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The relationship of individual anaerobic thresholds to total, alactic, and lactic oxygen debts after… Wiley, James Preston 1980

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THE RELATIONSHIP OF INDIVIDUAL ANAEROBIC THRESHOLDS TO TOTAL, ALACTIC, AND LACTIC OXYGEN DEBTS AFTER A SET TREADMILL  RUN  by  JAMES PRESTON WILEY B.P.E., The U n i v e r s i t y  of B r i t i s h Columbia, 1977  A THESIS SUBMITTED IN PARTIAL FULFULLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF PHYSICAL EDUCATION  in THE FACULTY OF GRADUATE STUDIES School o f P h y s i c a l E d u c a t i o n and R e c r e a t i o n  We accept t h i s  t h e s i s as  to the r e q u i r e d  conforming  standard  THE UNIVERSITY OF BRITISH COLUMBIA April  ©  1980  James P r e s t o n Wiley,  1980  DE-6  In p r e s e n t i n g  t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f the r e q u i r e m e n t s f o r  an advanced d e g r e e a t the U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and I f u r t h e r agree that permission f o r s c h o l a r l y p u r p o s e s may by h i s r e p r e s e n t a t i v e s .  for extensive  study.  copying of this thesis  be g r a n t e d by the Head o f my Department o r It i s understood that copying or p u b l i c a t i o n  o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be a l l o w e d w i t h o u t my written  permission.  Department Of  Physical Education  The U n i v e r s i t y o f B r i t i s h C o l u m b i a 2075 Wesbrook P l a c e V a n c o u v e r , Canada V6T 1W5  BP  75-51  1E  ABSTRACT Anaerobic threshold speed  was determined for 20 male uni-  versity students using a continuous treadmill protocol.  The onset of  anaerobiosis was determined by analyzing excess C O 2 elimination. following week, a l l subjects ran at the per hour for 10 minutes. after this run.  The  median speed of 7.25 miles  Recovery oxygen consumption was monitored  Application of double exponential equations by  computer and subsequent integration, calculated Total, Alactic, and Lactic Oxygen Debts.  Subjects who ran above their V ^  (group L-V-^)  had significantly (p < .05) higher total, lactic and alactic debts than those subjects who ran below their"V-.., (group H-V_,„). TAM TAM J  &  r  The total debt  showed a significant (p < .05) negative correlation (r=-.77) to group L'-V_^.  in  ;  This appears to be due to the increasing lactic debt,  that was also significantly (p < .05) negatively correlated (r=-.73) to V_ ,.' TAM sll  Group H-V_.,, did not exhibit this characteristic. TAM r  This study  demonstrates that V_.„ is a c r i t i c a l factor in determining oxygen debt TAM and therefore, work above this point results in the onset of metabolic acidosis, which may limit the optimal running speed for a given distance.  TABLE OF CONTENTS ABSTRACT  i i  LIST OF TABLES  v  LIST OF FIGURES  .  v  ACKNOWLEDGEMENT  v i i i  Chapter 1 STATEMENT OF THE PROBLEM Introduction Statement of the Problem Hypotheses Rationale Assumption Delimitations Limitations Definitions 2  3  4  ..... ...... ;  7  Historical Considerations Anaerobic Metabolism and Oxygen Debt . . . . . Anaerobic Threshold .......  METHODS AND PROCEDURES  . . . . . . . . .  7 8 12 24  Subjects Testing Procedures Data Analysis  24 24 25  RESULTS AND DISCUSSION  27  Results Discussion 5  1 1 3 3 4 5 5 5 5  REVIEW OF THE LITERATURE I. II. III.  i  27 41  SUMMARY AND CONCLUSIONS  46  iii  REFERENCES APPENDIX A  48 Sample Calculation of Oxygen Recovery Debts  . . . . 54  APPENDIX B Anaerobic Threshold Curves to Determine Individual V  56  APPENDIX C  78  T A M  Individual Oxygen Recovery Curves  iv  LIST OF TABLES  Table 1. I n d i v i d u a l S u b j e c t C h a r a c t e r i s t i c s 2. I n d i v i d u a l Double E x p o n e n t i a l E q u a t i o n s f o r S u b j e c t s i n Group L - V ^ 3.  I n d i v i d u a l Double E x p o n e n t i a l E q u a t i o n s f o r S u b j e c t s i n Group H-V-.., TAM I n d i v i d u a l A l a c t i c , L a c t i c , T o t a l and R a t i o Oxygen Debts f o r Group L - V _ „ TAM I n d i v i d u a l A l a c t i c , L a c t i c , T o t a l and R a t i o Oxygen. Debts f o r Group H-V_._, TAM Means o f T o t a l , A l a c t i c , L a c t i c and R a t i o Debts f o r Groups L-V_ .. and H - V _ „ ^ TAM TAM M u l t i v a r i a t e A n a l y s i s o f Dependent V a r i a b l e s T o t a l Debt and R a t i o Debt M u l t i v a r i a t e A n a l y s i s o f Dependent V a r i a b l e s A l a c t i c and L a c t i c Debts C o r r e l a t i o n C o e f f i c i e n t s f o r Subjects i n Group L - V C o r r e l a t i o n C o e f f i c i e n t s f o r Subjects i n Group H - V  28 29 30  r  4.  A  32  r  5.  6.  4  7. 8. 9. 10.  A  33  38  39 39  T A M  40  T A M  40  v  LIST OF FIGURES Figure 1.  Graph of Total Debt and Individual V_,^  34  2.  Graph of Alactic Debt and Individual V_,^  35  3.  Graph of • Graph of AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve AT curve Recovery Recovery Recovery Recovery Recovery Recovery Recovery Recovery Recovery  4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34.  Lactic Debt and Individual V_.. TAM Individual Ratio Debt and subject CN subject ML subject RR subject TB . . subject DG subject DD subject BV subject GS subject DA subject RF subject HB . . . . . . . . . . . subject AB subject AO . subject GT . . subject DH subject JL subject JO subject DM subject DW subject SP . . . . subject DH curve subject CN curve subject ML curve subject RR . . curve subject TB . . . curve subject DG curve subject DD curve subject BV curve subject GS curve subj ect DA x  vi  .  36 37 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 79 80 81 82 83 84 85 86 87  35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45.  Recovery Recovery Recovery Recovery Recovery Recovery Recovery Recovery Recovery Recovery Recovery  curve curve curve curve curve curve curve curve curve curve curve  subj ect RF subj ect HB . subj ect subj ect subj ect subj ect subj ect JL subj ect subj ect subj ect subj ect  88 89 . • 90 . . . 91 92 93 94 .. . . . . . . . . 95 . 96 97 98  vll  ACKNOWLEDGEMENT  The author extends a p p r e c i a t i o n  t o t h e members o f the committee  (Dr. E. C. Rhodes [Chairman], Dr. K. C o u t t s , Dr. J . Ledsome, and Dr. R. Schutz) f o r t h e i r work and e f f o r t t h a t molded t h i s t h e s i s t o i t s p r e s e n t form.  In p a r t i c u l a r , t h e author i s i n d e b t e d t o Dr. Rhodes, whose  p a t i e n c e and a s s i s t a n c e thanks a r e extended  guided t h e d i r e c t i o n o f t h i s t h e s i s .  t o Mr. D. Dunwoody, whose t e c h n i c a l  time, and f r i e n d s h i p made t h i s t h e s i s  viii  possible.  Special  assistance,  CHAPTER 1  STATEMENT OF THE  PROBLEM  Introduction Anaerobic  metabolism and  i t s e f f e c t on r e c o v e r y was  gated by H i l l , Long and Lupton (1924). terms as moderate and  first  investi-  At t h a t time, such work l e v e l  severe were used to d e s c r i b e the e x e r c i s e t h a t p r o -  duced e i t h e r immediate or extended r e c o v e r y oxygen uptake s i t u a t i o n s . Since t h i s c l a s s i c r e c o v e r y curve curve  i n v e s t i g a t i o n , r e s e a r c h e r s have l a b e l l e d  the oxygen  (Margaria et a l . , 1933), q u a n t i f i e d the oxygen r e c o v e r y  (Henry & DeMoor, 1950), and attempted to e x p l a i n the oxygen  r e c o v e r y curve  (Huckabee, 1958b).  However, no one has  examined oxygen  r e c o v e r y curves w i t h r e s p e c t to i n d i v i d u a l onset of a n a e r o b i c Although exercise  some a n a e r o b i c m e t a b o l i c a c t i v i t y c o n t i n u a l l y o c c u r s  of  during  (Graham, 1978), o n l y when l a c t a t e d i f f u s e s i n t o venous b l o o d  from the muscle t i s s u e s i s i t p o s s i b l e to e a s i l y monitor ity.  metabolism.  anaerobic  flow  activ-  T h i s i n c r e a s e i n venous l a c t a t e p r o d u c t i o n appears to be the  result  the imbalance between l a c t a t e p r o d u c t i o n and o x i d a t i o n i n the muscle  tissues  (Graham, 1978).  been the standard (1933) f i r s t  L a c t a t e i n the venous c i r c u l a t o r y system  f o r examining work i n t e n s i t y s i n c e M a r g a r i a  examined phases of l a c t a t e p r o d u c t i o n and  et a l .  removal.  R e s u l t s of these e a r l i e r i n v e s t i g a t i o n s (Margaria et a l . , 1933; & Edwards, 1934b) and more r e c e n t r e s e a r c h demonstrate the f o l l o w i n g : 1  has  Margaria  (Hermansen & S t e n s v o l d ,  1972)  2 1.  there is a greater lactate concentration for higher work intensity,  2.  lactate levels stabilize for a l l but the very extreme levels of steady-state work, and  3.  long recovery periods are associated with high lactate levels. These early investigations imply that there is a point in the tran-  sition from aerobic to anaerobic work intensity where anaerobic metabolism starts to play a very important role.  This increased role is marked  by the appearance of lactate in the blood in excess of resting levels. I n i t i a l investigation (Issekutz & Rodahl, 1961; Naimark et a l . , 1964;  Wasserman et a l . ,  1964;  Wasserman & Mcllroy, 1964) monitored this phenom-  enon and associated cardiovascular-respiratory changes.  This resulted  in non-invasive procedures involving measurement of respiratory gas exchange to determine the onset of metabolic acidosis. The term "Anaerobic Threshold" (AT) which has become synonymous with the onset of metabolic acidosis, was originally defined by Wasserman et a l . (1964) as " . . .  the work level (oxygen uptake) above which the  subject develops metabolic acidosis."  This definition i s convenient in  that i t is the limit of predominately aerobic energy production (as determined by normal blood lactate levels) but misleading in that no detectable anaerobic activity has occurred (again as determined by blood lactate level).  Most studies use Wasserman's definition of the AT but  some define the AT as the point where blood lactate production just exceeds i t s removal (Davis et a l . , 1976). The study of the role of the AT and i t s relationship to human performance has only just begun.  This phenomenon has been used for testing  hospital patients (Wasserman & Whipp, 1975), athletes (Costill, 1970), and models of hypernea (Whipp, 1977).  However, the role of the onset  3  of anaerobic metabolism and i t s relationship to recovery analysis has not yet been investigated. Statement of the Problem The purpose of this study was to investigate the effects of individual Anaerobic Threshold Levels on the accumulation of an Oxygen Debt for a set treadmill run. Hypotheses 1.  Subjects with an Anaerobic Threshold speed (V--^) above the speed of the treadmill (group -H-V-,^) have a lower Total Oxygen Debt than those with a V ,„ below the speed of the treadmill (group L-V^.^,). TAM TAM Group H-V , -has a lower Alactacid Debt than group L-V_,„. m  2. 3.  m  Group H-V__„, has a lower Lactacid Debt than group L-V__ f TAM ° TAM Group H-V_,._, has a lower ratio of Lactacid Debt to Alactacid Debt TAM y  4.  than group L ~ V  TAM>  5.  Group H-V_,^ has a lower Total Recovery Time than group L-V_,^.  6.  Within group L-V  7.  V TAM Within group L-V  , Total Oxygen Debt is a decreasing function of  , Alactacid Debt i s a decreasing function of 1 JTli 1  V TAM 8.  Within group l  -  ^ ^ '  Lactacid Debt is a decreasing function of  TAM"  V  9.  Within group rj,^> L-V  t n e  ratio of Lactacid Debt and Alactacid Debt  is a decreasing function of V .„. m  10.  W i t h i n group L - V „ ^ , T o t a l Recovery time i s a d e c r e a s i n g °  11.  f  V  function  TAM'  W i t h i n group H-V_.„, T o t a l Oxygen Debt and V_,,,, a r e u n r e l a t e d . TAM ° TAM J  12.  W i t h i n group H-V„.„, A l a c t a c i d Debt and V_ „, a r e u n r e l a t e d . TAM' TAM A  &  13.  r  W i t h i n group H-V..,,,, t h e L a c t a c i d Debt and V _ , a r e u n r e l a t e d . ^ TAM TAM w  &  14.  W i t h i n group H-V ^ , and  15.  t  n  e  r a t i o of L a c t a c i d Debt t o A l a c t a c i d  Debt  are unrelated.  W i t h i n group H-V_. , T o t a l Recovery time and V_._, a r e u n r e l a t e d . ^ TAM TAM . w  a  J  Rationale The A n a e r o b i c T h r e s h o l d i s t h e t h r e s h o l d o f appearance o f venous l a c t a t e as a r e s u l t o f a n a e r o b i c metabolism.  As d e f i n e d , V . „ i s t h e TAM m  r u n n i n g speed above which t h e s e m e t a b o l i t e s s t a r t t o i n c r e a s e . f o r e , a s u b j e c t r u n n i n g a t a speed above h i s V ^  There-  w i l l have a l a r g e r  c o n t r i b u t i o n of a n a e r o b i c metabolism as an energy s u p p l y , r e l a t i v e t o a s u b j e c t r u n n i n g a t a speed below h i s V T h i s suggests t h a t A l a c t a c i d and L a c t a c i d Debts w i l l be l a r g e r f o r the  s u b j e c t r u n n i n g a t a speed above h i s  I f this i s true,  i m p l i e s a l a r g e r T o t a l Oxygen Debt and a l o n g e r Recovery The d i f f e r e n c e between t h e t r e a d m i l l speed and j e c t s r u n n i n g f a s t e r than t h e i r V  should be r e f l e c t e d  c o n t r i b u t i o n o f a n a e r o b i c metabolism t o t h e i r a c t i v i t y .  this  Time. ( f o r t h e subi n a larger T h i s suggests  t h a t t h e components o f Oxygen Debt and V^^. w i l l be i n v e r s e l y  related.  Furthermore, i t i s s p e c u l a t e d t h a t t h i s r e l a t i o n s h i p w i l l be approximately  linear.  5 This relationship is not expected for those subjects running below their V TAM Assumption That an oxygen consumption during recovery after exercise can be described by a double exponential equation. Delimitations This study is delimited by: 1.  the sample type (college age males),  2.  the sample size,  3.  the sample's fitness levels,  4.  the speed of the treadmill at which the subjects w i l l run, and  5.  the length of the time each subject is on the treadmill before recovery.  Limitations This study's results are limited by: 1.  data collection capabilities of the Beckman Metabolic Measurement ,Cart and the Hewlitt Packard Data Acquisition system interfaced with it,  2.  the method of Anaerobic Threshold determination, and  3.  the individuals' metabolic responses to the protocols.  Definitions 1.  Alactacid Oxygen Debt (Alactic Debt):  theOxygen Debt incurred as a  result of decreased body oxygen stores and high energy phosphate compounds (the fast exponential recovery phase). 2.  Anaerobic Threshold:  the point above which excess C O 2 production  6 is associated with elevated blood Lactate levels as a result of anaerobic metabolism. Excess Lactate (XL):  as calculated by Huckabee (1958a) is  XL = (L -L ) - (P -P )(L /P ) no n o o o where: L  L P P 4.  = concentration of blood lactate at time of sampling  n  = concentration of blood lactate at rest  o  = concentration of blood pyruvate at time of sampling  n  = concentration of blood pyruvate at rest  o  Excess  C O 2  is the excess of carbon dioxide in expired air-  ( E X C C O 2 ) :  (Issekutz & Rodahl, 1961) expressed as: ExcC0 = VC0 - V0 ARQ 2  2  2  where: C0 O2  2  is the total expired carbon dioxide is the total expired oxygen  ARQ is the resting RQ (determined by Issekutz & Rodahl as .75). 5.  Lactacid Oxygen Debt (Lactic Debt):  the oxygen debt in excess of  Alactic Debt due to lactate oxidation and higher metabolic rate (the slow exponential recovery phase). 6.  Total Oxygen Debt:  the total oxygen required during recovery in  excess of resting. 7.  Total Recovery Time:  the time from cessation of exercise until  oxygen consumption returns to resting. 8.  V_,. : the treadmill speed corresponding to the Anaerobic TAM  Threshold.  CHAPTER 2  REVIEW OF THE LITERATURE  I.  H i s t o r i c a l Considerations E a r l y r e s e a r c h of a n a e r o b i c metabolism and it's r e l a t i o n s h i p t o  physical activity  i n v e s t i g a t e d oxygen uptake curves a f t e r e x e r c i s e .  f i r s t major study was completed t h i s i n v e s t i g a t i o n , the f i r s t  by H i l l ,  attempt  Long and Lupton  (1924).  The  In  at quantifying exercise exertion  w i t h r e g a r d t o r e c o v e r y time and b l o o d l a c t a t e accumulation was made. The d i s t i n c t i o n between moderate and severe e x e r c i s e and the p o s s i b l e r o l e of l a c t a t e i n d e t e r m i n i n g oxygen consumption a f t e r e x e r c i s e was investigated.  R. M a r g a r i a  (Margaria et a l . , 1933;  Margaria  & Edwards,  1934a, b) f u r t h e r r e s e a r c h e d oxygen consumption a f t e r e x e r c i s e and developed  r e c o v e r y oxygen concepts  s t u d i e s supported  t h a t a r e s t i l l p o p u l a r today.  a two-phase r e c o v e r y system where t h e i n i t i a l  These fast  r e c o v e r y component was termed the ' a l a c t a c i d debt' and t h e l a t e r , r e c o v e r y component was d e s i g n a t e d the ' l a c t a c i d debt'. the f a s t repayment phase repays repayment phase corresponds Berg  f a s t energy  slower  The concept  supply systems and t h e slow  t o b l o o d l a c t a t e removal was h y p o t h e s i z e d .  (1947) a p p l i e d a s i n g l e e x p o n e n t i a l e q u a t i o n t o oxygen and  carbon d i o x i d e r e c o v e r y curves a f t e r e x e r c i s e , as suggested ( H i l l e t a l . , 1924).  that  earlier  T h i s e q u a t i o n v a r i e d dependence between i n d i -  v i d u a l s and c o n d i t i o n s , and Berg advanced no p o s s i b l e i n t e r p r e t a t i o n  7  8 for these results.'  Later, Henry and DeMoor (1950) applied •-•"1" !  double exponential equations to the oxygen recovery curve.  This double  exponential equation f i t the data points extremely well and provided a mathematical way to separate the oxygen recovery curve into alactacid and lactacid components, as f i r s t suggested by Margaria et a l . (1933).  The  following year, Henry (1951) postulated that the alactacid component is an essential phase of any physical exertion due to equilibria processes that the exponential function predicts.  This added support to his double  exponential model. Subsequently, Henry and DeMoor (1956) investigated the effect of varying work intensity on a bicycle ergometer with regard to oxygen debt. Their findings demonstrated that increased work intensity produced a larger oxygen debt and recovery lag.  Double exponential equations  accurately described a l l recovery curves and showed an increasing lactacid component with increasing intensity. It is from this model, developed by Margaria et a l . (1933) and Henry and DeMoor (1950), that current recovery curve analysis is based. II.  Anaerobic Metabolism and Oxygen Debt In 1958, W. E. Huckabee produced a series of articles (Huckabee,  1958a, b, c) that attempted to explain anaerobic metabolism and i t s relationship to oxygen recovery curves.  He proposed, based on lactate:  pyruvate equilibria relationships, that oxygen debt is directly related to Excess Lactate (XL) production.  Excess lactate traced the oxygen  recovery curve exactly and could be used as a predictor for total oxygen debt.  This was in direct contrast to the concept of a two-phase  recovery curve as supported by Margaria et a l . (1933) and Henry and  9 DeMoor (1950). This finding was subsequently investigated by other researchers. Knuttgen (1962), using four different work rates on a bicycle ergometer, observed that past a " c r i t i c a l level of work," there was a rapid increase in oxygen debt.  This increase in debt was paralleled by rises in lactate  and excess lactate. oxygen debt.  However, this parallel rise underestimated the  At the lower levels of work, where the fast component of  recovery was dominant, the level of lactate or excess lactate could not predict the oxygen debt. Margaria et a l . (1963) also observed no increase in venous lactate until a specific exercise intensity.  However, below this work inten-  sity, an oxygen debt is s t i l l measurable and possibly attributable to the alactacid component.  This study once again demonstrated the pos-  sible existence of a two-phase oxygen debt payment. An editorial in the Annals of Internal Medicine by R. E. Olson (1963) cautioned the use of "Excess Lactate" as a means of signifying anaerobiosis.  The assumption that the lactate/pyruvate ratio reflects  the DPNH/DPN ratio in the c e l l and that the latter reflects aerobic capacity at a given time is disputed. The presence of at least two DPNH/DPN pools, with at least one membrane (mitochondrial) separating hydrogen ions, suggests that the lactate/pyruvate ratio may not reflect mitochondrial deficiency as much as limited hydrogen transfer from the cytoplasm to the mitochondria. Wasserman et a l . (1965) also challenged Huckabee's debt explanation, as i t directly conflicted with their concept of anaerobic threshold  (Wasserman & McTlroy, 1964; Wasserman et a l . , 1964).  Their find-  ings support the concepts of Knuttgen (1962) and Margaria et a l . (1963). This was especially true at the low work levels (subthreshold) where  10 Wasserman et a l . (1965) found excess lactate to predict less than ten percent of oxygen debt. Thomas et a l . (1965) also investigated the excess lactate concept. In a repetition of Huckabee's original works (with slight procedural modifications), Thomas failed to demonstrate the correlations for excess lactate that Huckabee presented.  The investigation did reconfirm the  role of lactate and pyruvate as indicators of anaerobic activity, however,  made no attempt to further delineate the mechanisms. Research next advanced to the analysis of the compartments of  oxygen debt.  Piiper et a l . (1968), utilizing intact dog gastrocnemius  muscle, observed a decreased concentration of high energy'phosphates (specifically creatine phosphate) and oxygenated myoglobin during stimulated exercise.  They hypothesized that the return to resting values  is accomplished during the alactacid phase of recovery (as mathematically predicted by Henry [1951]). DiPrampero and Margaria (1968).conducted a similar experiment with inovivodog gastrocnemius muscle.  Their analysis showed a decreasing  creatine phosphate concentration with increased exercise intensity while the concentration of ATP and ADP at steady-state remained constant. This decrease and the linear relationship between exercise oxygen uptake at steady-state and alactacid oxygen debt (reflecting phosphagen resynthesis half reaction time) supports creatine phosphate's resynthesis role in alactacid oxygen debt. Cerretelli (1969) continued this line of investigation to determine energy equivalents for the lactate and alactic phases (associated with decreased phosphagen stores).  The linear equation for energy expendi-  ture for these two phases, based on modern hypotheses, does not  11 significantly differ from that calculated by Margaria et a l . (1934). This fact supports the two component oxygen debt hypothesis empirically. Direct evidence of creatine phosphate synthesis after exercise was investigated by Piiper and Spiller (1970).  Their data shows almost com-  plete phosphagen resynthesis in.the f i r s t two minutes of recovery. During this time period, the alactacid oxygen recovery component appears to explain the resynthesis of phosphagen.  Total oxygen debt over-  predicted high energy phosphate resynthesis. In 1970, a very extensive investigation into oxygen debt and submaximal exercise was completed by Knuttgen.  Oxygen debt was observed  after steady-state work performed at varying maximal oxygen uptake percentages and at different work durations.  In his discussion, he stressed a  two-phase recovery system as i t accurately described a l l the results. The slow component increased exponentially, and the fast component increased linearly after an i n i t i a l basal rate as work intensity increased. Only the slow component increased as the time duration : increased at a constant work load.  This did not correlate with exercise blood lactate  levels, therefore adding more doubt to the postulate that the slow component is related only to blood lactate levels.  Knuttgen (1970) sug-  gested that other.factors, such as hormones, body temperature, and electrolyte changes might also be considered. A comparison of steady-state and nonsteady-state exercise, employing different time durations for a constant workload was researched by Whipp et a l . (1970).  Their conclusions suggested that the difference seen in  efficiency between short and longer term exercise related to oxygen debt, could be compared to: 1.  delay of alactic debt repayment during steady-state, and  2.  lactate accumulation during the second and third minutes of  12 e x e r c i s e due to d e l a y i n oxygen s u p p l y . They d i d not attempt  to s e p a r a t e the r e c o v e r y curve i n t o two  components.  A s i m i l a r experiment by McMiken (1976) demonstrated work i n t e n s i t y to  be a key i n p r e d i c t i n g oxygen debt  (r=0.83).  He a l s o showed t h a t  s t e a d y - s t a t e and n o n s t e a d y - s t a t e debt were q u i t e s i m i l a r , but d i d not imply t h a t the p r e v i o u s i n t e r p r e t a t i o n o f Whipp et a l . (1970) was  incor-  rect. Current a n a l y s i s of r e c o v e r y has c e n t e r e d on supramaximal (Katch, 1972;  diPrampero  & Morton,.1978). of  the a l a c t a c i d  et a l . ,  1973;  Freund & Gendry, 1978;  Roberts  The main t h r u s t of t h i s work has been the examination component of r e c o v e r y .  The d e l a y of a n e x p o n e n t i a l  r e t u r n to b a s a l l e v e l xvas seen i n diPrampero's  (1973) work which  gested a d e l a y e d r e t u r n of metabolism to normal  (1978) demonstrated  the r e l i a b i l i t y  sug-  (perhaps an a n a e r o b i c  r e c o v e r y phase as suggested by C e r r e t e l l i et a l . [1974]). Morton  work  Roberts and  of t h i s t e c h n i q u e to d e t e r -  mine maximum a l a c t i c debt but so f a r o n l y a few p i e c e s of a v e r y l a r g e p u z z l e have been found.  III.  Anaerobic Threshold Since M a r g a r i a et a l . (1933) monitored l a c t a t e b u i l d - u p i n the  b l o o d , most i n v e s t i g a t i o n has i n v o l v e d the time c o u r s e of l a c t a t e d u r i n g and a f t e r e x e r c i s e . "critical  However, i n 1962,  Knuttgen  l e v e l of work" where l a c t a t e f i r s t  l e v e l of work had been seen b e f o r e (Huckabee, t i o n appears to be the f i r s t I s s e k u t z and Rodahl  to attach  (1962) n o t i c e d a  appears i n the b l o o d . 1958), but t h i s  This  investiga-  s i g n i f i c a n c e to i t .  (1961) observed a l a c t a t e breakaway u s i n g the  change i n the R e s p i r a t o r y Q u o t i e n t and Excess C0 .. 2  T h i s was  the  first  13 attempt at noninvasively following lactate levels in the blood (a correlation of 0.918 for lactate and Excess C0 was found). 2  Wyndham et a l . (1962) speculated that the onset of metabolic acidosis limited prolonged physical activity.  Their investigation demon-  strated that the "trained" individual had a higher tolerance to this anaerobic onset.  These ideas were novel, but the use of Huckabee's  (1958a) Excess Lactate concept in determining the start of anaerobiosis detracted from their study. Williams et a l . (1962) investigated the anaerobic threshold and heat stress.  Anaerobic activity appeared to increase at about 1.4 to  1.5 liters per minute of oxygen uptake above resting in the hot environment (about 95°F) and at about 1.6 to 2.2 liters per minute of oxygen uptake above resting in the comfortable environment (about 70°F).  This  suggests that the anaerobic metabolism mechanism is initiated faster at the higher temperature.  Excess lactate determined the onset of anaerobic  activity therefore the results must be interpreted with caution. In 1964, three studies from Karlman Wasserman's laboratory in Palo Alto, California, introduced the concept of the "Anaerobic Threshold" (AT).  The AT was originally defined by Wasserman et a l . (1964) as  ". . . the work level (oxygen uptake) above which the subject develops metabolic acidosis."  The AT and i t s relationship to varying levels of  work rate was investigated by Wasserman et a l . (1964).  Their findings  reiterated Wyndham et al.'s (1962) postulate, that activity may be prolonged for workloads at or below the AT. The continual observation of blood lactate may be inconvenient, so Naimark et a l . (1964) investigated respiratory exchange variables concomitant with lactate and bicarbonate changes.  When lactate and  14 bicarbonate were compared to the Respiratory Quotient and Excess C O 2 production (as similarly done by Issekutz & Rodahl [1961]) identical time of change was observed and a correlation (r) of 0.98 was calculated for excess C0 and bicarbonate concentration. 2  Trained athletes did not  exhibit an increase i n the Respiratory Quotient until a much higher work level (oxygen uptake) compared to that of the untrained, suggesting a delay in onset of anaerobic activity.  Their bike protocol for detecting  the AT proved very reliable when checked the following day. The third investigation (Wasserman & Mcllroy, 1964) utilized breathby-breath analysis of end-tidal gas concentrations to calculate the Respiratory Quotient to determine the AT for patients in a'hospital. monitoring procedures were administered cardiovascular deficiency.  AT  to determine some forms of  This provided an added measure of safety to  testing procedures not available under maximal stress test conditions. Wyndham et a l . (1965) also investigated the AT comparing normal and hospital patients.  Although excess lactate was used to determine the  point of onset of anaerobic metabolism, the relationship between maximum oxygen uptake and the AT was introduced.  The use of a 'percent' of  maximum oxygen uptake would allow comparisons of normal and cardiac patients, however, exclusion of work-load would limit: 1.  absolute improvement recognition after rehabilitation, and  2.  comparisons to other studies with subjects of.different maximal oxygen uptakes. The result of investigation of the AT and i t s application to  hospital patients was the review of muscular exercise analysis in normal and disease situations (Wasserman & Whipp, 1975).  Results of previous  investigation, especially relating to the AT, and i t s application to  15 patient diagnosis was  reviewed.  Bouhuys et a l . (1966) conducted an exhaustive analysis of-the relationships between blood lactate and excess C O 2 , respiratory quotient, blood pH and bicarbonate.  Although significant correlations (p < .001)  existed between lactate and a l l variables, the highest correlation was to excess C © 2 (r=.796).  As with a l l predictors, there were discrepan-  cies which led these researchers to conclude that the best index of metabolic acidosis i s blood lactate.  However, they do agree that the  respiratory quotient and excess C O 2 are good indicators of 'change' in blood lactate levels. A study relating AT and training was conducted by Williams et a l . (1967).  This investigation showed that daily exercise of four hours,  continuing from four to six weeks, increases the percent of maximum oxygen uptake at which anaerobiosis occurred (maximum oxygen uptake also increased).  Analysis of anaerobiosis consisted of excess lactate  determination therefore limiting interpretation of the results. The following year, Williams et a l . (1968) conducted a similar, but more extensive study on endurance trained males.  The results showed  that blood lactate and excess lactate levels were significantly different when determined after the sixth and sixtieth minute of exercise.  A  more important finding was the difference in the AT as predicted by lactate and excess lactate (this difference was significant according to the authors, but no confidence limits were given).  This again casts doubt  on the use of excess lactate as previously discussed. Wasserman et a l . (1967) investigated work intensity and duration of exercise on a bicycle ergometer related to the AT.  The concepts of  "pay-as-you-go" (aerobic) and "credit" (anaerobic) energy supply terms  16 relating to exercise could explain the different metabolic  responses.  Increased oxygen debt, as a result of increased.work intensity showed an increased debt proportion due to lactate oxygen equivalents.  At a sub-  threshold (moderate intensity as described by Wells et a l . [1957]) work intensity, activity could be maintained by a l l subjects for a minimum f i f t y minute period.  However, only four of ten subjects completed the  heavy work intensity (just above the AT) for the f i f t y minute period. No one completed the f i f t y minutes at the very heavy work load.  This  suggests that the AT i s the limit of aerobic work capacity and that activity above this work level requires an increasing amount of anaerobic energy supply.  The finite limits of energy production from the pyruvate:  lactate system and i t s feedback on aerobic energy supply (Wenger & Reed, 1976) appears to restrict prolonged activity. Nagle et a l . (1970) investigated lactate accumulation during runs of 64 to 89 percent of maximum oxygen uptake.  The untrained subjects  demonstrated higher blood lactate levels for the same work intensity (as also shown by Wasserman et a l . [1967]) as compared to the trained subjects.  This again suggests'that the AT may limit muscular activity as  a result of anaerobiosis. Costill (1970) was the f i r s t investigator to apply the AT to distance running. ' As might be predicted from previous studies, increased distance produced, decreased steady-state blood lactate levels.  The  limited energy supply of anaerobic metabolism and discomfort from tissue acidosis (Simonson, 1971) probably determines the optimal running speed for a given race and therefore blood lactate level. The relationship of work intensity above the AT to oxygen uptake time course to steady-state was investigated by Whipp and Wasserman  17 (1972).  A slower rise to oxygen uptake steady-state was observed in  a l l instances for work intensity above the AT.  It was also shown that  the trained individual had a higher level of work intensity before anaerobic energy supply started. More recently, Whipp et a l . (1979) utilized f i r s t order kinetics to demonstrate the oxygen uptake rise to steady-state at subthreshold exercise (time constant about 45 seconds).  However, depending on the  type of suprathreshold exercise (ramp., sinusoid, or square wave) a twocomponent rise to steady-state was only occasionally noted.  Discrep-  ancies between individuals suggested that a two-component or delayed rise is not necessary for suprathreshold activity. The concept of continued anaerobic activity and removal of lactate during submaximal exercise was reviewed by Hermansen and Stensvold (1972).  They failed to show lactate production for individuals at a  work intensity below 60-80 percent maximum oxygen uptake (normally termed the AT).  However, observations during exercise, where blood  lactate is produced, showed that the major area for lactate oxidation i s in skeletal muscle.  Graham (1978), in a comprehensive review of lac-  tate changes during exercise, showed that lactate may be produced in muscle but does not necessarily diffuse into the blood, thereby masking anaerobic activity.  This suggests lactate's continual oxidation in  muscle (active or inactive fibers). Wasserman et a l . (1973) summarized breath-by-breath non-invasive techniques for detecting the AT.  A most significant result of the  study was that for non-invasive AT determination, a one minute work increment rate was found to be optimal for showing the change in metabolism at the start of anaerobiosis.  This permits stress testing duration  18 to be kept at a minimum.  Reproducibility of the AT was shown to be  exact over one hour, four hour, one week and nine month intervals. Breath-by-breath analysis (Wasserman et a l . [1973]) also solved a major problem of AT detection.  Hyperventilation may obscure the AT  when observing expired volume or C O 2 .  Observation of end-tidal oxygen  and carbon dioxide eliminates the chance of misinterpretation. Koyal et a l . (1978) researched the feasibility of using oxygen pulse in determining the AT.  In a very restricted sample (maximum  oxygen uptake less than 3.75 liters per minute), they observed that the curvilinear plateauing of oxygen pulse coincided with the AT.  The  leveling of oxygen pulse iridicatesvimpending maximum oxygen uptake. An important change in testing technique of the AT was researched by Volkov et a l . (1975). of  He utilized the Excess  C O 2  ( E X C C O 2 )  concept  Issekutz and Rodahl (1961) to determine the AT during a speed  increase treadmill protocol. which the AT occurred.  The result was a running speed (  V T A M  )  a t  Volkov et a l . (1975) suggested that this AT  speed could be used as an index of running efficiency. Application of the AT to distance running was investigated by Weiser et a l . (1978), Sucec (1979) and Farrell et a l . (1979).  Sucec  (1979) used the AT, expressed as m i l l i l i t e r s oxygen per kilogram per minute, in a regression equation to predict one and two mile times (the other regression factors were percent body fat, running efficiency, oxygen debt, lean body weight, and maximum oxygen uptake).  Correlations  of 0.98 and 0.91 were calculated for the one and two mile times respeet ively. Farrell et a l . (1979) determined the running speed at which the "onset of plasma lactate accumulation" (OPLA) occurred.  This speed and  19 mean marathon speed were correlated significantly at r=0.98.  Mean  marathon pace was 0.28 miles per hour above the mean treadmill pace at OPLA.  Weiser et a l . (1978) also applied this concept and showed a  correlation of 0.92  (p < 0.001) for 3.2 kilometer race pace-and treadmill  speed at the AT. Wasserman's continued basic AT research (Wasserman et a l . 1975) demonstrated the involvement of the carotid body: in"metabolic responses - = during exercise. .  The carotid body appears to be the mediator for  ventilatory change for suprathreshold activity to maintain blood homeostasis.  This respiratory compensation is the major factor stabilizing  metabolic acidosis.  These findings also support the use of non-invasive  measurements for AT determination. Whipp's model (Whipp, 1977) of hyperpnea during exercise also used the AT as a division of exercise intensity for determining ventilatory control.  The carotid bodies seem to be the main mediators of control  for suprathreshold work intensity.  Subthreshold mediation of hyperpnea  seems to be much more complex (as reviewed by Whipp [1977.]). AT determination for three different modes of exercise was analyzed by Davis et a l . (1976).  Similar results for maximal oxygen uptake and  percent of maximum at the AT were obtained for treadmill walk/run exercise and cycling.  Significantly lower results were obtained in a l l  instances for arm cranking.  Their results also show that the AT is  reproducible for their three exercise protocols. Wiswell et a l . (1979) also investigated the relationship between the AT calculated for a bicycle ergometer and a treadmill.  For each  modality, oxygen uptake at the AT and the maximum oxygen uptake were significantly correlated, but the AT (expressed as a percent of maximum  20 oxygen uptake) for each exercise was significantly different (p :< 0.01). This suggests that, similar to maximal oxygen uptake (Str^mme et a l . , 1977;  McKay & Bannister, 1976), the AT may be exercise specific, as  suggested by Davis et a l . (1976).  However, Stamfor et a l . (1978) showed  that for one- versus two-legged cycling, the AT occurs at the same relative percent of maximum 0 uptake, though at a lower absolute work 2  load.  Maximum oxygen uptake was lower for one-legged cycling.  These  findings suggest that the AT and.maximum oxygen uptake may be influenced differently under the same conditions. MacDougall (1977) investigated the different training techniques of athletes and their AT.  His observation suggests that the longer  distance trained athletes have a higher AT.  He also illustrates a hypo-  thetical example of two identical athletes, except for different ATs, who run a marathon.  The higher AT individual would have a significant  advantage i n supplying aerobic energy requirements that could be equated to distance advantage i f constant stress to the cardiovascular system was maintained. Comparison of sprinters and endurance runners are described by Roberts et a l . (1979).  The sprinters exhibited:  1.  lower maximum oxygen uptake,  2.  lower AT (percent maximum oxygen uptake),  3.  lower oxygen uptake at the AT,  4. lower muscular capillarization, and 5.  higher percent fast twitch fiber composition.  These data suggest that the sprinters have lower endurance capacity, especially when considering the AT and capillarization.  However, com-  parison of the AT and muscle fiber composition by Green et a l . (1979)  21 showed no correlation between the two, thereby limiting conclusions about AT and muscle fiber composition.  No muscular sites were reported in  this abstract. Davis et a l . (1979a, b) investigated the effect of endurance cycle ergometry training on maximum oxygen uptake and the AT for middle-aged men.  Both significantly increased over the nine week program.  Again  this suggests increased AT with endurance type training. Weltman et a l . (1978) tested the use of the AT as a measure of submaximal fitness.  They observed that the individuals with a high  oxygen uptake at metabolic acidosis had: 1.  a faster rate to steady-state oxygen consumption,  2.  a lower steady-state oxygen uptake, and  3.  a faster return to resting values after exercise.  These findings are in agreement to those of Whipp and Wasserman (1972). The use of heart rate as a training index was investigated by. Katch et a l . (1978).  These researchers suggest that the use of the  relative percent of maximum heart rate concept is not a good training concept when considering whether or not an individual is producing lactate.  Individuals appear to have different heart rates corresponding  to their AT, therefore caution i s stressed until the AT has been determined . Patton et a l . (1979) also investigated heart rate and i t s relationship to the AT.  Their data suggest that heart rate does not vary  between individuals (trained or untrained) at their AT.  However, their  heart rates at the AT were higher compared to Wyndham et a l . (1965) and Wasserman et a l . (1973). Ivy et a l . (1979) examined the effect of substrate availability  22 on the AT.  Increased blood glucose had no effect on the AT.  Increased  blood free fatty acids significantly lowered the AT and absolute blood lactate.  This has definite implications in calculation of the AT and  Ivy suggests that standards should be adopted for AT determination. Pendergast et a l . (1979) compared performance on an arm crank exercise between kayakers and sedentaries.  Kayakers were observed to  have higher absolute ATs but not relative ATs (they also had higher maximum oxygen uptake).  Their methodology of determining the AT con-  sisted of separated exercises at different work loads for different time durations.  This resulted in an early.lactate release for some sub-  threshold exercises that lasted for :more than thirty seconds.  .They  suggest that this lactate release may be used with the AT to evaluate muscle training. Kindermann et a l . (1979) analyzed the aerobic-anaerobic transition with reference to the AT and blood lactate concentrations.  They suggest  three categories for this transition as a result of their investigations. This classification system includes: i.  The Aerobic Threshold (blood lactate less than 2 mmol/1, similar to the AT)  ii.  The Aerobic-anaerobic Transition (blood lactate 2 to 4 mmol/1)  iii.  The Anaerobic Threshold (greater than 4 mmol/1 blood lactate).  These investigators found that under conditions of the Aerobic Threshold, activity could be maintained format least two hours.  The Aerobic-anaerobic  Transition period'of activity could be maintained for one hour. conditions of the 'Anaerobic Threshold  1  For  only periods of exercise less  than an hour in duration could be maintained. The AT has been applied to many diverse testing situations.  The  majority seem to be concerned with very specific but unrelated areas of application.  .Research of the AT is needed to explain the conclusions  of these studies.  This would allow applied research to progress with  a sound knowledge base.  CHAPTER 3  METHODS AND  PROCEDURES  Subj e c t s Twenty male s u b j e c t s were s e l e c t e d f o r t h i s study from v o l u n t e e r s from the U n i v e r s i t y of B r i t i s h Columbia p a r t i c i p a n t s were s u b j e c t i v e l y chosen  student p o p u l a t i o n .  (from a n a l y s i s o f t h e i r  p a t t e r n s ) t o a c h i e v e a continuum of A n a e r o b i c T h r e s h o l d (AT)  The activity levels.  T e s t i n g Procedures The t e s t i n g format c o n s i s t e d o f two s e s s i o n s . week, h e i g h t , weight, a n a e r o b i c t h r e s h o l d speed consumption uptake  (VO2  (VO2  max)  D u r i n g the  first  (V,j,^). arid maximum oxygen  were determined f o r each s u b j e c t .  R e s t i n g oxygen  r e s t ) and r e c o v e r y oxygen uptake a f t e r the s e t t r e a d m i l l r u n  were determined d u r i n g the second week. VO2 protocol.  max  and V  were determined u s i n g a c o n t i n u o u s t r e a d m i l l  As a warm-up, each s u b j e c t walked on the t r e a d m i l l a t  m i l e s per hour f o r t e n minutes.  Subsequently, the t r e a d m i l l was  3.5 set  at 4.0 m i l e s per hour and then i n c r e a s e d o n e - h a l f m i l e per hour a t the end of each minute u n t i l v o l i t i o n a l Heart r a t e was  fatigue.  monitored by d i r e c t ECG u t i l i z i n g an A v i o n i c s 4000  c a r d i o g r a p h w i t h o s c i l l i s c o p e and ST d e p r e s s i o n computer and E x p i r e d gases were c o n t i n u o u s l y sampled b o l i c Measurement Cart  display.  and a n a l y z e d by a Beckman Meta-  (BMMC) i n t e r f a c e d i n t o a H e w l i t t Packard 3052A 24  25 Data Acquisition system for fifteen second determination of respiratory gas exchange variables. Maximum oxygen consumption was determined by averaging the highest four consecutive fifteen second oxygen uptake values.  The anaerobic  threshold speed was determined by examining excess C0 (Volkov et a l . , 1975).  2  elimination  The determination of the AT was consistent with  the definition by Wasserman et a l . (1964). subjects into a low and high V  In order to dichotomize;the  grouping a median speed (V^g^) f °  r  t n e  twenty V , scores was calculated as 7.25 miles per hour. TAM mll  Resting oxygen consumption was determined during a ten minute s i t ting period at the start of the week 2 assessment.  Each subject then  walked on the treadmill at 3.5 miles per hour for five minutes. the warm-up, each subject ran at  After  for ten minutes, during which time  expired gases were continuously monitored by the aforementioned systems. At the completion of the ten minute run, each subject sat until he either returned to his V0  rest (minimum time sitting was set at twenty  2  minutes) or had recovered for thirty minutes. Data Analysis The last four oxygen uptake values were averaged during the median speed run to determine time zero oxygen uptake for recovery.  This value,  plus each fifteen second value of oxygen uptake during recovery were f i t with a double exponential equation of the form: V0 where  V0  2  2  = a^e  -ait  -a t + ae + a3 2  2  = oxygen consumption at time 't'  a 1 = alactic linear parameter a±  = alactic non-linear parameter  '\ a2 = l a c t i c l i n e a r  26  parameter  0^2 = l a c t i c n o n - l i n e a r  parameter  a3 = asymptotic VO2 r e s t The U.B.C. Computing Centre's program P:3R utilized  f o r t h i s computation.  based on Henry  (Dixon & Brown, 1979)  I n i t i a l n o n - l i n e a r parameters were  and DeMoor (1950).  Each e q u a t i o n was  i n t e g r a t e d over t h i r t y minutes t o d e t e r m i n e . t o t a l  oxygen debt, a l a c t i c oxygen debt, and l a c t i c oxygen debt A).  was  Each i n d i v i d u a l ' s l a c t i c oxygen debt was  oxygen debt to g i v e the l a c t i c : a l a c t i c Time t o r e s t i n g oxygen uptake  ratio  (see Appendix  d i v i d e d by the a l a c t i c  (ratio  debt).  ( r e c o v e r y time) a f t e r e x e r c i s e  was  determined by o b s e r v a t i o n of f o u r c o n s e c u t i v e r e c o v e r y r e a d i n g s e q u i v a l e n t t o each s u b j e c t ' s i n i t i a l  r e s t i n g oxygen uptake.  T o t a l oxygen debt and r a t i o debt f o r s u b j e c t s w i t h a n a e r o b i c t h r e s h o l d speeds above (group H-V  ) and below (group  L-  Vrj,^j)  the  median speed were a n a l y z e d f o r d i f f e r e n c e between means by the U.B.C. Computing Centre program MULTIVAR ( F i n n , 1977). m u l t i v a r i a t e F ( H o t e l l i n g T ) was 2  where s i g n i f i c a n c e o c c u r r e d .  Significant  (p <  .05)  f o l l o w e d by u n i v a r i a t e F to determine  A l a c t i c oxygen debt and l a c t i c  oxygen  debt were s i m i l a r l y a n a l y z e d . Correlation coefficients  ( r ) were c a l c u l a t e d f o r the oxygen debt  v a r i a b l e s and V_.,„ f o r the H-V_._, and L-V_... groups by the U.B.C. ComputTAM TAM TAM J  i n g Centre program  SIMCORT (Le, 1979).  f o r s i g n i f i c a n c e from z e r o .  Each c o e f f i c i e n t was a n a l y z e d  CHAPTER 4 RESULTS AND DISCUSSION Results The twenty subjects were tested, as per the schedule of week 1. One subject (RR) was added to the twenty after the median speed determination to attempt to further complete the AT continuum between 5 and 6.5 miles per hour.  Their results are summarized in Table 1.  vidual anaerobic threshold curves and determination of V ^  Indi-  appear in  Appendix B. Oxygen uptake curves during recovery in week 2 appear in Appendix C.  Subject JO experienced coughing during the recovery period there-  fore his recovery data are not included in this study. Double exponential equations calculated for each subject appear in Table 2. These functions are represented as a continuous line on each recovery curve in Appendix C. The computer program P:3R did not produce a resting asymptotic value for subjects ML, BV, RF and DW.  This appears  to be due to insufficient data points at base line oxygen consumption. Therefore, a lower limit for this baseline was set as 0.225 liters of oxygen per minute.  This value appears to be a limit for V0 rest. 2  The  resulting equations are included in Table 2 and Table 3. The computer program could not i n i t i a l l y calculate a double exponential equation for subject GS.  Observation of his recovery curve  data points showed that three points (at 1.0, 1.24 and 1.50 minutes) seem to deviate from the regular pattern seen in a l l subjects. 27  Removal  TABLE 1 Individual Subject Characteristics  Subject  CN ML RR TB DG DD BV GS DA RF HB AB AO GT DH JL JO DM DW SP JH  Age (years)  Height (cm)  Weight (kg)  TAM , (mph) V  , ™ 2 max (ml/kg/min.)  22 25 24 21 24 24 21 22 23 24 20 21 26 24 21 22 24 23 24 21 23  175.6 174.0 188.4 179.5 174.0 179.1 184.5 188.7 175.6 166.1 170.2 181.1 174.9 176.7 179.3 177.5 180.8 174.9 178.6 182.1 176.4  77.9 82.7 95.5 71.1 60.5 75.5 88.5 99.4 77.9 62.5 65.0 74.5 63:4 78.4 69.3 67.1 79.0 63.3 79.8 68.0 71.0  4.0 5.0 6.0 6.5 6.5 6.5 6.5 6.5 7.0 7.0 7.0 7.5 7.5 7.5 8.0 8.0 8.0 8.5 8.5 10.5 11.0  42.6 46.6 52.2 46.3 59.2 49.1 67.9 46.6 58.5 55.4 50.0 62.5 58.6 57.0 57.4 64.8 69.2 76.2 52.0 68.0 72.7  X  22.8  178.0  74.8  7.3  57.8  SD  1.60  5.37  10.57  1.5  9.4  29 TABLE 2 Individual Double Exponential Equations for Subjects in Group L-V^^.  Exponential Equation  V0 rest (1/min.)  + 1.221e"*  + .441  .317  2  ^TAM  Subject  4.0  CN  3.269e"  5.0  ML  2.923e~ '  6.0  RR  6.5  1,076t  168t  140t  +  .181e"'  P97t  + .225  .273  3.472e"  1,169t  +  .151e""  075t  + .265  .320  TB  2.453e"  1,510t  +  .391e""  206t  + .309  .280  6.5  DG  2.417e"  1-411t  +  .407e~"  185t  + .253  .240  6.5  DD  2.985e '°  90t  +  .285e""  103t  + .278  .291  6.5  BV  2.949e"  1-020t  +  .210e"'  038t  + .225  .369  6.5  GS  -2 4^41- 33912.595e ^ - ^ + i.056e + .310  .291  7.0  DA  3.228e '  7.0  RF  7.0  HB  1  _1  H  z  168t  +  .132e~"  079t  + .408  .349  2.038e"  1,264t  +  .063e""  160t  + .225  .223  1.921e"  1,787t  +  .727e~*  406t  + .277  .259  _1  30 TABLE 3 Individual Double Exponential Equations for Subjects in Group H-V „ ^ TAM mA  V0 rest (1/min.) 2  TAM  V  S u D  J  e c t  Exponential Equation  7.5  AB  2.342e ' _1  169t  7.5  AO  1.181e"  6,690t  7.5  GT  8.0  DH  8.0  .609e" ' 9  1.992e"  1,393t  2.713e" ' 1  J L  562t  160t  +  .106e~"°  + 1.143e~' + 1.941e~ + +  32t  + .268  .312  751t  + .289  .286  1-153t  .476e"' .165e"-  419t  068t  8.5  DM  1.515e"  4,083t  +  .953e"-  8.5  DW  3.189e"  1,236t  +  .137e""°  10.5  SP  11.0  JH  2.052e " + -? Q17t1.729e + _1  820t  + .272 + .273  + .217  .246 .230 .230  + .210  .211  + .225  .301  .186t"' + .302 - 559t.611e ' + .278  .318  709t  23t  225t  3 3 ) , z  .271  31 of the data points at 1.25 and 1.50 minutes allowed for proper curve f i t ting.  Therefore, his results are calculated without these two data  points. Each equation was integrated over thirty minutes to determine alactic, lactic, and total oxygen debts (Appendix A).  The ratio of  lactic to alactic debt (debt ratio) was then calculated for each subject. These results appear in Table 4 and Table 5.  Graphs of these results  versus individual anaerobic threshold speeds (  V  )  a r e  T A M  represented in  Figures 1 to 4. It was not possible in this study to determine exact recovery time. The fluctuation of the fifteen second values for oxygen uptake (Appendix C) prevented the attainment of four consecutive equivalent values of resting oxygen uptake.  The asymptotic value calculated by the computer  program P:3R never exactly matched the resting value determined, therefore comparison of these two values would not be valid. hypotheses 5, 10, and 15 were not tested.  Thus,,  TABLE 4 Individual Alactic, Lactic, Total and Ratio Oxygen Debts for Group l ^ ^ -  Lactic Debt (liters)  Ratio Debt (Lactic/ Alactic)  Total Debt (liters)  TAM  Subject  Alactic Debt (liters)  4.0  CN  3.04  7.22  2.38  10.26  5.0  ML  2.56  1.76  0.69  4.33  6.0  RR  2.97  1.80  0.61  4.77  6.5  TB  1.62  1.89  1.17.  3.52  6.5  DG  1.71  2.19  1.28  3.90  6.5  DD  2.74  2.64  0.97  5.38  6.5  BV  2.89  3.76  1.30  6.65  6.5  GS  1.07  3.12  3.92  4.19  7.0  DA  2.76  1.51  0.55  4.28  7.0  RF  2.19  0.39  0.24  2.00  7.0  HB  1.07  1.79  1.66  2.87  V  33  TABLE 5 Individual Alactic, Lactic, Total and Ratio Oxygen Debts for Group H-V .  Lactic Debt (liters)  Ratio Debt (Lactic/ Alactic)  TAM  Subject  Alactic Debt (liters)  7.5  AB  2.00  2.04  1.02  4.05  7.5  AO  0.18  1.52  8.44  1.70  7.5  GT  0.06  1.68  27.18  1.75  8.0  DH  1.43  1.14  0.79  2.57  8.0  JL  2.34  2.11  0.90  4.45  8.5  DM  0.37  1.34  3.81  1.72  8.5  DW  2.58  2.97  1.15  5.55  10.5  SP  1.13  0.83  0.73  1.95  11.0  JH  0.59  1.09  1.85  1.69  V  Total Debt (liters)  10 -J Total Debt (liters)  O  ® 2 scores  9  8  Individual scores  J  7 -\  6  J  5  J  4  J  3  J  o O  o  o o o  o  o  o  o  2  J  O O  -T  MED Figure 1.  Graph of Total Debt and Individual V, TAM  T 8  n  9  1  r  10  1  11 V (mph) TAM  O  Individual  Scores  4 H Alactic Debt (liters) O  3 H  O O  O O  O  2 H  o  8 i  o  H  -7?-  o  o  o o 10  F i g u r e 2.  V MED Graph of A l a c t i c Debt and I n d i v i d u a l V TAM  11 TAM  (mph)  O  7 i  Individual  Scores  O  Lactic Debt (liters) 6  5  H  4 1  o o  2 H  o  o  o o o  1 H  o o  o  8  o  r 5 Graph of L a c t i c Debt and I n d i v i d u a l  o  o  O  O  10 MED  Figure 3.  o  V, TAM  11 V  TAM  (mph)  O  I n d i v i d u a l Scores  ®  2 Scores  4 Scores  Ratio Debt  30 H o 25 -\ 20 H 15 10 H  11 0  o  O  77^  o  o  ©  8  o —I—  MED F i g u r e 4.  Graph of I n d i v i d u a l R a t i o Debt and V  o  O  10 TAM  O  (mph) CO  TAM  38 Individual oxygen debt values were averaged for those subjects in group L-V  and in group H-V  .  These values appear in Table 6.  TABLE 6 Means of Total, Alactic, Lactic and Ratio Debts for Groups L - V and H-V TAM  Group L-V..,, TAM H-V„ TAM 1U  Total Debt Mean SD  Ratio Debt Mean SD  Alactic Debt Mean SD  Lactic Debt Mean SD  4.74  2.33  1.34  1.04  2.19  0.77  2.55  1.78  2.83  1.43  5.10  8.65  1.19  0.95  1.64  0.65  The difference between means for total debt and ratio debt for the two groups show a significant T  2  statistic (p < .05).  Univariate t  values subsequently calculated for each dependent variable show this significance to be attributable to the difference between total debt means only (p <-:.01).  Despite the large difference in means for ratio  debt, significance was not obtained due to the large variance in group ^~^TAM (  D r o u  8 t about by subjects AO and GT). n  These results are sum-  marized in Table 7. These results indicate rejection of the null hypothesis and therefore acceptance of hypothesis 1;  that subjects in group H-V_^ accumu-  late a lower oxygen debt compared to those subjects in group ^-V_^. However, the statistical calculation does not indicate acceptance of hypothesis 4; that there i s a difference between subjects i n groups H-V .„, and L-V „ in the calculation of a debt ratio (lactic debt/ TAM TAM m  alactic debt).  mA  39 TABLE 7 Multivariate Analysis of Dependent Variables Total Debt and Ratio Debt Dependent Variable Total Debt Ratio Debt  T  2  4.84  .022  9.78  .006  1.43  .248  The difference between means for alactic and lactic debt for the two groups also exhibit a significant Hotelling T  (p < .01).  Univar-  iate t values calculated for each dependent variable show this s i g n i f i cance to be attributable to both the difference between alactic debt means (p < .01) and lactic debt means (p < .05).  These results are  summarized i n Table 8. These results indicate acceptance of hypotheses 2 and 3;  subjects  in group H-V,., accumulate lower lactic and alactic debts than subjects ° TAM m  in group L-V  J  TAM  . TABLE 8  Multivariate Analysis of Dependent Variables Alactic and Lactic Debts Dependent Variable Alactic Debt Lactic Debt  T  2  6.02  p<  .01  t  p<  12.1  .003  4.6  .047  Pearson correlation coefficients calculated for groups H-V  and  40 L-V-.,,, for V_.., with total, ratio, alactic, and lactic debts appear in TAM TAM r r  Tables 9 and 10. TABLE 9 Correlation Coefficients for Subjects in Group L-V, TAM  v TAM  Total Debt  Ratio Debt  Alactic Debt  -.767*  -.210  -.506  Lactic Debt -.729*  *Significantly different from zero at p < .05. The results in Table 9 indicate acceptance of hypotheses 6 and 8; subjects i n group L-V^^ have a negative linear relationship between total and lactic debts and V .„. These results do not indicate accepTAM tance of hypotheses 7 and 9, therefore a decreasing function between m  ratio and alactic debts and V_._- is not evident. TAM  It i s apparent from  observation of Figures 2 and 4, that both alactic and ratio debts are unrelated to V_,^. TABLE 10 Correlation Coefficients for Subjects i n Group H-V Total Debt v TAM  -.279  Ratio Debt -.375  Alactic Debt -.478  Lactic Debt -.098  The probability that the correlation coefficients for group  H _ V T A M  (Table 10) are different from zero are as.follows (Rohlf & Sokal, 1969): 1. Total debt  (r = -.279)  .4 < p < .5  2. Ratio debt  (r = -.375)  .2 < p < .4  41 3.  Alactic debt (r = -.478)  .1 < p < .2  4.  Lactic debt  .5 < p < .9  (r = -.098)  These probabilities suggest that there is not a linear relationship between total, ratio, and lactic debts and V , . The correlation coefTAM ficient for alactic debt and V_... i s very close to the confidence limit TAM m  w  J  (p < .05) to reject the null hypothesis, suggesting a linear relationship.  However, as the correlation coefficient only accounts for 23%  of the variance of the sample, there i s strong evidence to suggest that there i s also no linear relationship between alactic debt and  As  a result, the null hypothesis is not rejected and remains as stated in hypotheses 12, 13, and 14; there is no decreasing.^function between total, alactic, and lactic debts and V_... in group H-V_... ' TAM TAM A  Discussion The computation of double exponential equations to describe recovery oxygen consumption is consistent with the theory of decreasing payment (exponential decay) of alactic and lactic debts.  The equations  calculated (Tables 2 and 3) are similar to those reported by Henry and DeMoor (1950).  The non-linear (a ) and linear ( a ^ parameters are of  the same order of magnitude, except for those subjects in this study with small fast recovery component (subjects AO, GT and DM).  Total  debt, for a l l individuals, falls within limits of previous studies (Hill et a l . , 1924;  Margaria et a l . , 1933;  Henry & DeMoor, 1950;  Knuttgen,  1962). Comparison of computer calculated asymptotic V0 determined V0  2  rest values reveal similar results.  2  rest values and  However, subjects  CN, BV, and DW exhibit differences greater than sixty m i l l i l i t e r s of  42 oxygen per minute.  Subject CN experienced difficulty i n completing the  10-minute run, so that an elevated asymptotic V O 2 rest i s normal (Hill et a l . , 1924).  The large variation i n the resting data of subjects BV  and CN may be attributed to the fact that i n i t i a l resting oxygen consumption values are not indicative of their true resting state, and secondly, the lower limit of 0.225 l i t e r s of oxygen per minute set for the computer may be low. The resulting integration of the equations to determine contributions to the total debt in this investigation, indicates that the processes involved are not as simple as originally proposed by Margaria et a l . (1933).  Calculation of debt ratios from the data reported by Mar-  garia et a l . (1933), Henry and DeMoor (1950), and Knuttgen (1970), show a range from 0 (mildest exercise) to about 2.  These ratios do not  compare with the ratios of 8.44, 27.18, and 3.81 of subjects AO, GT and DM respectively.  Subjects in H-V_,^ who were working at a mild inten-  sity of exercise did not exhibit ratios of 0. At these low intensities of exercise, Berg (1947) argues that the recovery processes are better explained by a single exponential equation, thus supporting H i l l et a l . (1924) and Margaria et a l . (1933).  If this  is true, single exponential equations should better describe the recovery curves i n group H-V^^. H-V  Post hoc analysis of recovery curves i n group  show a mean residual sum of squares of .79 (SD=-56) for a single  exponential curve f i t (computer program P:3R [Dixon & Brown, 1979]).  The  double exponential curves calculate a mean residual sum of squares of .68 (SD=.37).  A correlated t-test of significance show these means to be  significantly different at a level of .05.  This suggests that the double  exponential equation better describes recovery oxygen data points i n group  This result  s u p p o r t s e v i d e n c e by Knuttgen (1970), t h a t t h i s slow  r e c o v e r y phase does e x i s t .  T h i s phase may  be r e s p o n s i b l e f o r a r e t u r n  of ". . . i n t e r r e l a t e d m e t a b o l i c , t h e r m a l , e l e c t o l y t i c and  hormonal  changes which the body undergoes d u r i n g e x e r c i s e . . . " t o n o n - e x e r c i s e levels.  P i i p e r and S p i l l e r  (1970) a l s o observed t h i s slow phase i n  i n t a c t dog gastrocnemius muscle s t i m u l a t e d t o do e x e r c i s e t h a t d i d not produce blood l a c t a t e i n excess of r e s t i n g v a l u e s . similar conclusion i n their D u r i n g oxygen d e f i c i t  They came to a  investigation. f o r m a t i o n , b o t h a n a e r o b i c L g l y c o l y s i s and  s t o r e d h i g h energy phosphates c o u l d be the energy s o u r c e s . repayment  phase  D u r i n g the  (oxygen d e b t ) , t h e s e energy s o u r c e s would be r e p l e n i s h e d  at a r a t e dependent on t h e p h y s i o l o g i c a l make-up and the t r a i n i n g of the i n d i v i d u a l . debt r a t i o  level  T h i s might account f o r some of the v a r i a b i l i t y i n  scores.  R e s u l t s of t h i s study i n d i c a t e t h a t r e c o v e r y oxygen consumption i s adequately d e s c r i b e d by a double e x p o n e n t i a l e q u a t i o n .  However, t o  r e f e r to these two phases as t r a d i t i o n a l l a c t i c and a l a c t i c may  be q u e s t i o n a b l e .  components  The r e m a i n i n g d i s c u s s i o n w i l l use a l a c t i c  and  l a c t i c terms to r e f e r t o the f a s t and slow r e c o v e r y phases r e s p e c t i v e l y . The r e l a t i o n s h i p between a n a e r o b i c t h r e s h o l d speed (V^,^) and r e c o v e r y oxygen consumption i s demonstrated i n the r e s u l t s of t h e hypotheses t e s t e d .  The s i g n i f i c a n t d i f f e r e n c e f o r t o t a l debt means f o r the  two groups i s c o n s i s t e n t w i t h o b s e r v a t i o n s of Wasserman et a l . (1965) and Wasserman et a l . (1967).  The n o n - s i g n i f i c a n t  linear  relationship  (r=-.279) c a l c u l a t e d f o r group H-V_^. suggests t h a t r u n n i n g speeds below V_  A M  result  i n a r e l a t i v e l y s t a b l e oxygen debt.  The  significant  44 negative linear relationship (r=-. 767) calculated for group ^-V ^ indicates that running at increasing speeds above V  results in  JL Jr_C*i  increasing total debt. The compartmentalization of total debt, into alactic and lactic debts, also produces a significant difference between the two debt components of each group (Tables 7 and 8). ;  The only significant negative  correlation calculated was for lactic debt and V_„, for group L-V TAM TAM  < w  s  (r=-.729).  v  These results suggest that the increasing total debt for  group L-V,-,^ i s a result of increasing lactic debt.  This may be a t t r i -  butable to increased blood lactate levels as suggested by Margaria et a l . (1933), Henry and DeMOor (1950), Wasserman et al. (1967), and Knuttgen (1970). Linear relationships were not observed between alactic debt and V„.„ in group L-V_.„ or between alactic and lactic debts and V .„ in TAM TAM TAM r  group H-Vrj,^.  m  Therefore, the relatively stable alactic component i n  each group, and the relatively stable lactic component i n group ^-V suggests the significant difference between debts i s a result of running above or below V . , „ . TAM  This implies that V_,„ is also c r i t i c a l in deterTAM r  mining alactic and lactic debt components for a constant running speed. Oxygen debt may be used as a criterion of fatigue and therefore used as a measure of work intensity (Simonson, 1971).  The relationship  between oxygen debt and V_, in this study shows that V ..„ is useful in TAM TAM J b  w  determining primarily aerobic work intensity.  m  Its relationship to  increased anaerobic activity and resultant metabolic acidosis through increased utilization (recruitment) of fast twitch muscle fibers and/or increased anaerobic activity in slow twitch muscle fibers, may be very important i n determining running speed for a given distance.  45 This relationship has recently been investigated by Weiser et a l . (1978), Farrell et a l . (1979), and Sucec (1979).  These investigators  illustrated very significant correlations between onset of anaerobic metabolism and race time.  This gives strong evidence to the role of  V „ , in determining efficient running speed for a given distance. TAM o r . o m  In conclusion, the anaerobic threshold speed (V  ) appears to be  the c r i t i c a l speed in determining total oxygen debt in a running exercise above V „ . m<  TAM  V_„, also appears to be c r i t i c a l in determining the TAM  *^  size of the debt components, alactic and lactic.  6  Further investigation  is needed to clarify the mechanisms involved in determining the total debt and i t s components.  Once this i s accomplished, the effect of  training techniques on V_,_, and the onset of anaerobic metabolism w i l l ^ TAM b  be elucidated.  CHAPTER 5 SUMMARY AND CONCLUSIONS Work intensity variation has been shown to change oxygen debt after exercise (Hill et a l . , 1924;  Wasserman et a l . , 1965).  Knuttgen (1962)  noticed a " c r i t i c a l level" of work intensity which seemed to drastically affect oxygen debt size.  This level of work intensity i s associated  with the onset of metabolic acidosis termed the Anaerobic Threshold (Wasserman et a l . , 1964).  This study attempted to clarify the relation-  ship between anaerobic threshold and oxygen debt after a set treadmill run. Twenty male university students were chosen to achieve a continuum of anaerobic threshold levels.  Week 1 testing sessions determined each  subject's anaerobic threshold running speed (V-^) using a continuous treadmill protocol. VC0  ^AM  (calculated to excess C0  2  a l . , 1975).  ^xAM  s c o r e s  identified by analyzing expired V0  W a s  r a n  2  2  and  values) every fifteen seconds (Volkov et  8 d from 4.0 miles per hour to 11.0 miles per e  hour. During the second testing session, each subject ran at the predetermined median speed of 7.25 miles per hour for ten minutes, which separated the subjects into two groups: 1.  Group H-V  —those subjects running below their  2.  Group L-V_. —those w  and  subjects running above their V_... r  Each subject's expired gases during the exercise and recovery stages were analyzed every fifteen seconds by a Beckman Metabolic Measurement Cart 46  47 interfaced into a Hewlitt Packard 3052A data acquisition system. Each individual's recovery oxygen consumption data were described with a double exponential equation by a computer.  The resulting equation  was integrated to determine total oxygen debt and i t s alactic and lactic components.  The ratio of lactic/alactic debts was also calculated.  Analysis for a significance between means (Hotelling T ) for groups 2  H-V .„ and L-V , for the debt variables revealed significant (p < .05) TAM TAM m  m  w  &  r  differences for total, alactic, and lactic oxygen debts. was evident for the ratio of lactic/alactic debt. L-VTAM  No difference  The subjects in group  had larger oxygen debts and debt.components compared with those sub-  jects in group H-V,.,.. TAM &  m  r  These results suggest that V ,„ is c r i t i c a l in TAM m  determining the oxygen debt after a run. Analysis for a linear relationship between V-^j and oxygen debt for each group showed total oxygen debt (r=-.767), and lactic debt (r=-.729) to be significantly (p < .05) linearly related to V only.  in group  L -  V  T A M  This suggests that the faster the running speed i s above an  individual's V ,„, the larger the total debt. TAM &  m  a result of the increasing lactic debt.  This increasing debt is •  This was not evident in group  " TAMThe accumulation of an oxygen debt is indicative of fatigue  H  V  (Simonson, 1971). '  The fact that V_._, is a c r i t i c a l factor in determining TAM  oxygen debt suggests that work above this point results in the onset of metabolic acidosis, which limits the optimal running speed for a given distance.  These findings support the evidence by Farrell et a l . (1979)  and Weiser et a l . (1978) that the onset of metabolic acidosis is related to average race running speed.  Further investigation should determine  the mechanisms by which V_,„,, is determined and changed. TAM 3  48  References Berg, W. E. Individual differences in respiratory gas exchange during recovery from moderate exercise. Amer. J. Physiol., 149: 597-610, 1947. Bouhuys, A., Pool, J., Binkhorst, R. A., & van Leeuwen, P. Metabolic acidosis of exercise in healthy .males-.. J. Appl. Physiol. , 21_(3) : 1040-1046, 1966. Cerretelli,'.-P..., Ambrosoli, G., & Fumagalli, M. Anaerobic recovery in man. Eur. J. Appl. Physiol., 34: 141-148, 1975. Cerretelli, P., diPrampero, P. E., & Piiper, J. Energy balance of anaerobic work in the dog gastrocnemius muscle. Amer. J. Physiol., 217(2): 581-585, 1969. C o s t i l l , D. L. Metabolic responses during distance running. Physiol., 28(3): 251-255, 1970.  J. Appl.  Davis, J. A., Frank, M. H., Whipp, B. J., & Wasserman, K. 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Integration of an  exponential function i s determined by the formula: t Jr ae- c t , dt = a -a Q  -at  a -a  Integration of subject DH's equation . -1.393t , _ ._, -0.419t 1.992e + 0.476e o o 0  over thirty minutes would be: .30  Alactic debt = / 1.992e 1  -1.393t  0  1.992 dt = -1.393  -1.393(30) 6  1.992 -1.393  = 1.43 Lactic debt  /  30  -0.419t dt = 0.476e  ' 0  0.476 -0.419(30) -0.419 e  1.14 Total debt  = Alactic debt + Lactic debt = 1.43 + 1.14 = 2.57  Debt ratio  = Lactic debt * Alactic debt = 1.14 * 1.43 = 0.79  0.476 -0.419  APPENDIX B  Anaerobic Threshold Curves to Determine Individual V  56  EcxCO-. (ml/kg/min)  26 H 24  20  16-H 12 8H  AH i  4  |  5 V  i  |  6  i  1 7  T A M  F i g u r e 5.  AT curve subject CN  1  1 8  r-  1  1  9  10 speed (mph)  1  ExcC0  2  (ml/kg/min) 20.  18  16  14  12 _J  10  8  J  ~i 10 TAM Figure 6.  AT curve subject ML  —:—r 11 Speed (mph) n  ExcC0  2  (ml/kg/min) 28 242016X  128-  ~~[  9 TAM Figure 7. AT curve subject RR  1  1  10 Speed (mph)  ExcCO  20 "  (ml/kg/min) 18 ~ 16  -  14  -  12  -  10  8 6 . 4  -  2  -  10  Figure 8.  AT curve subject TB  TAM  Speed (mph)  ExcC0  2  (ml/kg/min) 18-  16-  14-  12-  10-  8-  6-  4-  0 4  5  6  7 TAM  Figure 9.  AT curve subject DG  8  9  10  11  Speed (mph)  62  4= D.  r- o  e  T3  ii  CO  00  o  n01 XI  3  W  01 >  3 5 <u u  «-N CO C CN  CM 00 O _! CJ —  a x  i-i  S  -aCM  o  CN  -]—i vO  1 CM  1  1 00  1  1 -3-  3  r-  oo  O  ExcC0„  10  (ml/kg/mln) 8 -  7 -  6 5 -  2 -J  •  *  1 TAM  Figure 11. AT curve subject BV  1  9  I  10 Speed (mph)  ExcC0 (ml/kg/min) 2  32 2824201612 -  4  5  6  7 V  Figure 12. AT curve subject GS  T A M  8  9  10 Speed (mph)  ExcCO„  20  18 H (ml/kg/mln) 16  H  14 12-^ 10 J  8-J  4J 2A  T 8 TAM Figure 13. AT curve subject DA  —'  1  10 Speed (mph)  ExcCO„  9-1 8"  (ml/kg/min)  7-  6543H  2H H  10 TAM  Figure 14. AT curve subject RF  Speed (mph)  ExcC0  2  (ml/kg/min) 28  H  24  H  20 16  12  H  8-J  4-^  ">  r 5  Figure 15. AT curve subject HB  10 TAM  11 Speed (mph)  ExcCO(ml/kg/min) 2826242016 H 12 H 8H  4 H T 5  n  r 6  Figure 16. AT curve subject AB  10  11 12 Speed (mph) ON CO  ExcC0  2  (ml/kg/min)  -i  4  1  5  Figure 17.  r  1 6  '  1 7  r  V, TAM  8  1  AT curve subject AO  i—•—r —r —r - 1  11  - 1  12 Speed 13 (mph)-  ExcC0  2  (ml/kg/min) 28-  24  H  20 H 16-4 12-4  4-J  i i « i—i—i—i—r 4  5  6  7  8 TAM Figure 18. AT curve subject GT  10  ~~i— —r 1  11  12  13 Speed (mph)  20 ExcCO„ (ml/kg/min) 1614 H 12  io H  6H 4H 2H  —r 10 Figure 19. AT curve subject DH  TAM  -•  n —  n  Speed (mph)  ExcC02 (ml/kg/min) 18 16 14 H 12 H 10 -1  8H  T  10 TAM Figure 20. AT curve subject JL  T  11 Speed (mph)  73  Q. e  o>  Cu CO  o  o  1—  01 J3 03  01 •> U 3 U  5  01 3 00  u  00 — / > >_> •H C .-I  o  eg  o  u  00  20 4 ExcCO„ 18 -| (ml/kg/min) 16 -1 14 12 _ 10 _ 8 J  4J • • •  i— —i— —i— —r 1  6  1  1  7  8  9  V Figure 22. AT curve subject TAM DM  11  12  I  13  1  1  14 Speed (mph)  ExcC0  2  (ml/kg/min) 32 28 24 20 16 -\ 12  4 H  10 TAM Figure 23. AT curve subject DW  11 Speed (mph)  36 H ExcCO32 H (ml/kg/min) 2824201612-J 8 J  4-4 0 4  1  5  1  I ' 1 6  7  1  I  1  8  1 ' 1 9  10  —!—•—|—i—|—i—|  >~  , 11 TAM  Figure 24. AT curve subject SP  12  13  14  15 Speed (mph)  ExcC02  361 32 "i  (ml/kg/min) 2824201612••-  1—' 7  1— 8  1  1 9  •  ' — I — ' — I — — I — ' — I ~I— r 1  10  11  12  VTAM Figure 25. AT curve subject DH  13  14  1  1  r  ~  15 Speed (mph)  APPENDIX C Individual Oxygen Recovery Curves  78  Figure 26.  Recovery curve subject CN  F i g u r e 27.  Recovery curve s u b j e c t ML  Figure 29.  Recovery curve subject TB  00  5H  Time (min) Figure 31.  Recovery curve subject DD  Figure 32.  Recovery curve subject BV  Figure 34. Recovery curve subject DA  5-4  vo  2  (1/min)  4-4  3-4  2J.I  ~i  8 Figure 35.  10  12  Recovery curve subject RF  14  16  18  20  r  1  r  1  r  22  24  26  28 Time  30 (min)  CO CO  Figure 37. Recovery curve subject AB  5H  3-J  1 8  10  12  Figure 38. Recovery curve subject AO  14  16  18  20  22  . 24  26  1  r  28 30 Time (min)  Figure 39.  Recovery curve subject GT  Figure 40.  Recovery .curve subject DH  5J  o  i  0  1 2  Figure 41.  1 4  1——i 6  8  1 10  1——i 12  Recovery curve subject JL  14  1 16  r 18  1 20  1 22  1 24  1 26  1 28 Time  r 30 (min)  Figure 42.  Recovery curve subject DM  5 -A  Time (min) Figure 44.  Recovery curve subject SP  5-]  Figure 45.  Recovery curve subject JH  Time (min)  

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