A REVIEW OF FLIGHT

Fuel and Blood Chemistry in Racing Pigeons

Luckily for us as fanciers, over these many years, pigeons have been the subject of a
great deal of investigation, especially in the areas of muscle and the study of energy for
flight. As a matter of fact, the very foundations of all modern knowledge of the
physiology (refers to the normal chemistry and function) of muscle in many species,
including birds, animals and of course, humans, were established by early work
conducted by a researcher named Szent-Gyorgyi who used the breast muscles of pigeons
in his biochemical experiments.

These early investigations by Szent-Gyorgyi ultimately led to the important discovery of

the key energy cycle, known as the Krebs Citric Acid Cycle, named for Hans Krebs who
also used the breast muscles of pigeons in the universally acclaimed work that led to this
major discovery.

In summary, because the biochemical steps in the whole process are extremely complex,
the work done by Krebs showed that, within cells of the body, in the presence of oxygen,
the major fuels, glucose and fat are metabolized (roughly means "broken down" or
"utilized") in the tissues in a stepwise fashion, to the final products, carbon dioxide and
water. Through these complex chemical processes to reach the final products of carbon
dioxide and water, important chemical by-products are produced along the way as spin-
offs in the whole process.

One of these very important compounds represents the key to all work, and is called
adenosine triphoshate, ATP for short. Thus, the energy for work of any kind -- and in
particular the energy for flight, which interests us the most -- is developed in muscle
fibers through the production of this very important high-energy compound known as
ATP. It is a potent compound that in every-day terms is comparable to the steam that
once provided the energy to drive the old railroad locomotives. Regardless of the fuel
being used -- fat or glucose -- the main purpose is to produce ATP to supply the energy
for the different phases of flight, including the launch, rapid sustained cruising flight,
dodging bursts of speed at any time, braking to land, etc...

ATP derives its power from the three -- hence, tri phosphate -- important high- energy
phosphate bonds that make up this compound. These three high-energy phosphate bonds
provide the energy, delivered through the massive breast muscles, that allows the wings
of our birds to beat on an average of about 9.5 times per second during the explosive
launch phase of flight, and on an average of about 5.5 times per second at cruising speed
for the duration of the training toss or race, however long it might be. (Indeed, the two
great breast muscles of the pigeon are so powerful that one of them alone is capable of
developing a force about ten times the weight of the bird -- definitely, tremendous
power.)
Through all of this it is significant to realize that it was the pioneering work of Szent-
Gyorgyi and Krebs on the breast muscles of pigeons that established the dynamic
biochemical foundations of energy production in various species of animal’s right
through to the level of humans.

As noted previously, the major fuels used to provide the energy for short or prolonged
flight are glucose and fat, and particularly the latter when prolonged flight is required. In
the explosive launch phase of flight -- and indeed whenever explosive or dodging bursts
of speed are needed during the course of flight -- glycogen, the storage form of glucose in
liver and muscle, is metabolized rapidly to glucose in the white muscle of the breast
muscles.

In turn, the glucose obtained from the breakdown of glycogen is metabolized in the white
muscle to produce the ATP needed to provide energy for this rapid action.

Note that for the launch phase of flight, glucose from the breakdown of glycogen is the
chief fuel needed. So powerful and energy-demanding is the launch phase that within 10
minutes or so, all of the glycogen in the white muscle of the breast is completely utilized,
and to all intents and purposes, the white muscles cease to function because of the
depletion of their main source of energy. Importantly, during the course of the flight, the
white muscles are recharged with glycogen to provide the energy for sudden dodging and
other explosive bursts of energy, which would be needed if there were attacks by falcons
or other raptors, for example, or for any situation requiring a sudden, dramatic increase in
speed.

The process of re- fueling the white muscle with glycogen originates in the liver which
has abundant supplies of glycogen. In the liver, glycogen is broken down to glucose for
transport through the blood stream to the white muscles in the breast. In the white
muscles, glucose is once more built up into abundant supplies of glycogen that are ready
for the kinds of emergencies described earlier.

If the white muscles are once again depleted of glycogen by any of these emergencies,
they are subsequently re-fueled, so that even at the prolonged flight time of 18 hours, the
white muscles seem to contain useful amounts of this fuel. It is important to be aware as
well that some of the glucose liberated to the blood stream from the liver is critically
needed as a source of fuel for the brain which can use only glucose as a source of energy.
Thus, it is critical that there be a steady flow of glucose from the liver, and is one of the
major reasons that birds normally have a high level of glucose circulating in the blood
stream.

By contrast, fat in the red muscle of the breast is metabolized to produce the energy from
ATP needed for the prolonged effort of flights that range from regular training tosses to
short, middle, long distance and endurance performances. For sustained flight, within a
short time, there is a shift from the initial use of white muscle which utilizes glucose for
the launch phase, to the use of red muscle which utilizes chiefly fat for the duration of the
flight.
Supplies of fat are present in the red muscle and as they are depleted during flight, they
are replenished from depots in the body cavity. The flow of fat to the red muscle occurs
from the fat depots where the fat is converted to fatty acids for transport through the
blood stream into the red muscles. Although the white muscles seem to be quite well
supplied with glycogen even at 18 hours of flight, by contrast at this same time, the red
muscles, which are the real workhorses of the system, are found to be severely depleted
of fuel, so that the onset of fatigue becomes imminent and even critical.

As noted in an earlier article, in migratory birds, migratory fat is distinguished from

winter fat. Migratory fat is built up rapidly in the body cavity of these birds in preparation
for prolonged flight, and is also completely utilized at the end of migration. Winter fat,
located under the skin, is used primarily as insulation against the cold.

This fat, combined with a certain amount of shivering to create heat helps to keep the bird
warm in a cold environment. The use of fat as insulation is a much more efficient method
of retaining heat than the continued wasteful use of fuel for shivering to generate heat.

In looking at racing pigeons, it seems likely to me that the fat that is built up rapidly each
week in the body cavity among the intestines, etc., is probably of the "migratory" type
because it is well utilized and may be almost completely depleted, depending on the
severity of the race, by the end of the race. From a biological point of view then, pigeon
races each week may well represent a repeated migratory phase of their lives.

As also noted in an earlier article, the breast muscles of the pigeon contain two types of
muscle - red and white - with red muscle by far the most abundant. Why does red muscle
appear red or red-brown, and white muscle white or at least pale? The difference lies in
the fact that red muscle contains myoglobin, a complex, pigmented chemical structure
that is similar to hemoglobin, the compound that imparts the red color to blood.

Both hemoglobin and myoglobin contain iron, which gives them their red color, and both
have a great capacity to hold oxygen for transfer to tissues. Red muscle fibers contain
abundant myoglobin which holds and then transfers plentiful supplies of oxygen to the
muscle structures known as mitochondria -- the powerhouses of all cells, where ATP is
generated -- that utilize the fat in the production of energy for flight.

White muscle fibers contain little or no myoglobin, and as a result they have developed
the capacity to utilize glycogen in the absence of oxygen (called anaerobic metabolism:
"an" = "without"; "aerobic" = "oxygen") in the production of energy for their explosive
role in flight. Also, because they don't contain the pigmented compound myoglobin, they
are very pale, and are described as white. Thus, the great breast muscles of the domestic
chicken or turkey are comprised mainly of white fibers, and appear pale for that reason.
By comparison, the massive breast muscles of the pigeon and any cut of beef for that
matter are good examples of muscles that contain mainly red fibers.

To digress momentarily, cells in various organs of the body are of different types, shapes
and functions. For example, cells of the liver look like sugar cubes or dice; some brain
cells resemble a spider or an octopus; skin cells, especially those near the surface, are flat
like pancakes, and so on. When muscle cells are separated out from the main breast
muscles by delicate micro techniques, they are seen to be elongated or thread-like, and in
fact, resemble fine threads or fibers. Hence, by convention, muscles cells are usually
described as fibers.

If a lengthwise section of muscle is cut out of the breast of a pigeon, fixed in formalin to
stop all chemical processes, processed in a series of chemicals such as alcohols and
xylene, embedded in wax, cut very thinly, placed on a glass slide, and then stained with
certain dyes, it can be examined by use of a microscope. A lengthwise section of stained
muscle has the appearance of a number of cigar- shaped fibers lying side by side and end
to end. Closer inspection reveals that there are fibers of two different diameters, one
broad and the other narrow. The broad fibers have a relatively smooth looking interior,
whereas the narrow fibers have a rough interior because of the amount of "machinery"
and fuel they contain.

To obtain a different look at the microscopic appearance of the muscle, we can also cut
our original piece of muscle in cross section and stain it to get an end-on view of these
fibers. Here, we see that the fibers are arranged in bundles that are separated from one
another by connective tissue. When we look at the muscle in cross section, we see even
more clearly that there are fibers of two different diameters, one a broad diameter and the
other, a narrow diameter fiber, and further that, for the most part, the broad diameter
fibers are located on the outside edge of each bundle, whereas, the smaller diameter fibers
are located more deeply within a bundle.

The broad diameter fibers are known as white fibers, and the narrow diameter fibers are
the red fibers. In diameter, the white fibers are over twice as broad as the red fibers. In
cross section, all of the fibers regardless of their diameter, whether they are red or white,
are round or oval. Fibers may extend the entire length of a muscle, or may form long
strands by connecting to other fibers.

If we were to count several series of 100 fibers, we would see that the red fibers vastly
outnumber the white fibers; for every 100 fibers, approximately 85-95 are red fibers and
only 5-15 are white fibers. Note the point: the great breast muscles of the pigeon contain
both red and white fibers, but the muscle appears red to the eye because of the
predominance of the red fibers.

Both types of fibers are present prior to hatching, a finding that indicates that the
differentiation between the two is genetically determined, and not the result of
specialized activity after hatching. Incidentally, as an athletic individual becomes
conditioned for racing, the number of fibers doesn't increase, but rather the existing fibers
simply enlarge to handle the increased work load.

Red fibers are abundantly supplied with microscopic thread-like blood vessels called
capillaries that cris cross and interconnect over the surface of each of these fibers. This
extensive meshwork of blood vessels is necessary because the red fibers utilize fat in a
system that requires oxygen (aerobic metabolism), and a steady flow of oxygen-rich
blood to these fibers ensures that, along with good supplies of fuel that is also delivered
in these blood vessels, they are able to function rapidly and efficiently for many hours at
a stretch. On the other hand, the white fibers are very poorly supplied with blood vessels,
since they don't require oxygen to function.

Now, if we cut another thin cross section of muscle, place it on a glass slide, and apply a
stain that detects only glycogen, we see that there is very intense staining of the white
fibers, which indicates clearly, an abundance of glycogen in these fibers - about 10%.
However, there is also some less intense staining of the red fibers, an indication that they
contain glycogen also, but much less - about 2.5%.

If we cut another cross section of the breast muscle and this time apply a stain that detects
only fat, we see that there is very intense staining of the small red fibers, but no staining
whatsoever in the broad white fibers. It becomes immediately clear that the chief fuel for
the broad white fibers is glycogen, and for the more abundant narrow red fibers, the chief
fuel is fat, although as noted, lesser amounts of glycogen are also present in these red
fibers. Overall, analysis of a section of the breast muscles containing a mixture of both
red and white fibers indicates a glycogen content of 3.5% and a fat content that ranges
between 10-15%.

As well as their differences in fuel supplies, it is known that the two fiber types have
differing speeds at which they operate - called "twitch" speed. Although both twitch very
rapidly indeed, white fibers twitch much more rapidly than red fibers. Because white
fibers twitch so rapidly, they also utilize their glycogen fuel supplies very rapidly, and so,
tire out very rapidly. For this reason, it becomes obvious that white fibers cannot be used
for prolonged, hour upon hour flight.

So what is their main function? Because they twitch very rapidly and tire out very rapidly
as well, white fibers are used for explosive actions, such as the launch phase of flight, or
for dodging bursts of speed to escape aerial predators, or in the braking process for
landing. So rapidly is the glycogen reserves utilized at launch for example, that it is
known that this complete depletion of glycogen in the white fibers occurs within about 10
minutes after the launch. To indicate how rapidly the white fibers can twitch, note that
birds shivering from cold, or those whose trembling wing tips indicate birds in top
condition, are using their white fibers for these actions.

By contrast, although they truly twitch very rapidly, the red fibers nevertheless twitch
much less rapidly than the white fibers, and as a result, they tire out much more slowly. It
becomes very obvious then that the red fibers, utilizing their abundant stores of fat, and
calling on further fat supplies from the body cavity when they are needed, are the real
work horses of the breast muscles, and can handle the challenge of rapid, sustained flight
over the many miles of the race course for many hours at a stretch.
There are those who believe that glycogen is the fuel needed for training tosses and short
races, and that possibly fat is needed only for more distance events. Some even
completely discount fat as a fuel, believing (incorrectly) as they do that glycogen is the
key fuel for any race of any distance. The well-established scientifically proved facts are,
however, that glycogen is the chief fuel for the launch and certain other explosive phases
of flight as noted previously, and that fat has been clearly established as the key fuel of
the red fibers for any flight from a short training toss to the longest endurance event.

The views of some commercial interests and others who propose "carbohydrate loading"
in racing pigeons seem to be at direct variance with the known and well established facts
about the fuels needed by pigeons for racing performance.

Having said that, I do believe however, based on facts from scientific literature, that the
timely use of sugars such as glucose and fructose (in addition to the use of high energy
grains) is highly valuable in the preparation of birds for racing, simply because both
sugars are known to be converted readily to fatty acids in the liver -- see earlier articles
for details. From the liver these fatty acids are exported in the blood stream to the red
muscle fibers where they are the major fuel for the sustained efforts of flights ranging
from short to long distance.

Now, some other interesting findings related to flight in pigeons. In Canada, experiments
on other important aspects of flight were conducted by Dr George and his colleagues,
with two flocks of racing pigeons. One flock of birds had been in regular training and
was considered to be fit birds. A second flock of birds had not been in training for three
months and were very likely in off condition. Both groups were transported to a release
point located about 48 km (approx 30 miles) from home. A third flock of birds, known as
controls (because they were rested and not exercised), was simply transported to the
release point and returned to the loft without being flown. This was done to determine
whether there was any effect on birds simply from the trip to the release point alone.

The Autumn day chosen for this study was clear, calm and sunny with a temperature of
6oC (about 43oF). Birds in the first two flocks were released in groups of 2-3. Birds in
the trained group homed in 60-80 minutes, whereas the group that hadn't been in training
for three months homed in 90-160 minutes. (After reading these results, one of my main
questions was: why did trained pigeons that should have been in good physical condition
take so long to arrive home from such a short distance? In my experience, on a calm day,
birds should fly a mile in 1 to 1 1/2 minutes, so from 30 miles I estimated that they
should have homed in about 45 minutes maximum. These results are certainly puzzling
but that is the way they were reported. I can only conclude that there were restraining
effects from factors we don't know about.)

As the birds arrived home, blood samples were collected in separate tubes from a wing
vein of each flown bird and from each bird that was just transported and not flown. These
blood samples were allowed to clot and the resulting serum (serum is the pale yellow
fluid that separates from the clot) was collected and then analyzed for a number of
constituents, and the results for each of the three groups were compared.
Compared with the rested birds, in those birds in the flock that had been in regular
training before this toss, there were significant increases in blood levels of glucose,
lactate (lactic acid), fatty acids, uric acid, and in the following hormones: growth
hormone, adrenalin, thyroid hormone, AVT (arginine vasotocin), and melatonin.
In the flock that had not been in training for three months, there were significant
increases in triglycerides (fat), free fatty acids, and the hormone glucagons, but no
significant increase in blood levels of corticosterone. Now before anyone begins to panic
and stumble over some of these big words, I will try to explain as well as I can, what
these results mean!

Read on.

To supply energy, fuel is needed for many tissues, among which the brain and breast
muscles are highly important. To liberate glucose and fatty acids, the release into the
blood stream of two hormones, namely growth hormone and glucagons, is important, so
their levels in the blood rise during flight. In turn, these hormones target the liver and the
fat depots where they stimulate chemical reactions that liberate glucose and fatty acids
into the blood stream for use as fuel by brain and muscle. Blood levels of adrenalin from
the adrenal glands located just ahead of the kidneys, also rise and aid in the release of
glucose into the blood.

Levels of lactate (lactic acid) in the blood rise as well during the launch phase of flight
when the white fibers are working at their maximum, and also if dodging bursts of speed
are needed on the way home, or if the birds have to work hard against a head wind, and
during the braking phase of landing. The reason is that the white fibers that are likely
used to a great extent during these intense muscular efforts, use glucose derived from
their stores of glycogen in a chemical reaction that doesn't require oxygen (thus, it is an
anaerobic process). In such an anaerobic state, one of the by products of the reaction is
lactic acid - hence its elevation in the blood stream.

As discussed in an earlier article, blood levels of uric acid and the hormones AVT and
melatonin also rise during flight, and are believed to be useful in preventing the birds
from overheating. Recall from an earlier article that the amount of heat produced during
flight in the pigeon has been estimated to increase about six times compared with amount
of heat produced by a resting bird. The most important site for the loss of most of the heat
from the body seems to be the legs - witness the dangling legs of some birds while they
exercise around the loft on a hot day.

Melatonin seems to exert its temperature-reducing effects by causing blood vessels near
the surface of the body to dilate (expand in diameter) in order to bring more heat from the
warmer core of the body to the surface. The hormone AVT also seems to be useful in
mobilizing fat from storage areas for use by the flight muscles, and by playing a role in
decreasing the loss of water from the body, along with its role in the regulation of body
temperature. Blood levels of uric acid also rise during flight in pigeons and may well be
another component of the complex chemical interactions that try to ensure heat stability
and water conservation during flight. It was mentioned in an earlier article that uric acid
levels are known to increase in wild birds during migration, and in human athletes
exercised to exhaustion.
Although blood levels of corticosteroids released from the adrenal glands -- which are
indications of a response to stress -- did not rise significantly during the test flights in
Canada, another unrelated study in Germany found that there were large increases in the
level of this hormone in pigeons flown greater distances than those flown in Canada.

In the German study, trained pigeons were flown from distances ranging through 115,
225, 314 and 557 km (about 70, 135, 188 and 330 miles, respectively). Flight times
ranged from 315 to 561 minutes (about 5¬ to 9« hours). Blood samples collected at the
end of these flights had a high level of the hormone corticosterone, likely a direct
response of the birds to the stress of flying these greater distances. As well, this hormone
which is released from the adrenal glands, combined with others mentioned earlier, may
help to mobilize glucose from the liver and some say, fat from body fat depots for use
during these prolonged flights.

The likely reason that blood levels of corticosterone didn't rise in the birds studied in
Canada, is that the distance and the time to home were not sufficiently stressful to induce
the release of this hormone. By contrast, the time and distances flown by the birds in
Germany called for much more strenuous effort, and therefore were sufficiently stressful
to cause the release of this hormone.

Sounds complex, doesn't it? Well, it is, and the foregoing explanations merely scratch the
surface of all of the events that occur in any living creature, human or otherwise, during
work, and indeed during life in general. Like any other living creature, the racing pigeon
is a complex animal endowed with beautifully intricate and co-coordinated biochemical
systems that convert food into the energy of life. I confess that I know and understand
very little about all of the complexity of these marvelously well meshed, intricate
systems, but I am fascinated that they work so astonishingly well and efficiently, not just
in pigeons but in all creatures, humans included.

In closing, I should say that my lack of solid knowledge on the whole subject reminds me
of a perhaps pertinent quotation from the ancient philosopher Socrates. It seems that
Socrates had an argument or disagreement with a friend or colleague, and after they
parted company, Socrates later wrote: "And so I left him, and as I went away, I said to
myself, 'Well, although I do not suppose that either of us knows anything, I am better off
than he is. For he knows nothing, and thinks that he knows. I neither know nor think that
I know. In this latter particular then, I seem slightly to have the advantage of him.'"

Like old Socrates, I neither know nor think that I know, but I am willing to expose and
share my ignorance with you!

Good racing!