ENERGY EXPENDITURE AND FATIGUE

EXERCISE PHYSIOLOGY AND EXERCISE PRESCRIPTION

Measuring Energy Expenditure

Direct calorimetry measures the body’s heat production to
estimate energy expenditure

Indirect calorimetry calculates energy expenditure from the
ratio of CO2 produced to O2 consumed
Measuring Energy Expenditure:
Direct Calorimetry

Substrate metabolism efficiency

40% of substrate energy
ATP

60% of substrate energy
heat

Heat production increases with energy production

Can be measured in a calorimeter

Water flows through walls

Body temperature increases water temperature
Figure 5.1
Measuring Energy Expenditure:
Direct Calorimetry

Advantage

Accurate over time

Good for resting metabolic measurements

Disavantage

Expensive, slow

Exercise equipment adds extra heat

Sweat creates errors in measurements

Not practical or accurate for exercise
Measuring Energy Expenditure:
Indirect Calorimetry

Estimates total body energy expenditure based on O2

used, CO2 produced

Measures respiratory gas concentrations

Only accurate for steady-state oxidative metabolism

Older methods of analysis accurate but slow

New methods faster but expensive

Douglas Bag
Measuring Energy Expenditure:
O2 and CO2 Measurements

VO2: volume of O2 consumed per minute

Rate of O2 consumption

Volume of inspired O2 − volume of expired O2

VCO2: volume of CO2 produced per minute

Rate of CO2 production

Volume of expired CO2 − volume of inspired CO2
Figure 5.2
Calculating Oxygen Consumption and
Carbon Dioxide Production
Calculating VO2 and VCO2 requires:
– Volume of air inspired
. (VI) .
– Volume of air expired (VE) .
– Fraction of O2 in the inspired
. air (FIO2)
– Fraction of CO2 in the inspired air (FICO2)
– Fraction of O2 in the expired air (FEO2)
– Fraction of CO2 in the expired air (FECO2)
Calculating O2 Consumption and CO2
Production (L/min)

. . .
VO2 = (VI x FIO2) – (VE x FEO2)

. . .
VCO2 = (VE x FECO2) – (VI x FICO2)
Respiratory Exchange Ratio

.
• The ratio between
. CO2 released (VCO2) and oxygen
consumed. (VO2) .
• RER = VCO2 / VO2
• The RER value at rest is usually 0.78 to 0.80
Measuring Energy Expenditure:
Indirect Calorimetry Limitations

CO2 production may not = CO2 exhalation

RER inaccurate for protein oxidation

RER near 1.0 may be inaccurate when lactate buildup 

CO2 exhalation

Gluconeogenesis produces RER <0.70

Measuring Energy Expenditure:
Isotopic Measurements

Isotope: element with atypical atomic weight

Can be radioactive or nonradioactive

Can be traced throughout body

13C, 2H (deuterium) common isotopes for studying

energy metabolism

Easy, accurate, low-risk study of CO2 production

Ideal for long-term measurements (weeks)
Energy Expenditure at Rest and During
Exercise

Metabolic rate: rate of energy use by body

Based on whole-body O2 consumption and

corresponding caloric equivalent

At rest, RER ~0.80, VO2 ~0.3 L/min

At rest, metabolic rate ~2,000 kcal/day
Energy Expenditure at Rest:
Basal Metabolic Rate

Basal metabolic rate (BMR): rate of energy expenditure

at rest

In supine position

Thermoneutral environment

After 8 h sleep and 12 h fasting

Minimum energy requirement for living

Related to fat-free mass (kcal  kg FFM-1  min-1)

Also affected by body surface area, age, stress, hormones,
body temperature
Resting Metabolic Rate and
Normal Daily Metabolic Activity

Resting metabolic rate (RMR)

Similar to BMR (within 5-10% of BMR) but easier

Doesn’t require stringent standardized conditions

1,200 to 2,400 kcal/day

Total daily metabolic activity

Includes normal daily activities

Normal range: 1,800 to 3,000 kcal/day

Competitive athletes: up to 10,000 kcal/day
Factors That Affect BMR

Age: BMR gradually decreases with age, generally because of a
decrease in fat-free mass

Body temperature: BMR increases with increasing temperature

Psychological stress: Stress increases activity of the
sympathetic nervous system

Hormones: Thyroxine from the thyroid gland and epinephrine
from the adrenal medulla both increase BMR
Metabolic Rate During
Submaximal Exercise

Metabolism increases in direct proportion to the increase in
exercise intensity

During exercise at a constant power output (work rate) VO2
increases from its resting value to a steady-state value within
1-2. minutes

There is a linear increase in the VO2 with increases in power
output (work rate) .
Increase in Oxygen Uptake with
Increasing Power Output

Reprinted, by permission, from G.A. Gaesser and D.C. Poole, 1996, “The slow component of oxygen uptake kinetics in
humans,” Exercise and Sport Sciences Reviews 24: 36.
Increase in Oxygen Uptake with
Increasing Power Output

Reprinted, by permission, from G.A. Gaesser and D.C. Poole, 1996, “The slow component of oxygen uptake kinetics in
humans,” Exercise and Sport Sciences Reviews 24: 36.
Energy Expenditure During
Maximal Aerobic Exercise

VO2max: The maximal capacity for oxygen

consumption by the body during maximal exertion

VO2max expressed in L/min

Easy standard units

Suitable for non-weight-bearing activities

VO2max expressed in ml O2  kg-1  min-1

Suitable for weight-bearing events

More accurate comparison for different body sizes
Maximal Capacity for Aerobic Exercise

Single best measurement of cardiorespiratory endurance and

aerobic fitness

Increases with physical training

Generally expressed relative to body weight (ml · kg-1 · min-1)

Normally active untrained college-aged students = 38-42 ml ·
kg-1 · min-1 for women and 44-50 ml · kg-1 · min-1 for men.

VO2max declines in active people after age 25-30 by ~ 1% per
year

Sex difference due to women’s lower FFM and hemoglobin
Relationship Between Exercise Intensity and
Oxygen Uptake in Trained and Untrained
Man
Anaerobic Effort and Exercise Capacity

No activity 100% aerobic or anaerobic

Estimates of anaerobic effort involve

Excess postexercise O2 consumption

Lactate threshold
Anaerobic Energy Expenditure:
Postexercise O2 Consumption

• O2 consumption requires several minutes to reach the

required steady state level at which the aerobic processes
are fully functional
• Oxygen deficit is calculated as the difference between the
oxygen required for a given exercise intensity and the actual
oxygen consumption
• Anaerobic effort can be estimated by examining excess
postexercise oxygen consumption (EPOC)—the mismatch
between O2 consumption and energy requirements
Anaerobic Energy Expenditure:
Postexercise O2 Consumption

O2 demand > O2 consumed in early exercise

Body incurs O2 deficit

O2 required − O2 consumed

Occurs when anaerobic pathways used for ATP production

O2 consumed > O2 demand in early recovery

Excess postexercise O2 consumption (EPOC)
Oxygen Requirement and Oxygen
Consumption During Exercise and Recovery
Factors Responsible for EPOC
• Rebuilding depleted ATP and PCr supplies
• Clearing lactate produced by anaerobic metabolism
• Replenishing O2 supplies borrowed from hemoglobin and
myoglobin
• Removing CO2 that has accumulated in body tissues
• Increased metabolic and respiratory rates due to increased
body temperature
Lactate Threshold

• It is the point at which blood lactate begins to accumulate

substantially above resting concentrations during exercise of
increasing intensity
• The rate at which lactate production exceeds lactate
clearance
• Usually expressed as a percentage of maximal oxygen uptake
• A high lactate threshold can indicate potential for better
endurance performance
• Lactate accumulation contributes to fatigue
Relationship Between Exercise Intensity and
Blood Lactate Concentration
Lactate Threshold and
Endurance Performance

Lactate threshold (LT), when expressed as a percentage of
VO2max, is one of the best determinants of an athlete’s pace in
endurance events. such as running and cycling. While untrained
people typically have LT around 50% to 60% of their VO2max,
elite athletes may not reach LT until around 70% to 80%
VO2max. .

.
Economy of Effort

As athletes become more skilled, use less energy for

given pace

Independent of VO2max

Body learns energy economy with practice

Multifactorial phenomenon

Economy  with distance of race

Practice
better economy of movement (form)

Varies with type of exercise (running vs. swimming)
Economy of Effort
Energy Cost of Various Activities

Varies with type and intensity of activity

Calculated from VO2, expressed in kilocalories/minute

Values ignore anaerobic aspects, EPOC

Daily expenditures depend on

Activity level (largest influence)

Inherent body factors (age, sex, size, weight, FFM)
Successful Endurance Athletes

1. High VO2max

2. High lactate threshold (as % VO2max)

3. High economy of effort

4. High percentage of type I muscle fibers

Fatigue and Its Causes

Fatigue: two definitions

Decrements in muscular performance with continued effort,
accompanied by sensations of tiredness

Inability to maintain required power output to continue
muscular work at given intensity

Reversible by rest
Fatigue and Its Causes

Mechanisms of fatigue depend on

Type, intensity of exercise

Muscle fiber type

Training status, diet

Four major causes

Inadequate energy delivery/metabolism

Accumulation of metabolic by-products

Failure of muscle contractile mechanism

Altered neural control of muscle contraction
Fatigue and Its Causes:
Energy Systems—PCr Depletion

PCr depletion coincides with fatigue

PCr used for short-term, high-intensity effort

PCr depletes more quickly than total ATP

Pi accumulation may be potential cause

Pacing helps defer PCr depletion

Energy Systems and Fatigue

PCr depletion

Glycogen depletion (“hitting the wall”)

Pattern of glycogen depletion from Type I and II fibers depends
on the duration and intensity of the activity

Glycogen depletion is selective to the muscle groups involved
in the activity

Depletion of liver glycogen to increase blood glucose increases
muscle glycogen utilization
Energy Systems—Glycogen Depletion

Fiber type and recruitment patterns

Fibers recruited first or most frequently deplete fastest

Type I fibers depleted after moderate endurance exercise

Recruitment depends on exercise intensity

Type I fibers recruit first (light/moderate intensity)

Type IIa fibers recruit next (moderate/high intensity)

Type IIx fibers recruit last (maximal intensity)
Fatigue and Its Causes:
Energy Systems—Glycogen Depletion

Depletion in different muscle groups

Activity-specific muscles deplete fastest

Recruited earliest and longest for given task

Depletion and blood glucose

Muscle glycogen insufficient for prolonged exercise

Liver glycogen
glucose into blood

As muscle glycogen , liver glycogenolysis 

Muscle glycogen depletion + hypoglycemia = fatigue
Glycogen Depletion in Different Muscle
Groups
Fatigue and Its Causes:
Metabolic By-Products

Pi: From rapid breakdown of PCr, ATP

Heat: Retained by body, core temperature 

Lactic acid: Product of anaerobic glycolysis

H+ Lactic acid
lactate + H+

Fatigue and Its Causes:
Metabolic By-Products

Heat alters metabolic rate

–  Rate of carbohydrate utilization

Hastens glycogen depletion

High muscle temperature may impair muscle function

Time to fatigue changes with ambient temperature

11°C: time to exhaustion longest

31°C: time to exhaustion shortest

Muscle precooling prolongs exercise
Figure 5.11
Fatigue and Its Causes:
Metabolic By-Products

Lactic acid accumulates during brief, high-intensity

exercise

If not cleared immediately, converts to lactate + H+

H+ accumulation causes  muscle pH (acidosis)

Buffers help muscle pH but not enough

Buffers minimize drop in pH (7.1 to 6.5, not to 1.5)

Cells therefore survive but don’t function well

pH <6.9 inhibits glycolytic enzymes, ATP synthesis

pH = 6.4 prevents further glycogen breakdown
Figure 5.12