11 Lung Physiology
11 Lung Physiology
11 Lung Physiology
So far, on the MAPP I have devoted a lot of attention to two major organ
systems that are intimately linked in exercise performance, the skeletal
muscles and the cardiovascular system. Now I want to factor a third, equally
important system into the equation, the pulmonary system. I am going to
discuss breathing and exercise.
The actual act of moving air in an out of the lungs is accomplished in much
the same way that a bathroom plunger works. Since the thorax is "sealed,"
Cycling Articles: Physiology 3 11. Lung Physiology
expanding its volume causes air to rush into the lungs to fill the relative
vacuum. When we inspire air, we do so by contracting the diaphragm, pulling
it down (similar to pulling up on the plunger). If you are breathing "correctly" at
rest, the main movement you will notice is your stomach bulging out a bit
during inspiration. This is because the diaphragm is pressing down against
the abdominal cavity. You are "belly breathing." At rest and low ventilation
volumes, this is enough. However, when we need to move a lot of air in and
out of the lungs, we not only increase the force of diaphragmatic contraction,
we also contract muscles attached to the rib cage. This pulls the ribs up and
out, further expanding the thorax and allowing more air to rush into to the
lungs. All of this takes muscular work, so inspiration is an "active process."
Expiration, or blowing the air back out, is basically a matter of relaxation when
we are at rest. The elasticity of the muscles and tissues is sufficient to push
the air out. The process becomes more demanding as we exercise harder.
During heavy exercise we exhale more forcefully and deeply in order to
ensure more rapid and complete exhalation of the old air. This process also
takes energy in the form of muscular work. So, the bottom line is that
breathing is not free to the body. And, it gets more energetically expensive at
high workloads. I will come back to this point later.
2. Gas exchange: Moving oxygen and carbon dioxide on and off the red
blood cells.
OK, now that we are moving the air in and out of our body, let's turn our
attention to the lungs, and "get cellular" in our thinking. The lungs do some
pretty cool stuff. The lung's challenge is to mix air and blood thoroughly and
rapidly so that gas exchange can occur. Here comes another analogy. Let's
say you have a gallon bucket (4 liters) of paint sitting with the lid open. How
long will it take that bucket of paint to dry out? Days and days. But, if we
spread the paint as a very thin layer over a very large wall, it will be dry in no
time. By spreading the paint out we increase the total area of exposed
surface between the paint and the air thousands of times and the water in the
paint is quickly evaporated. At any given instant at rest, the lungs spread
about 70 ml of blood (less than half of the volume of a coke can) in a "sheet"
of capillaries with a total surface area of 70 square meters. That is like
spreading a gallon of paint thin enough to paint a football field! The capillaries
are so narrow that the red blood cells actually have to squeeze through. This
also insures that the gas exchange across the red blood cell and capillary
membranes is lightning fast. Simultaneously the lungs move the inspired air
down a system of 23 branches of air passages terminating with about 300
million tiny spherical alveoli that form the terminal exchange tissue in the
Cycling Articles: Physiology 4 11. Lung Physiology
bronchial system. These two exchange systems, the alveoli for air, and the
capillaries for blood, are intertwined so microscopically close that oxygen and
carbon dioxide molecules diffuse across the membranes and equilibrate
almost instantaneously. Blood passes through the capillaries in about 0.8
seconds at rest and as little as 0.4 to 0.5 seconds during hard exercise. It is
during this very brief exposure period that all the gas exchange between each
red blood cell and the air in the lungs must take place before each return trip
to the body!
Now perhaps one of you physiology student types is doing some analytical
thinking that does like this: If resting cardiac output is 5 liters per minute, and
maximal cardiac output were say, 25 liters/min, that would mean that 5 times
as much blood passes through the lungs (and back to the heart and then to
the body) and those capillaries per minute at max. Why doesn't the lung
capillary transit time for each blood cell decrease from 0.8 secs at rest to
0.8/5 or a really fast 0.16 seconds during maximal exercise? Here is the
answer. Normally, the lungs only use a fraction of the total capillary volume
available. Small arterioles can regulate the entry of blood into portions of the
lungs. During exercise this restriction is gradually removed and the capillary
volume can increase by over 3 fold to about 250 ml. This helps to minimize
the decrease in capillary transit time. It does not eliminate it though. How
important is that? This is another point which we will hit on again.
Now let's think about how the blood fits into all of this. Blood serves several
important functions during exercise (heat removal, deacidification, glucose
delivery, hormone communication to name a few), but the one I want to focus
on is its role as a delivery truck for oxygen. About 40 to 50% of the total
volume of blood is made up of red blood cells (RBCs). For example, if your
"hematocrit" is 43, then 43% of your total blood volume is RBCs. More than
anything else, RBCs are just tiny, flexible sacks of hemoglobin. Each RBC
contains hundreds of hemoglobin molecules, and each hemoglobin molecule
has room to carry exactly 4 oxygen molecules. When I say "carry" I mean
"bind" in chemistry lingo. There is some very fancy chemistry going on here
that we would be in deep trouble without, but I will not try to explain it other
than to say that hemoglobin molecules are engineered to hold on to oxygen
tightly enough to carry it out of the lungs, but loosely enough to release it in
the capillaries feeding the skeletal muscles (and other organs of course). This
whole process is designed to function best when the atmospheric pressure is
near sea level. If we get 1500 meters or more higher than that, the system
begins to break down, and hemoglobin leaves the lungs without a full load of
oxygen. This is why it is more difficult to breathe and exercise at altitude.
So, for an average person with a hemoglobin of 15, the oxygen volume
contained in each liter of delivered blood will be:
15g/dl x 1.34 ml O2/g hgb x 0.96 saturation x (10dl/l) = 193 ml O2/ liter
blood.
Cycling Articles: Physiology 6 11. Lung Physiology
If we substitute in 12 for the hemoglobin concentration (someone with
anaemia) and 18 (a very high value occasionally seen in trained athletes at
high altitude), we see that for the same cardiac output, the volume of oxygen
carried by the blood would vary between 154 and 232 ml per liter, depending
on the hemoglobin value. It is not hard to see how the blood oxygen carrying
capacity affects the VO2 max. Remember, the muscles can only use what the
heart can deliver.
If hemoglobin concentration is higher, the blood can carry more oxygen. This
is an important point with relevance to altitude training, illegal EPO use,
gender differences in VO2 max, anaemia etc. (I should note here that there is
a downside to increasing haemoglobin concentration in the blood and that is
increased blood viscosity. The body normally maintains an appropriate
balance. If the blood becomes too thick, flow resistance increases and the
risk of blood embolism increases, hence the dangers of EPO use.) Second,
when the blood leaves the lungs it is normally fully saturated with oxygen.
This means that the lungs are very effective at ventilating the blood, even in
untrained folks. This is one of the reasons why in the big scheme of things we
basically disregard lung function as an area for improvement in the athlete's
endurance machine. But, this issue is worth taking a closer look at.
Sometimes you hear people say "I ran out of wind." Is that really possible?
Can we reach a point in exercise when ventilation just can't keep up with
demand? The answer is no, assuming you don't have acute asthma or some
other severe pulmonary dysfunction. We can measure a person's maximal
voluntary ventilation (MVV), the maximal volume of air they can breath in and
out while at rest, and compare it with their maximal ventilation during
exercise. What we see is that untrained people only use about 60 to 85% of
their maximum ventilatory capacity even at maximal exercise. For example
the MVV for an average male might be nearly 200 l/min. However, during a
treadmill VO2 max test, they reach a peak ventilation of only 140 l/min. Highly
trained athletes use more of their capacity, perhaps over 90%, but ventilation
capacity is still not a limitation on performance. Unlike the story with cardiac
output, even during maximal exercise, the ventilatory capacity is not maxed
out.
Another sport that has gotten some special attention from the ventilation folks
is rowing. This I know more about, so I will elaborate a bit. During rowing, the
body is squeezed up with the chest against the knees over 30 times a minute,
limiting diaphragmatic excursion. That might create some breathing problems,
but it is not the biggest issue. The real issue is the fact that rowers also use
the same abdominal and intercostal muscles used for breathing to support
the back during the powerful extension employed each stroke. Rowers
isometrically contract all of these muscles to apply a high interthoracic
pressure at the moment of the catch, when the oars take the water, to
reinforce the connection between oar, back and legs. It is impossible to
breathe and constrict all the abdominal and thoracic muscles at the same
time.
If my wife joins me at the rowing club for a workout on the rowing machines,
an interesting phenomenon occurs. Hilde is not a rower, so she always
seems to adjust her rowing cadence so that it matches mine. I don't think she
does it on purpose, but her rowing rhythm naturally entrains onto mine. This
is problematic when I am doing intervals and she is rowing steady state! Our
ventilatory system does the same thing. Ventilation tends to match with
running, cycling or rowing cadence in a consistent pattern. For example, in
cycling, we sometimes see athletes exhale in unison with the downward kick
of the same leg, every 2nd or third stroke. This entrainment process does not
seem to be a bad thing. In fact, since it is more prevalent in experienced
athletes, it is probably an adaptation that promotes efficiency by minimizing
the mechanical constraints to breathing created by limb movements.
New topic. Earlier in this breath-taking novel of mine, I discussed the issue of
hemoglobin saturation with oxygen. I said that the lungs were so good at
oxygenation that the blood always leaves the lungs saturated with oxygen,
even during hard exercise when cardiac output is high. This means every
RBC picks up a full load of oxygen before leaving the lungs. Now I am going
to contradict myself a bit.
Here is the central issue. The blood returning from the periphery must
eliminate its carbon dioxide load and fully re-saturate with oxygen during the
brief time it passes through the lung capillary network on the way back
through the heart and out to the body again. Normally this is not a problem. It
only takes about 0.45 seconds for the hemoglobin to become fully saturated
during its passage through the twisting capillaries. It takes even less time to
unload the CO2. Since the transit time is 0.8 seconds, there is time to spare.
Even during exercise there is enough time, unless......you are a really fit
athlete with a very high cardiac output and VO2 max.
Recent studies with highly trained endurance athletes (VO2 max over 70
ml/kg/min) have shown a significant degree of arterial desaturation. This
means that for the guys with the really big cardiac outputs, the blood is
rushing through the lungs so fast that hemoglobin hasn't taken on a full load
of oxygen before leaving for the muscles. The result is that instead of being
96 or 97% saturated with oxygen, the blood leaving the lungs may only be 89
or 90% saturated in the athletes with very high cardiac outputs and VO2 max.
Clifford et al. (1990) reported a drop in arterial saturation from 105 mm Hg at
rest to 88 mmHg during the last minute of a maximal rowing test in elite
rowers. This means that the arterial blood was becoming slightly hypoxic
when they reached very high workloads. All other things being equal, VO2
max might be up to 5 % higher or so in the elite types if the lungs could fully
saturate the blood at maximal cardiac output. Support for this assumption
comes from the fact that when well trained athletes breathe a higher
concentration of oxygen while performing in a lab, they reach a slightly higher
VO2 max.
What does this information mean in regards to how we train? Nothing. There
is nothing we can do to prevent this desaturation in folks with the really high
Cycling Articles: Physiology 11 11. Lung Physiology
cardiac outputs, short of having them wear an oxygen tank while performing.
Unless you have a VO2 max of about 5 liters/ min or higher, it is not really an
issue anyway. Consider arterial desaturation a small physiological tax on the
"cardiovascularly endowed."
The ventilatory muscles, like all muscles need oxygen to support continuous
exercise. It turns out that the diaphragm is one of the body’s best endurance
muscles, perhaps even in second place behind the heart. It has a high
percentage of type I fibers, a high capillary density, and high concentration of
oxidative enzymes, compared to skeletal muscles. Animal studies have
demonstrated that the diaphragm improves its endurance capacity
(mitochondrial enzyme concentration) with training, but not more than about
20-30%, because it is already pretty well equipped for chronic work. However,
with high intensity training, other muscles involved in breathing like the
internal and external intercostals and the abdominal muscles become more
active and also improve their endurance capacity. Since these muscles are
less trained to begin with, they respond more to endurance training. These
accessory breathing muscles are not trained at low exercise intensities but
become active when we really start moving a lot of air.
From the above, plus your own experience, you can figure out that breathing
becomes more demanding when ventilation rates get very high. There are
studies which have actually measured the oxygen cost of breathing at
different intensities. You might think of this as the tax on oxygen delivery. The
body has to deliver blood to the ventilatory muscles so that they can help the
lungs supply oxygenated blood to the rest of the body. To make things worse,
this “tax rate” increases when you are working at very high intensities. The
oxygen cost per liter of ventilation (VE) doubles from low to very high exercise
intensities.
The bottom line is that while the oxygen cost of breathing is only perhaps 3-
6% of total VO2 at low intensities, it can be as high as 10 to even 15% of total
VO2 in young adults with greater than average VO2 max. In the fit older
athlete, a high oxygen cost of breathing may occur at lower ventilation rates
due to the increased stiffness of the chest wall.
When you add this to the problem of desaturation, we see that there are
some additional limitations on maximal performance that enter into the picture
when we start dealing with extremely well trained endurance athletes. These
guys have such high cardiac outputs and work capacities that the ventilation
machinery starts to demand a lot of the total available oxygen in order to keep
the machine running.
To make things short and sweet it appears that they do fatigue. Tests of
maximal ventilatory function after a hard endurance session show a
temporary drop in, for example peak expiratory force. This is an indirect way
of measuring how much force the diaphragm can generate.
To date the best technique for answering this question is to unload the
ventilatory muscles and observe whether performance improves. These
techniques have included using helium oxygen mixtures and breathing assist
devices. Unfortunately, the methodology is not perfect. For example, using
lighter than air helium may make the hoses and mouthpiece easier to hold in
the mouth and actually decrease the cost of rowing. The results are unclear.
At workloads below 85% of VO2 max it appears the respiratory fatigue has no
influence on performance. However, the results of some studies suggest that
at intensities approaching VO2 max respiratory fatigue may contribute to
Cycling Articles: Physiology 13 11. Lung Physiology
performance limitations. The scientific jury has not reached a verdict on this
question.
So, if you made it to the end of this novel, you may be disappointed to learn
that there are no secret breathing tricks that will push you over the top. In
general the lungs are wonderfully equipped for doing their job. Training does
improve the ventilatory system in some ways, but it is not the weak link in
healthy athletes. In recent years, there have been a handful of studies
published where the impact of inspiratory muscle training on various aspects
of pulmonary and endurance performance have been investigated. This
involves essentially weight training for the breathing muscles, where
resistance is generated by using some kind of device that reduces airflow
during inspiration and forces the inspiratory muscles to work harder against
greater resistance. Neither peak pulmonary function nor maximal oxygen
consumption have been shown to change with this form of training. However,
a couple of studies have shown modest increases in either time to exhaustion
or time trial performance during cycling, using placebo controlled designs.
How does this work? Perhaps stronger inspiratory muscles allow high
ventilation to be achieved at lower breathing frequencies. This would
decrease the oxygen cost of breathing and free up some blood flow for the
working muscles. Perhaps.