind chill index was 2.8 (0.1-10.4) m/s, 6.4 (
1.3-13.2)°C, and 520 (176-1,239)
kcal · m ·
2 · h
1,
respectively. These figures highlight the variability in the weather
conditions over the period of testing. Five of the walks were completed
in cold, wet, and windy weather. The surface conditions on the walks
tended to vary with the weather. Snow and ice were regularly
encountered, along with high winds, as reflected by a high windchill
index; these factors represent walking in very demanding climatic conditions.
The rise in Tre, illustrated in Fig.
2C, is a typical response
during both exercise and the initial stages of cooling because of
peripheral vasoconstriction decreasing the return of cooled blood from
the periphery. Also reflected at this point were the apparent decreases
in 



356 ± 23 W · m2) was observed during the lunch stop and for
the subsequent 2 km after this, before gaining a positive balance
again.
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During the ~30-min stop at midwalk, three subjects showed a pronounced shivering response after ~15 min of rest. Tre fell ~1.0°C from the level observed during the prelunch walking. Before the lunch stop, two of these subjects showed early signs of exposure; symptoms included stumbling, withdrawal from voluntary conversation, and slowing down in pace. Decreases in thermal balance were also observed at this time. The majority of the subjects complained of feeling "very cold" and wanting "to speed up" during the period after the lunch stop: these subjective impressions usually lasted ~15-45 min, depending on the weather conditions.
Psychomotor performance. The POMS profile showed an expected increase in tension and confusion before the walk (P < 0.05) and an increase in fatigue (P < 0.05) postwalk relative to both before and at midwalk. Overall RPE from the start of the walk to the lunch stop and to the final part of the walk was 15 ± 2 and 13 ± 3, respectively. There was a small decrease in grip strength (45.4 ± 2.7 to 43.5 ± 2.8 kg/m2) from prewalk to postwalk (P < 0.01). Any changes in reaction time were less evident. The only significant changes in reaction time were evident in "one-finger" reaction time (P <0.01) and in the recorded errors (4 finger) (P < 0.05, not shown), both after completion of the walk. However, the normal circadian variation of an accelerated reaction time from morning to afternoon was clearly not evident (not shown).
Respiratory gas exchange and HR responses.
During the first ~5 km of ascent, RER increased from 0.82 ± 0.03 at base to 0.89 ± 0.02. During the descent, RER gradually fell to <0.84 ± 0.02 for the final 5 km of the walk (Fig.
4A). RER changes are reflected
in the oxidation rates of CHO and fat shown in Fig. 4B.
During ascent, both CHO and fat oxidation increased. After the first 4 km, CHO oxidation decreased for the duration of the walk, with fat
oxidation remaining elevated (Fig. 5).
Increases in the
O2 and HR were evident
during the rise in altitude over the first 4.5 km of the hill walk
(Fig. 5, D and C). From ~3 km until the descent
of the hill walk (~6 km), subjects were operating at ~50%
O2 peak with an average HR of 148 ± 8 beats/min; during the descent, this value fell to ~25-40% of
O2 peak with HR averaging 126 ± 5 beats/min (Fig. 5, D and C).
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Blood and urine constituents
There were no significant changes in plasma volume during the walk. Consequently, circulating concentrations have not been corrected for hemoconcentration. Table 3 gives concentrations of the blood constituents. The energy metabolites 3-OHB, lactate, glycerol, and NEFA increased from prewalk to midwalk (P < 0.001). In contrast, there was no change in TAG concentrations. Insulin increased significantly, whereas cortisol decreased significantly (P < 0.01) postwalk, relative to both prewalk and midwalk. Table 4 gives the concentrations of the urine catecholamine collections. Generally, the hill walk led to a marked elevation in urinary epinephrine and norepinephrine compared with the rest day. In addition, urine osmolality increased prewalk to postwalk (from 603 ± 86 to 744 ± 71 mosmol/kgH2O; P < 0.05).
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DISCUSSION |
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The main finding of this study was that a seemingly normal outdoor hill walk in adverse, but not uncommon, conditions led to a significant physiological stress on the body. Despite these stresses, which included dehydration, high thermal stress, and marked negative energy balance, subjects demonstrated only slight impairment in some of the measured psychomotor tests throughout the walk. Furthermore, the hill walk significantly altered the hormonal and metabolic milieu. Despite the large difference in energy intake and expenditure, a normal blood glucose level was maintained. The major source of energy, in the monitored walk, was an enhanced fat oxidation probably from adipose tissue lipolysis.
Physiological Responses
Pugh (37) described a
O2 cutoff point, above which individuals
exercising in a cold, wet, and windy environment would not experience
any influence on the physiological responses to exercise, i.e., drop in
core temperature, mental impairment, extreme fatigue, and exhaustion.
Below this point, there would be an obligatory increase in energy
expenditure and subnormal Tre and muscle temperatures. Weller et al. (45) suggested that the cutoff point is
likely to depend on factors, such as clothing insulation, body
morphology, mass, and body fatness (43), and may account
for the random nature of the hypothermic casualties described by Pugh
(35, 37). The present observations highlight the
variability in
O2 in response to the
hill walk, which is likely to depend on such factors as terrain,
gradient, weather condition, backpack weight, exercise intensity,
preceding diet, and thermal stress. It was only during the
high-intensity part of the walk that subjects reached this cutoff
point. This cutoff point was clearly variable. Because the hill walkers
in this study walked at their own pace, it could be cautiously
concluded that hill walkers do not consistently operate at, or above,
this cutoff level.
Weller et al. (45, 46) demonstrated heat loss was greater
in low-intensity (~30%
O2 peak) than
high-intensity (~60%
O2 peak)
walking, illustrating how the rate of heat production during
high-intensity exercise will offset heat loss to the environment more
effectively than low-intensity exercise. This observation has important
implications for hill walking in that, during the low-intensity phase
of a hill walk (e.g., walking downhill, navigation, and so on), an
increased heat loss relative to heat production may be experienced. The
factors already mentioned, along with individual variation in effective
cold stress under given environmental conditions, may lead to a
compromise in the ability to operate safely in the mountainous environment.
The observations of the present study regarding the thermal stresses involved in a hill walking event have provided some novel results. The results show generally higher values for the Tre profile than described in previous studies (42, 45, 46); this increase is most likely due to the elevated thermal insulation from the protective clothes worn by the hill walkers, causing a decrease in heat loss, which will subsequently increase Tre. The maintenance of normal core temperature during cold stress depends on the subject's ability to generate enough heat to offset heat loss to the environment (33, 46). This clearly was not a problem for the hill walkers because of high levels of prolonged exercise intensity.
When the subjects stopped for lunch and measurements midwalk for ~30
min, the exercise hyperthermia was canceled out by the decreased heat
production and increased heat loss through conduction and radiation.
The initial physiological responses to cold exposure to maintain core
temperature in the cold are peripheral vasoconstriction to reduce heat
loss and shivering to generate heat. Although shivering was not
quantified directly, pronounced shivering was noted in four of the
subjects. Once peripheral vasoconstriction is maximized, core
temperature can only be maintained by an increased heat production, i.e., shivering, which is thought to be the major contributor to the
cold-induced increase in heat production (13). The core temperature continued to fall after subjects began walking after lunch.
This temperature afterdrop has been reported in a number of
cold-water-immersion studies (17, 33) but, to our
knowledge, has not been reported in circumstances such as those of the
present study. Even though Tre did not drop below 36°C,
this afterdrop may describe the reason for hill walkers slipping into
the first stages of hypothermia after stopping for a rest. The
suggested mechanisms for the afterdrop are at present subject to
controversy (10, 17, 27, 44). The extent of this afterdrop
was also reflected by the high negative thermal balance (
356 ± 23 W · m2). Weller et al. (45)
reported similar values for thermal balance after 240-360 min of
walking in a simulated cold, wet, and windy environment with subjects
wearing a minimum of clothing. The high negative thermal balance
highlights the thermal stress placed on the human body, even when
apparently prepared for the prevailing weather conditions
One possible explanation for this continued drop in Tre may be the decrease in exercise intensity (and hence metabolic heat production) after lunch when the subjects were walking downhill. Studies of cold-water immersion have shown Tre to continue to fall after immediate removal from cold water, even when subjects are actively rewarmed (27). The extent to which this afterdrop can be limited by an increase in exercise intensity in this scenario merits further research.
Psychomotor Response
The unremarkable changes in the psychomotor tests demonstrate that, despite serious physiological stress, the subjects demonstrated normal motor control during the walk. The small decrease in grip strength suggests that, over the monitored walk, motor function was near normal, despite potential cooling of the peripheral tissues.Energy Balance
The recorded energy expenditure of 14.5 MJ highlights the high energetic cost of the 12-km hill walk and is comparable to the 12-MJ energy expenditure in the studies of Greenhaff et al. (18) and Maughan et al. (29). These latter studies were based on a flat 37-km walk, which corresponded to operating at 17% of
O2 max, with unrestricted access to
either a mixed or CHO diet, respectively. In contrast, subjects in the
present study operated at ~50% of
O2 max during the first 5-km on the predominately uphill section and at ~30% during the final 7-km, which was on the downhill section of the walk. This suggests that the
main determinant of energy expenditure, in a hill walk is the relative
difficulty of the walk, in terms of the gradient and terrain, both of
which will contribute to a greater level of exercise intensity.
Additionally, Maughan et al. demonstrated that a weight loss of 2 kg
over their 4-day walk was only apparent in the group who had a low-CHO
intake but was not observed in the group ingesting a high-CHO diet.
They concluded that the loss of body weight was a consequence of the
gradual use of the hepatic and muscle glycogen stores and loss of
associated water. In the present study over 1 day, the weight loss also
averaged 2 kg, with subjects consuming a mixed diet. The normal whole
body muscle glycogen pool amounts to some 400 g (21).
Water is stored in association with this glycogen in a ratio of
~3-4 g water/g glycogen (5, 34), suggesting that
the observed body weight loss of 2 kg in the present study may reflect
a small decrease in whole body glycogen level. Supporting this
postulated decrease in whole body glycogen was an observed negative CHO
balance of 200 g. In addition, this negative CHO balance of
200 g would equate to an ~800-g loss of body water, suggesting a
high element of dehydration, which was also reflected by the increased
urine osmolality concentrations. The fundamental difference in the
relative intensities of the walk, compared with that of Maughan et
al., may largely explain the magnitude of the weight loss in the
present study. However, subjects in the studies described (18,
19) had unrestricted access to food, whereas, in the present
study, they supplied their own food that they considered appropriate
for the conditions. The unrestricted food may have attenuated any
potential negative energy balance in the aforementioned studies. In the
present study, the negative energy balance, the failure to provide
enough fuel for the exercise duration and intensity, could well have
led to compromises in both physiological and psychological functioning if the duration were more prolonged. This suggestion generates important considerations for the hill walker with regard to nutrient intake both before and during the walk. For example, because the subjects were clearly in negative energy balance, a suggestion is that
they should take more food and/or foods with higher energy content to
help prevent this from occurring and provide a measure of protection if
the walk becomes unexpectedly prolonged.
Metabolic Responses
The measurements made at the midpoint of the walk showed an enhanced lipolysis, demonstrated by an almost fourfold increase in NEFA concentrations accompanied by high glycerol and 3-OHB concentrations. Fatty acids delivered from adipose tissue are the predominant fuel for sustained exercise at moderate intensity (2, 11, 24). There is usually a surge in plasma NEFA concentrations shortly after exercise, presumed to reflect a continued high rate of lipolysis when muscle NEFA uptake has suddenly diminished (22), and this may have been responsible for some of the elevation in NEFA concentration observed. The stimulus for lipolysis during exercise is mainly adrenergic (4), reinforced by decreased insulin concentrations. It is likely that the former stimulus was greater in our subjects than in many exercise studies because of the adverse conditions. Even though large variations were present in urine catecholamine concentrations, the results clearly indicate a general stress response to the hill walk compared with the prior day.In contrast, NEFA, glycerol, and 3-OHB concentrations were considerably lower at the end of the walk than midway. The food intake midwalk is likely to have influenced the pattern of lipid mobilization during the final part of the walk. Ahlborg and Felig (1) and Krzentowski et al. (26) showed that CHO feeding before or during mild-intensity, prolonged exercise decreased the amount of energy derived from fat oxidation and increased proportionally the amount of energy-derived blood glucose. However, in the present study, the decreased NEFA, glycerol, and 3-OHB concentrations postwalk coincided with no evident decreases in fat oxidation assessed by indirect calorimetry. This maintained fat oxidation might be accounted for by fatty acids derived from the body's intramuscular stores (40). The relatively high circulating plasma insulin levels recorded at the end of the walk (~12 mU/l) would be expected to lead to a decrease in adipose tissue lipolysis, whereas intramuscular lipoysis is not so readily inhibited by insulin (7, 19).
Hypoglycemia, which would affect both fatigue and the shivering response (20), was not observed in this study at any time. The data indicate that the liver was able to meet glucose requirements by a combination of glycogenolysis and gluconeogenesis, supplemented by the midwalk carbohydrate intake.
In summary, the pattern of substrate mobilization and utilization is likely to vary according to the intensity and duration of the walk, level of fitness, environmental conditions and preceding diet of the participant. It is apparent that the energy expenditure during the hill walk exceeded energy intake, apparent in both total CHO and fat oxidation rates for the walk. The large energy expenditure observed served to highlight the high energetic cost of such hill-walking events, even when completed over a relatively short duration. The observations generate important implications for hill walkers with regard to nutritional strategies for preventing some of the potential detrimental effects of operating at a marked negative energy balance. The negative energy balance may lead to a compromise in physiological function and safety if activity is performed over a prolonged period. Nevertheless, despite the physiological stress and the difference between energy intake and expenditure, blood glucose was maintained. The major source of energy, in the monitored hill walk, was an enhanced fat oxidation probably from adipose tissue lipolysis.
It is evident that more research on this topic is desirable. Future research should extend the findings of this study into hill-walking events over a longer duration. Furthermore, there is little in the literature with respect to repetitive high-intensity hill walks completed over 1-2 wk. Finally, the optimal fluid and nutritional strategies need to be quantified for activity of this form.
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ACKNOWLEDGEMENTS |
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The subjects in this study deserve our special thanks. We admire their bravery to volunteer and the enthusiasm and persistence they maintained, despite the arduous testing and climatic conditions. We acknowledge the help of Dr. N. Roberts for the analysis of the urine catecholamines.
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FOOTNOTES |
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The study was supported by Mars Incorporated.
Address for reprint requests and other correspondence: P. N. Ainslie, Research Institute for Sport and Exercise Sciences, Liverpool John Moores Univ., Liverpool L3 2ET, UK (E-mail: humpains{at}livjm.ac.uk).
The costs of publication of this article were defrayed in part by the payment of page charges. The article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.
Received 15 June 2001; accepted in final form 29 August 2001.
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