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Protein turnover rates of two human subjects during an unassisted crossing of Antarctica
BY M. A. STROUD A N D A. A. JACKSON Institute o f Human Nutrition, West Wing Level C, Southampton General Hospital, Tremona Road, Southampton SO9 4XY AND J. C. WATERLOW London School of Hygiene and Tropical Medicine, Keppel Street, London W C l E 7HT
(Received 31 October 1995 -Revised 12 April 1996- Accepted 29 April 1996)
During the Austral summer of 1992-3, two men, MS and RF, waked 2300 km across Antarctica in 96d, unassisted by other men, animals or machines. During the journey they ate freeze-dried rations, towed on sledges, that contained an average of 21.3 MJ/d of which 567 % was fat, 35.5 % carbohydrate and 7.8 % protein (98.8 8). Despite this high energy intake both men lost more than 20 kg in body weight due to their extremely high energy expenditures. Studies of protein turnover using [15N]glycineby the singledose end-product method were made before, during and after the journey, and these demonstrated considerable differences in the metabolic responses of the two men to the combined stresses of exercise, cold and undernutrition. However, both men maintained high and relatively stable levels of protein synthesis during the expedition despite the great exertion and the onset of considerable debilitation. This stability indicates the vital physiological function of protein synthesis. Protein turnover : Exercise : Undernutrition
During the Austral summer of 1992-3 two men made the first crossing of Antarctica, unassisted by other men, animals or machines. They travelled on foot, hauling sledges for 1&12 h every day. Each sledge had a starting weight of 222 kg and contained 100 days’ supplies of food and fuel. The journey, which has been described by Stroud (1993), started at the Weddell Sea and ended after 96 d at the Ross ice-shelf. Overall, they covered about 2300 km with a climb of 3500 m to the highest part of the Antarctic polar plateau. During the course of the journey the men consumed an average of 21.3 MJ/d which, in order to save weight, was derived mainly from fat. Based on measurements of energy expenditure made on an expedition of similar nature (Stroud, 1987) it was expected that loss of body weight would be limited to 10 kg. In the event this was an underestimate and weight losses exceeded 20 kg. At the end of the journey the two men were on half-rations. In the course of the expedition a number of scientific studies were made including measurements of whole-body protein turnover, since little is known about the response of this key function to very severe exertion continued over a long time, especially when combined with negative energy balance. Protein turnover was measured 1 week before the expedition; five times during the expedition on days 9/10,19/20,49/50,76/77, and 92/96 (the two subjects being tested on different days); and twice following the journey, 2 and 3 weeks after return. The present paper reports the results of these measurements.
Urine volumes were measured and a sample retained for later analysis. Protein turnover Protein turnover was measured with [15N]glycineby the single-dose end-product method (Fern et al. Energy intake Before and after the expedition normal foods were consumed ad libitum. Stamford.166 M. NH. 1996). Herts.00 hours to 07. was eaten at 07. Welwyn Garden City. Urinary creatinine was determined by the Jaffrereaction and urinary NH. mean daily energy intakes were calculated making an allowance for any unused food. biscuits. East Midlands) before and after the expedition. were fit men aged 37 and 48 years respectively. These semi-quantitative records were analysed independently by two dietitians. and urea-N were removed stepwise from urine .39 kJ/g and that of fat was taken as 39. and urea were measured by the Berthelot reaction. The energy value of protein was taken as 18. chocolate bars.00 hours. Lean-tissue losses were assumed to be 73 % water and 27 O h protein. During the expedition the subjects ate a specially formulated pre-packed diet (Be-Well Nutrition. S T R O U D A N D OTHERS METHODS Subjects The subjects.00 hours the next morning.7 kJ/g (Brouwer. 99 atom per cent 15N) dissolved in water.. These isotopes were given to the subjects in order to measure energy expenditure using the doubly-labelled water method. Both men gave informed written consent for all experiments. 1965). On each occasion the measurements were made after overnight fast. Measurements of body composition were also made 10 d before and 7 d after the expedition using underwater weighing (UWW) with measurements of residual lung volumes by He-dilution spirometer (Gould Pulmonet 111). West Bromwich.) containing a high proportion of fat in order to provide a high energy : weight ratio. Raviv-Aran.00 hours. Body masses were recorded in the UK (Salter KR120 scales. Israel). defaecation and the voiding of urine. Breakfast. 1981). Lincs. whereas fat losses were considered to be 100% fat. The amount of each food was estimated and recorded. soups and hot chocolate drinks. Urine was collected in containers with thymol crystals as a preservative until 19.30 hours. Body composition estimates were calculated on days 0 and 50 of the expedition from the dilution space of the stable isotope l*O and on day 95 of the expedition from the dilution space of 'H. and the main meal was taken at 21. Energy expenditure Mean daily energy expenditure was estimated for the first 50 d and the last 45 d of the expedition by combining mean dietary energy intakes with the mean daily energy deficits calculated from losses of lean tissues and fat over each period of the expedition. and were also recorded at 10 d intervals during the journey with a miniature load-cell portable scale (Miniscale. MS and RF. but not weighed. A second urine collection was made from 19. All food was weighed and packaged into daily ration bags at the beginning of the expedition and.00 hours and the subject then took 100 mg [15N]glycine(Promochem. The bladder was emptied at 07. A. at the end of the expedition. a study that is to be reported elsewhere (Stroud et al. consisting of hot cereal and butter. Analysis was performed before departure with food values taken from manufacturers' nutritional information about their products. This consisted of freeze-dried meals supplemented with butter. soup and chocolate bars were eaten at intervals throughout the day.
the isotope excreted in urea.5.7 % came from fat. The value of e.E/e. is the I5Nexcreted in 24 h. collections were measured in a triple collector mass spectrometer (SIRA 10. synthesis rates were calculated only from Q. Since this could not be measured directly. Since blood samples were not taken. when only one meal was eaten.).): E. For the flux based on urea (Q. where d is the dose of 15N(mg) (Waterlow et al. Because the measurement of Q. was collected by further alkaline aeration. + N loss in creatinine is taken as a proxy.Eu(12)/e. Hence Q. The . and 425 g carbohydrate. for which E. VG Isogas.8 at 37' for 3 h in EDTA buffer. (Q. In each test 15Nabundances were calculated after subtracting the background abundance in the urine sample collected before the dose of labelled glycine. where I is the intake and E is the total urinary N excretion.1. Middlewich.PROTEIN TURNOVER D U R I N G A N T A R C T I C CROSSING 167 samples for mass spectrometry. The 15Nabundances in the NH. Over the second part of the expedition the average daily intake was 222 MJ consisting of 103 g protein. when the bulk of the food was consumed.) after liberation of N. is more open to error.-free urine estimated to contain 2 mg urea-N was then treated with urease (EC 3. 35. and were determined only for the 12 h periods following the dose since it could not be assumed that rates of protein synthesis and breakdown remained the same during the night.(12) = urea N excreted in 12 h (g). + E.(12) = d. needed to be corrected for the isotope retained as urea in the body at 12 h. 293 g fat. For the flux based on NH. Over the first 50 d the average daily intake was 19. Rates of protein synthesis (S) and breakdown (B) were obtained from the usual equation: Q = S + E = B+I.): E.S0. Therefore the appropriate value for e. it was assumed that there was no significant change in the size of the body urea pool. as during the day. 1995). In the later stages they also developed frostbite of the hands and feet which became chronically infected. in 12 h (mg).5) at pH 7. This was then aerated into 15 ml of 0*005~-(NH. both men lost considerable weight and became significantly debilitated by starvation towards the end of the walk.9 MJ consisting of 95 g protein. by LiOH.. The amount of urine estimated to contain 2 mg NH.( 12) = NH. before the resulting NH. Despite the high energy intake. The volume of NH.(12) = I5N excreted in NH.(24). RESULTS The dietary intake provided an average over the whole journey of 21. The error introduced by this assumption is considered later. and e.-N was made up to 14 ml with water and 10 M-NaOH to adjust the pH to 11-12. -N excreted in 12 h (g). it was assumed that all the labelled urea remaining in the body at 12 h had been excreted by 24 h (Grove & Jackson. Ches.5 % from carbohydrate and 7.8 % from protein. 1978b).3 MJ/d of which 56. and 557 g carbohydrate. The flux Q was calculated from the general equation : Q(gN/12 h) = d. for 3 h before 1-2ml was used for mass spectrometry. 338 g fat.
the pre. In the latter 45 d there was less good agreement with isotope values of 18.8 22. If the urea pool turns over with a half-life of about 10 h (Waterlow et al.1 Day 95 Day +7* 2.8 95.0 MS RF 0. The isotope data indicated that MS lost 9.9 kg fat and actually gained 0 1 kg lean tissues.3 MJ/d in MS and 38.168 M.0 89.8 kg lean tissues. values bear this out. 16N in urea (e.and post-expedition values are less accurate than those made during the expedition.3 17. after considerable food intake and weight gain. N intake. corrected by adding lo%. and possibly MS7. are also shown in Table 3. Body weight and body composition of two men.9 MJ/d in RF. before. during and after an unassisted crossing o f Antarctica.3 Day 50 8.5 1. It also shows the percentages of the dose of 15N excreted in NH.7 kg lean tissue during the first 50 d. estimates of body composition from the stable isotopes and UWW are shown in Table 1. There were two cases in which excretion of [15N]ureawas suspiciously low in the period 0-12 h: MS4 and RF3.5 MJ/d in RF. were also exceptionally low in both men and hence tend to support the fact that fat stores were essentially zero on the last day of the journey. it can be calculated that about 8 YOof the [“Nlurea formed will still be retained in the body at 24 h. (e. As discussed in the Methods section.6 Day 50 59. whilst during the last 45 d they were 26. will be far more serious if urine is lost in the first few hours after the dose than if it is lost later on. by direct measurement with [l’N]urea.1 MJ/d in RF. obtained a value of 10% for retention at 24 h and so values of total [l’N]~reaexcreted. and gives body weights and estimated energy and protein intakes at the time of each test. (1981).6 MJ/d in MS and 23.1 MJ/d in MS and 35. The fact that some of the D :N ratios may be . RF3. It also contains some general comments on the condition of the subjects and their situation at the time.0 71.9 Day 0 74. An incomplete collection.0 0.4 Day 0 22.) and 15N in NH. Combining the intake data with the estimated changes in body composition yielded energy expenditures during the first 50 d of 28. In an attempt to clarify these possible losses. Fern et al. At the end of the expedition the isotope data in both men suggested that fat content was zero.9 Age Day (years) -10 37 48 69.). together with the output of creatinine.8 MJ/d in MS and 23. Similar differences were also seen during the latter 45 d of the expedition when MS lost 5. Table 3 shows the 24 h urine volume. These results suggest that the urine collections were incomplete and the NH. even if the loss is small.1 MJ/d for RF (Stroud et al. These values for the first 50 d were similar to those estimated using isotope-labelled water which gave values of 29. creatinine. STROUD A N D OTHERS Table 1. whereas R F lost 22. measured using isotope dilution or underwater weighing Body weight (kg) Subject Body fat (YO) Day +7 584 76.0 * Body fat determined using underwater weighing. The results suggests incomplete collection in MS4. although in both men the isotope dilution spaces were exceptionally large suggesting that there had been considerable increases in the water content of the lean body mass. Table 2 shows the timing of the protein turnover studies. output and the apparent N balance. during the first 12 h and in urea from &12 and from 12-24 h. whereas RF lost 2 2 kg fat and gained 0-4 kg lean tissue. A.3 72. 1978u).8 Day 95 53. Table 3 also shows the excretion during the 12 h daytime period compared with that during the night (D :N) for a number of variables : N.2 Day -lo* 16.5 kg fat and 0. The UWW values for body fat at 7 d post-expedition.3 2. 1996).8 kg fat and 5.
22 6.5 D N T 72. Temperature -35" Recovering in UK Recovering in UK 19.03 26.0 D N T 91.15' Position .79 deg 37 rnin South.2 T 78.85 12.8 D N T 71. 10 MS 71.81 deg 37 min South.18' Position .00 hours (D).86 1797 8.35 Protein intake (g/d) MS RF 248 213 54 50 34 34 88 84 54 57 42 34 96 91 66 66 34 31 100 97 Test no. M S and RF.14 13.72 13.04 12. 20 68. 96 53.3 77. When they reached the ice-shelf of Ross Bay. General conditions Gaining weight before departure Position . corrected for an estimated 10 % retention of e.before. by subtraction of the 12 h N output.39 17. they were so exhausted that they would probably have died if they had tried to reach open water.87 deg 57 rnin South.83 deg 10 rnin South. having successfully completed the crossing of the Antarctic continent. 19. High synthesis rates were maintained throughout the expedition by both men. particularly in eA.3 Post 14 Post 20 58.with little effect on creatinine.22 6. Beyond Pole on plateau at 12400 feet.PROTEIN TURNOVER D U R I N G A N T A R C T I C CROSSING 169 Table 2.9 T Energy intake (MJId) MS 31.) and NH. RF and MS frostbitten.00-07. Episodic hypoglycaemia.7 72.97 8.56 11.50 24.82 3.86 deg 58 rnin South. by the time of test 5 (days 92 and 96).64 1'5.74 4.3 76 39 115 39 17 56 72 49 121 39 17 56 92. conditions and body weights o f two men.18 15. they had been on half rations for 8 (RF) and 12 (MS) d. DISCUSSION It is evident from the weight losses alone that this expedition entailed extremely hard work.00-19.17 RF 2959 10. At the end of the journey. low when that for creatinine is not particularly reduced may be due to a small loss in an early urine specimen having a large effect on the isotope output. Onset of extreme work. Reached land and beginning to climb. Temperature -28" Position . Constantly cold and hungry.49 17.25 5.60 18.60 18.00 hours ( N ) and total 24 h period ( T ) Body weight (kg) Day* Pre 7 9.00 9. 76 56.65 21. The estimates of flux based on urea (Q.) are shown in Table 4 (with Q. (Q. 50 59.0 T 95.0 RF 92.60 17. while MS had frostbitten fingers. Edge of Polar plateau at 8000 feet. The first part of the journey involved climbing 3500 m and in many places the terrain was made difficult and dangerous by crevasses and ridges of iced snow. Marked fatigue and onset of hunger. marked muscle pains. .8 615 203 185 144 162 * The first number is for RF and the second for MS.58 6.32 2629 9. maintained for a period of over 3 months. Good physical condition.0 D N T 76. during and after an unassisted crossing of Antarctica on test days with estimates of energy and protein intakes for 07.88 25.42 22. Position. Both subjects at times experienced symptoms of hypoglycaemia and RF had severe frostbite of the feet. Temperature .25 6. Temperature -48' Position .1 D N T 71. Temperature .95 6. RF frostbitten. Back down on Pacific ice-shelf.8 49. at 24 h) along with synthesis rates derived from Q.27 21.49 26.
9 16.9 13.6 12.3 11.0 7.7 3.90 223 2.2 7.35 0. apparent nitrogen balance.7 1.96 048 0.8 7.2 7. and D : N excretion valuesfor total nitrogen.4 13.32 1.4 1. Urine volume (ml) Intake 4.6 13.87 1.4 9.1 15.87 1.0 -2. “N isotope in urea ( e .68 1.2 5.2 3-3 4.oo 0.83 1.2 12.64 0.3 9-0 19.6 2450 1120 1840 2980 4860 4320 3130 3320 2450 2370 2900 3000 3700 5010 1720 1210 296 25.96 1.1 1 2.13 > z 099 17.0 32.69 1.09 1.0 24.1 Output* 13.7 12.5 9.66 0.5 39. creatinine.0 16.2 1.70 9. .85 6.36 2.2 11.5 8.0 7.7 21.72 152 037 1.6 5.24 0.2 0. creatinine excreti0n.56 0.25 1.7 1.6 14.2 82 171 8.3 12.7 14.percentage of 15N excreted in ammonia.03 1.36 042 0.75 16.63 029 1.15 0.1.49 0.9 12.00-19. Urine volume.95 0.3 * t Not corrected for non-absorbed nitrogen.1 .0 3.9 12.24 1.4 11.2 15.9 12.5 14.6 7.5 2.8 4.7 .0 7.5 19.5 22.00 hours ( N ) and total ( T ) .88 0.50 1.2 1 0.0 126 9.0 18.03 1.7 U 14.5 230 34.08 1.55 0.4 195 13.2 12.85 3.3 78 11.1 0.2 106 13.6 14.56 012 0.) in two men (MS and RF) making an unassisted crossing of Antarctica 15N excreted in urea (“h of dose) e .93 1.5 4.4 12.3 -6.9 -2. percentage of 15Ndose excreted in urea 07.25 1.97 1.9 3. 19.90 076 0. nitrogen intake and output.3 9.27 1.79 1.3 5.8 7.5-1 2.9 Test no.47 0. ) and “N isotope in ammonia (e.0047.1 5.43 044 0.96 1.30 1.5 5.6 8.Table 3 .8 6.69 MS 0 MS 1 MS 2 MS 3 MS 4t MS 5 MS 6 MS 7 RF 0 RF 1 RF 2 RF 3 t RF 4 RF 5 RF 6 RF 7 -6 3 .05 6.26 1.4 5.13 1.7 9.4 152 137 11.4 6.5 20. Urine collection suspected to be incomplete.2 3.73 0.16 1.30 8.9 15.1 13.8 5.4 16.5 167 Creatinine (g/d) l6N excreted in ammonia (% of dose) D T N Corrected T Total N D:N Creatinine D:N D:N eA D:N U 1.59 1.6 5.96 0.4 3.6 2.8 9.8 78 15. Nitrogen (g/d) Balance 15.45 0.3 12.4 8.03 073 1.10 0.82 1.7 0.39 1.5 153 18.103 12.88 1.29 0.6 11.13 0.3 2.8 3.15 079 1.83 043 039 1.09 1.00 hours (D).5 4.3 19.8 1.6 7.9 10.
57 5.3 37.37 9.4 47.46 829 565 39.14 6.3 31.8 57.40 4.5 45.2 31. nitrogenpux estimated from urea (Q.8 - C MS 0 MS 1 MS 2 MS 3 MS 4* MS 5 MS 6 MS 7 Mean (MS 1-5) 9.3 16.27 (SD 0. and protein synthesis rates estimated from Q .86 3.5 21.42 8.0 35.60 9.0 64.4 50.8 - 5.12 - 3.5 59.2 1 2. 2.83 ' 4.3 51.12 1.2 38. Q 2 .01 ~ 28.55 7.04 9.:Q.0 1 3.02 3.9 - 8.6 42-6 1.44 2.04 9.34 8. 0 7.43) z 2 Q * z 53.5 78.0 41.96 1. N loss N loss Q" Q.7 67.52 1221 761 - 59.06 623 3.9 43.8 59-0 75.00 hours.12 11.97 6.4 20.49 11.04 4.3 46.61 0 91 1.77 67.25 I 6.6 68.Nitrogen losses 07.1 26.7 23.5 83.2 61.57 1.5 57.3 46.77 2.48 1.0 - 288 2.71 - C P z 0 < 0 66.07 6.34) =! w RF 0 RF 1 RF 2 RF 3* RF 4 RF 5 RF 6 RF 7 Mean (RF 1-5) - z m * Urine collections were suspected to be incomplete.9 1 680 5.1 36.35 1.58 3.22 2.1 49.07 8.7 502 26.94 2.3 63.24 2.25 - S (mg N/kg per h) S (g protein/kg per 12 h) 4.Table 4.4 35.4 (g/12 h) (mg N/kg per h) (mg N/kg per h) (mg N/kg per h) Q.82 3. ( S )for two men ( M S and R F ) making an unassisted crossing of Antarctica T I P 0 e 2 2 1 Test no.63 11.5 92.24 6.35 6.2 1.0 34.) and ammonia (QJ.00-19.23 8.5 1 5-09 4-58 (SD 1.7 35.22 - 9 H P n c.01 '7.07 8.
This approach gives mean losses of 0. STROUD A N D OTHERS Under these conditions accurate collection. which are also remarkably close to the values derived from estimates of body composition changes. A. These values are almost identical to the mean of the three values of apparent N balance for the same period (MS1-3. but one that must have occurred at a critical time. GI losses of unabsorbed dietary N and other faecal losses normally range from 1-10 YOof dietary intake depending on the type of diet. in this case it would be misleading to make a comparison with the arithmetic mean of the two pertinent measurements (MS4 and 5.8 g N/d. there must have been some additional losses via the gastrointestinal (GI) tract. However. These comparisons of N balance as assessed by two independent techniques suggest that non-accounted N losses must have been very small during the expedition. recording and storing of specimens is extremely difficult and collection of urine in the daytime even risked penile frostbite. approximately 25 O h of . the actual amount of uric acid excreted does not normally rise with an increased protein intake and hence for the present subjects the value of 3 YOof urinary N losses being in this form would seem most likely. the estimates of protein synthesis rates will have inaccuracies dependent on the additional N excreted and it is therefore of interest to examine the likely scale of such unaccounted N losses. pre-packed. but not those in uric acid which may account for 3-5 YOof urinary N for individuals on a diet of about 70 g protein/d. and creatinine.4 g N/d for RF. therefore. RF4 and 5). NH.9 g N/d for RF. because the men were constantly hungry during the expedition. In principal. NH. Overall. In the case of this expedition the low intake of dietary fibre and the probability of enhanced colonic salvage would make it likely that GI losses were at the lower end of this scale and hence that less than 1 g N/d was lost by this route. soon after the dose of isotope had been given.9 g N/d for MS and gains of 0. and other N sources excreted are irrelevant to the calculation of flux which depends only on 15N abundance in the end-products. The results must be examined with this in mind. almost surprising that. During the course of the expedition.2 g N/d for RF. only two tests. It is interesting that it was MS who was in apparent negative N balance from both bodycomposition studies and estimates of N intake and urinary losses. Skin losses are usually about 20 mg N/kg body weight and so would amount to a maximum of 1. whereas RF tended to be in positive balance. However.172 M. we believe that total unaccounted N losses are unlikely to have exceeded 4g/d and would have led to overestimates of protein synthesis rates of only 5-10 %. However. RFl-3) which gave losses of 4. a better mean for the latter 45 d period must be calculated by assuming that N balance during the 34 d up to day 84 was that measured in MS4 and RF4. since the men reduced to half rations on day 84.6g/d. simple daily rations. while for 1 1 d beyond the switch to half rations it was that measured in MS5 and RF5. However. one for each subject. The urinary losses accounted for in t h s study included those in urea. However. MS tended to be in negative apparent N balance. presumably due to efficient colonic salvage. had to be rejected because of losses of urine. although the outputs took no account of faecal and other losses. This does not imply a large loss. there was essentially no plate wastage and with pre-weighed. the skin and nonmeasured urinary uric acid.8 g N/d for MS and gains of 0. the amounts of urea. It is.7 g N/d for RF. the estimates of dietary intake are likely to be very accurate.9 g N/d for MS and gains of 0. In this case maximal additional urinary N losses would be of the order of 0. This would suggest that non-urinary N losses during the period were very small. Instead. The rates of change of lean tissue can be calculated from the body composition data and for the first 50 d there were losses of 4. Garrow (1974) reported that when obese subjects lose weight through dietary restriction. as far as we can tell from the internal evidence. For the second 45 d period the body composition data suggested a similar pattern of changes with losses of 0 8 g N/d for MS and gains of 0. therefore.
although much smaller. in recovery tests 6 and 7. The Antarctic expedition provided a scenario in which all three of these elements affecting protein turnover were combined: intense exertion. The D: N ratios for 15N in NH. However. with breakdown predominating to produce a negative N balance (Tomkins et al. although variable.PROTEIN TURNOVER D U R I N G A N T A R C T I C CROSSING 173 losses are lean. whereas Q.25-1-30 (Fern et al. Waterlow. Jeevanandam et al. were more varied but probably have little meaning. Lemon (1991) concluded that protein requirements in very active individuals may be as high as 160% of the recommended dietary allowance. unpublished results). and for the successful collection periods during the expedition it averaged 2. It seems likely. if this was the case one might have expected similar lean losses in RF. However. The two estimates of flux that are obtained from the end-products urea and NH. synthesis and breakdown. and Millward et al. On the food side. both these reviews assume that individuals are in energy balance and during this expedition this was not the case. The ratio Q. the differences in N balance between the two men suggest that they behaved very differently in their metabolic responses to the stresses of the expedition and this individuality was also evident in the pattern of their N excretion.. In a review. (1994) concluded that in fully trained individuals. Values obtained for this ratio in normal subjects are in the range 1. both components of protein turnover.87 in MS. since far more NH. some degree of infection and trauma. 1983. and food deficit. In such children the rate of protein turnover is reduced (Waterlow. 1981. Since MS. and Q. Q. There is evidence that acute physical exercise leads to an increase in body protein breakdown and a decrease in synthesis (e. an observation that was also made during an expedition to the Andes. The end-product method of measuring protein turnover with [15N]glycinerests on a number of assumptions which have been justly criticized. Wolfe et al. but nothing is known about the effects of very-long-term exertion. he also noted that there is a decline in this exercise-inducedprotein requirement with continued training. The dietary protein requirements for men undertaking endurance exercise of this nature are unclear. 1986). was pulling a sledge of equal weight to RF. are not additive. where day and night collections were made for 24 d (J. therefore. The main purpose of the present study was to examine how the basic biological process of protein turnover responded to conditions of extreme stress and negative energy balance imposed over a long period.. 1984). is biased towards the metabolism in the splanchnic bed because urea is synthesized in the liver. was almost always higher than this. It is suggested that Q. Fern & Garlick. are increased. N losses associated with physical activity are minimal. Whatever the cause. QU:QA . whereas the opposite tended to be the case for RF. therefore. expedition. give us a glimpse of the relative rates of protein synthesis in these two parts of the body. In both RF and MS Qu:QA. In subjects exposed to infection or trauma. However.: Q. is excreted by day and hence small fluctuations in night-time excretion have a big effect on ratios. the difference in N balance results might reflect the relatively higher exercise intensities at which MS must have worked for some of the journey. and hence energy deficits alone could have been responsible for the 109 % lean loss in MS. MS excreted more N and more 15Nin urea during the night than the day. 1992). is probably the muscles.. but are separate estimates of whole-body N flux. For example. is biased towards the peripheral tissues because the main source of glutamine. The D :N ratios for creatinine were close to unity in both men. Following the in MS was well below the normal level. the precursor of urinary NH. perhaps indicating a relatively greater flux through the visceral organs with high protein feeding. but there is no other method that can be used in the field.05 in R F and 1. C. that protein oxidation was needed to fuel exertion. 1983). particularly during moderate-to-high-intensity exercise when it is likely to have provided 5 1 0 % of energy supply.g. may. perhaps the best analogy is the severely malnourished child who for a long time has been short of energy and protein.
Legaspi. The concept of the single body pool of metabolic nitrogen in determining the rate of whole body nitrogen turnover. (1994). & Rennie. Horowitz. C.Clinical Science 68. Report of sub-committee on constants and factors. B. Botwell. McNurlan. British Journal o f Nutrition 74. J. R. P.. 491-507. average values for the four successful measurements made during the course of the expedition were also higher than might have been expected at 3. M. Blaxter. This indicates the vital role of protein synthesis in physiological function as long as life is not extinct. Human Nutrition: Clinical Nutrition 39C. Proceedings o Stroud. E. Constantin-Teodosiu. D. Garlick. B. L. C. M. Clinical Science 61. J. Fern. should be used (Fern & Garlick. Nutrition and energy balance on the “Footsteps of Scott” expedition 1984-86.. (1993). E165-El74. H. Energy expenditure using isotope-labelled water (‘H. M. Ritz. that the overall message from these findings is that even in the face of great exertion. either arithmetic or harmonic. A. M. therefore. Golden. J.174 M. & Waterlow. (1978~). The excretion of isotope in urea and ammonia for estimating protein turnover in man with 15N-glycine. Shadows on the Wasteland. Garlick. A. E. (1974). P.. In Energy Metabolism. W. Protein turnover in man measured with 15N: comparison of end products and dose regimes. 313-324. Energy Balance and Obesity in Man. b). Lowry.. 4148. Isotopic determination of amino acid-urea f Applied Physiology 56. protein synthesis can be maintained. 93.. (1991). (1981).. STROUD A N D OTHERS perhaps indicating that restoration of lost muscle was taking priority. Wolfe. R. 15-37. skeletal muscle enzyme activities and plasma biochemical parameters in man during 95 days endurance exercise with inadequate energy intake. J. Amsterdam: Elsevier and North Holland Press. Garlick. since values of Q. Stroud. P.. Fern et al. protein metabolism and protein f the Nutrition Society 53. Waterlow. REFERENCES Brouwer. Fern. M. & Millward.217-228. J. Protein Turnover in Mammalian Tissues and in the Whole Waterlow. J. F. Pacy. interactions in exercise in humans. & Sawyer. N. W. C. Influence of increasing dietary intake on whole body protein kinetics in normal man. 1985a. A. E. F. (1992). Fern. Coward. J. P. F. Waterlow. D. E. B. (1965). H. probably because Q. P. However. J. J. S. R. M. (1978b). Lemon. D. Body. 85-89. which might also account for the high values seen in the first post-expedition assessments. B. P. & Waterlow. Physical activity. marked negative energy balance and serious physical deterioration. G . Garrow. J. Grove. although in RF low values were not seen at this time.. & Jackson. E. R. A. The mean values for protein synthesis in normal human subjects are between 2 and 3 g protein/kg per 12 h (Fern et al. is subject to a number of influences such as changes in acid-base status. L. 1983. P.’’O). (1985~).. Human Nutrition: Applied Nutrition 41A. However. Measurement of protein turnover in normal man using the end-product method with oral [15N]glycine: comparison of single-dose and intermittent-dose regimens.426-433. Fern & Garlick. C. J . (1983). (19856). (1986). 271-282. S . A. D. C. A. Protein-energy Malnutrition. A. probably due to overfeeding. 223-240.. M. Schofield. Jeevanandam. P.. & Shaw. Nadel. (1995). Tomkins. Apparent compartmentation of body nitrogen in one human Fern. in the present study were very variable. editor].. & Waterlow. J. requirements.. M. 1983) and hence the values for the pre-expedition tests were markedly raised in both men. Amsterdam: North Holland. L. J. & Brennan. London : Academic Press. A. A. pp. W. The combined effects of infection and malnutrition on protein metabolism in children.. 1981. Greenhaff. and we believe. Millward. and QA. Clinical Science 65. Does exercise alter dietary protein requirements? Advances in Nutrition and Top Sport 32.. J... p. P. The rate of whole body metabolism in the whole body of man measured with 15N glycine and uniformly labelled 15N wheat. the synthesis rates given in Table 4 were derived only from Q. B. Wole.. American Journal o f Physiology 4. 221-229. Stroud. G. J. Journal o Printed in Great Britain . C. & Waterlow. 91-107. J. Garlick. J. & Garlick. C. & Garlick. P.Crossing Antarctica with Sir Ranulph Fknnes.3 g protein/kg per 12 h for MS and 4.. European Journal o f Applied Physiology (In the Press). M. A. J. (1984). E. 441443 [K. (1983).. London: Jonathan Cape. J. exercise performance. M. London: Edward Arnold. (1987). (1996). Clinicai Nutrition 5. J. Human Nutrition: Clinical Nutrition 37C.6 g protein/kg per 12 h for RF. H. subject: its consequences in measuring the rate of whole body protein synthesis with 15N. Garlick. J. M. For the best estimate of total N flux it has been proposed that an average of Q.