The Physiological Infrastructure Model of Work Capacity
Functional fitness focuses on work capacity across broad time and modal domains. In plainer terms, fitness is the ability to perform a combination of tasks at the highest sustainable rate. This is a useful definition for comparing athletes but tells us little about how an individual athlete may improve their work capacity.
This article presents a cohesive model of work capacity based on four physiological parameters: motor unit recruitment, motor learning, phosphocreatine re-synthesis, and pH equilibrium. We show that increasing work capacity requires optimizing particular neuromuscular aspects of performance, while maximizing sustainable energy production. Our model enables coaches and athletes to identify the source of any work capacity gaps and correct them with targeted training interventions.
Muscle Contractions
The basic physiological activity of work capacity is muscle contraction. The key steps of muscle contraction are:
A nerve impulse is sent to muscle fibers, signaling demand for contraction. The collection of muscle fibers activated by an impulse is called a motor unit.
The impulse causes changes in electrical potential and release of calcium ions, preparing the muscle for contraction.
ATP provides energy for contraction. The part of the muscle fiber which contracts is called the myofibril.
These are the basics. A more technical discussion of the process of contraction, including video and graphics, can be found here.
MUSCLE FIBER CHARACTERISTICS
All fibers contract via the same process, but not all fibers are created equal. Fibers are commonly divided into fast twitch and slow twitch, but there are more differences than just contraction speed. Fibers vary in force production, energy production, and fatigue resistance. While there is a genetic component to the distribution of fibers by type, training can change muscle fiber characteristics.(1) In addition to pure fast and slow, hybrid fibers blend the characteristics of slow and fast fibers, (1) so that an individual’s musculature exhibits properties along a continuum from slow to fast. This chart summarizes some characteristics of muscle fibers.
Muscle fiber recruitment and utilization
Individual muscle contractions generally last 20-100 milliseconds, depending on fiber type. There are three phases to a contraction:
Contraction period; tension increases. (~0-20ms)
Relaxation period; tension decreases. (~20-100ms)
Interval between contractions. Varies based on the activity.
We don’t use every muscle fiber on every rep. For example, a 30% deadlift recruits fewer fibers than 90%. Toes to bar recruits different fibers than box jumps. During sustained activity, the brain cycles fibers on and off. As demand increases (heavier weight, more reps, increased intensity) additional fibers are recruited, and the interval between contractions decreases.
The sequence of recruitment is unclear; early research supported ascending size order of motor unit recruitment,(2) but newer findings indicates that larger, faster motor units be recruited first in response to mechanical demands of a task.(3) Regardless of the sequence, motor unit recruitment is task specific.
ATP and Energetics
Now that we’ve reviewed the basics of muscle fiber recruitment and contraction, let’s turn to the energy side. ATP provides energy to cellular processes, but very little ATP is stored in cells. Instead, cells produce ATP to meet increased energy demands. The immediate source of ATP is phosphocreatine (PCr). Cells can generally store 4 to 5 times more PCr than ATP. Experiments dating back to 1970 showed that muscle contraction was interrupted when PCr was depleted, though ATP concentration remained high.(4)
More recently, Imaging shows that during a contraction, ATP levels remain constant while PCr declines.(5) During the contraction period, ATP is consumed. PCr immediately resynthesizes ATP(6), ensuring a steady supply of energy. During the relaxation period, PCr is resynthesized from glycogen breakdown and glycolysis.(7, 8) Glycolytic processes and lactate production begin with the first muscle contraction. This has been observed repeatedly; experiments have shown significant lactate accumulation from intense exercise lasting only 5 seconds(9) or 10 seconds.(10) Between contractions, glycogen is resynthesized from plasma glucose.(7) The energy for this re-synthesis of glycogen is provided by the oxidation of lactate.
Ongoing exercise increases the concentration of ADP, signaling mitochondria to increase oxidative phosphorylation.(6) Energy produced in mitochondria is shuttled to myofibrils by phosphocreatine. We now have a clearer view of energy production:
- PCr is a store and shuttle of phosphates. During a muscle contraction, PCr instantly replenishes ATP. PCr also shuttles energy from the mitochondria to working muscles. Strictly speaking, PCr is not an energy system, as it does not extract energy from macronutrients, but simply moves and stores energy.
- Glycogen supplies ATP beginning immediately upon the commencement of exercise(7, 8, 10) and continuing for the duration of exercise.(11) Glycogen breakdown refills PCr pools during the relaxation phase of muscle contraction. Glycolysis has been shown to provide up to 20% of ATP for muscles working at sustainable intensity(11) even during longer duration activities.
- Mitochondria refill ATP and PCr pools in the interval between contractions. The concentration of ADP is a signal for the mitochondria to oxidative phosphorylation.
The Physiological Infrastructure model
The idea of three energy systems operating independently based on the duration of an activity is misleading. These energy systems function like gears in a machine, and each system is active during every muscle contraction. The sustainable ATP supply (and work rate) is governed by the interaction between glycolytic and oxidative processes. The respective contribution of each system depends on the demands of the task and the athlete’s training status. Rather than focusing on energy systems, we introduce the Physiological Infrastructure Model, which defines capacity in terms of neuromuscular activity and cellular energetics.
The neuromuscular parameters: recruitment and motor learning
Capacity begins with the recruitment of muscle fibers in response to an external stimulus, such as a barbell. Motor unit recruitment is influenced substantially by strength. A stronger athlete recruits a smaller proportion of muscle fibers than a weaker athlete to do the same work, delaying fatigue. Likewise, a stronger athlete can work at higher intensity than a weaker athlete, while using a similar proportion of motor units. For many tasks, strength is indispensable to work capacity.
Capacity is also enhanced by motor learning. Motor learning is a relatively permanent change in the nervous system, resulting from practice, producing improved movement quality. Consider double unders: proficient athletes turn the rope mostly with their wrists, and only jump high enough to complete the reps. Less experienced athletes will incorporate their upper arm and shoulder and may jump more than necessary. Proficiency improves work capacity by minimizing wasted energy and increasing work rates. For a stunning example of proficiency, check out Molly Metz doing 153 double unders in 60 seconds.
The energetics parameters: PCr re-synthesis and pH equilibrium
Motor unit recruitment and motor learning define the pool of active muscle fibers during a workout. Conditioning enhances the rate of energy production and resistance to fatigue of these muscle fibers. Therefore, conditioning is downstream from muscle fiber recruitment. Unlike monostructural capacity sports in which a small collection of motor units repeat the same task, conditioning for functional fitness should focus on
Maximizing sustainable energy levels for any task and/or duration
Producing repeatable above-threshold bursts of energy without reaching exhaustion
Energetics refers to the collection of physiological inputs to energy production, including mitochondrial density and enzyme content, capillary blood supply, stroke volume, glycolytic enzyme content, PCr quantities stored, proton buffering, and lactate shuttling. Rather than attempting to measure them all, we focus on two comprehensive parameters of energy production: PCr re-synthesis and pH equilibrium. These parameters integrate numerous factors and apply across the entire spectrum of motor units, so they provide a framework for training design for athletes at any level.
Phosphocreatine re-synthesis
During intense exercise, PCr concentration declines. Most of us have experienced this; it’s that feeling of hitting the gas, but the engine won’t go. But it’s not just a feeling. Experiments have shown that during intense exercise, performance is more closely related to the re-synthesis of PCr than to the level of lactate accumulation. There is also evidence of a strong relationship between the decline in PCr and a decline in force during high intensity exercise. (6) The extent of PCr breakdown is primarily dependent on the intensity of muscle contraction. Recovery of PCr during non-fatiguing exercise is almost exclusively oxygen dependent, but during very intense exercise, PCr recovery occurs in two stages
- An oxygen-dependent fast phase, beginning immediately upon cessation of exercise. Athletes with substantially depleted PCr have been shown to recover ~65% of their pre-exercise PCr after 90 seconds. (6)
- A pH dependent slow phase, which completes the recovery. After exercise of sufficient intensity, in which pH is substantially decreased, full recovery of PCr can take 10-15 minutes.
The likely mechanism of delayed PCr recovery after intense in reduced-pH environments is because low pH inhibits the ability of skeletal muscle to resynthesize ATP and therefore PCr. (6,11)
pH equilibrium
If you’ve pushed very hard and perhaps collapsed to the floor after a difficult workout, you’re familiar with reduced pH. Sometimes we call this being over the redline. Intense exercise recruits more glycolytic fibers, and glycolytic reactions produce lots pf protons, lowering pH if you cannot buffer or transport them. Low pH adversely impacts capacity due to:
- Decreased contraction velocity and ATPase activity. As pH decreased from 7.0 to 6.0, contraction velocity and ATPase activity decreased 27% and 25%, respectively. Isometric force production decreased 45%. (12)
- Reduced oxidative activity. A decline in pH inhibits the rise in ADP concentration, weakening the signal for increased mitochondrial activity, and limiting oxidative phosphorylation during exercise (11) resulting in lower sustainable work rates.
Possible decreased force production. One experiment showed that a drop in pH from 7.0 to 6.5 reduced isometric force by about 35%. (6) Another experiment showed that force decreased up to 44%, depending on muscle fiber type. (13) However, recent evidence has challenged the relationship between reduced pH and decreased force production, (14) so this remains somewhat unclear.
Athlete development in this framework
While effective training and athlete development programs should be individualized to an athlete’s goals and starting point, there are several principles which apply to training generally:
Effective strength and power training stresses a broad spectrum of motor units with bias toward the faster end. As athletes progress and technique improves, the bias toward intensity may increase. Training should also develop explosiveness (rate of force development) and utilize accessory work to correct imbalances resulting from previous athletic activities.
Skills practice should begin by prioritizing acquisition, retention or transference of a skill. Less experienced athletes generally benefit from low-stress repetition, whereas advanced athletes should encounter more randomized practice and contextual interference (15, 16)
Conditioning focuses on sustainable work rates. After correcting any imbalances, oxidative capacity across the broadest possible spectrum of motor units will enhance PCr recovery during any task. This can be accomplished with Polarization and interval work. High intensity intervals increase buffer capacity significantly,(17) enabling cells to maintain higher pH, which delays fatigue and facilitates more complete replenishment of PCr. Advanced athletes should encounter zero rest, mixed intensity and variable work-rest intervals.
Conclusion
We’ve described a model which aligns work capacity with physiology. Our model rests on four parameters which collectively encompass nearly all aspects of performance for fitness athletes: motor unit recruitment, motor learning, PCr re-synthesis and pH equilibrium. This framework enables coaches and athletes to quickly identify gaps in work capacity and personalize training interventions to maximize the value of training time. This framework is applicable to athlete at all levels, and can be disaggregated for specific goals. We look forward to employing it more broadly to enhance athlete performance and coaches’ understanding of the physiology of performance.
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References:
(1) https://pubmed.ncbi.nlm.nih.gov/10998639/
(2) https://journals.physiology.org/doi/pdf/10.1152/jn.1965.28.3.599
(3) https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1664648/
(4) https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4898252/
(5) https://pubmed.ncbi.nlm.nih.gov/9530118/
(6) https://pubmed.ncbi.nlm.nih.gov/12238940/
(7) https://www.ncbi.nlm.nih.gov/pmc/articles/PMC14608/
(8) https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1130636/pdf/biochemj00144-0029.pdf
(9) https://pubmed.ncbi.nlm.nih.gov/13942520/
(10) https://pubmed.ncbi.nlm.nih.gov/6618929/
(11) https://jeb.biologists.org/content/204/18/3189
(12) https://pubmed.ncbi.nlm.nih.gov/2842489/
(13) https://link.springer.com/article/10.1007/BF00713922
(14) https://academic.oup.com/ptj/article/81/12/1897/2857605
(15) https://pubmed.ncbi.nlm.nih.gov/21524808/