9. Lucchini R., and J. M. Sogo. Replication of transcrip- and assembly.

In: Chromatin Structure and Gene

tionally active chromatin. Nature 374: 276-280, 1995. Expression, edited by S.C.R. Elgin. Oxford, UK: IRL,
10. Podust, L. M., V. N. Podust, J. M. Sogo, and U. 1995, p. 49-70.
Hiibscher. Mammalian DNA polymerase auxiliary pro- 13. Sogo, J. M., H. Stahl, T. Koller, and R. Knippers. Structure
teins: analysis of replication factor C-catalyzed prol if- of replicating simian virus 40 minichromosomes. The
erating cell nuclear antigen loading onto circular dou- replication fork, core histones segregation and terminal
ble-stranded DNA. Mol. Cell. Biol. 15: 3072-3081, 1995. structures. J. Mol. Bio/. 189: 189-204, 1986.
11. Smith, S., and B. Stillman. Purification and character- 14. Waga, S., and B. Stillman. Anatomy of a DNA replica-
ization of CAF-1, a human cell factor required for tion fork revealed by reconstitution of SV40 DNA
chromatin assembly during DNA replication in vitro. replication in vitro. Nature 369: 207-212, 1994.
Ce// 58: 15-25, 1989. 15. Wolffe A. P. Implications of DNA replication for eu-
12. Sogo, J. M., and R. A. Laskey. Chromatin replication karyotic gene expression. J. Ce//Sci. 99: 201-206,199l.

Physiology and Biophysics of the

100-m Sprint
Richard L. Summers

The IOO-m sprint is the most exciting of Olympic events, and the winner is often
called the “world’s fastest human.” The physiology of this race is unique and
involves our most primordial survival mechanisms. A computer simulation
predicts the theoretical “physio/ogicaP’- limits for human achievement in
this event.

P erhaps of all Olympic

controversy
contests, the 100-m
sprint evokes more interest, emotion,
than any other event. Lasting no
and
current record
9.84 s. Regardless
question
is held by Donovan
of these achievements,
still remains,
Bailey at

Is there really a human

the

more than -10 s, the ultimate of all sprints has limit to this fastest of all races? Although the
featured some of the greatest duels in Olympic answer may never be really answered, by com-
history. Who can forget the history of the famous bining our current knowledge of physiology and
“Chariots of Fire” race of the 1924 Olympics, the biophysics with the use of computer simulations,
Carl Lewis-Ben Johnson showdown in Seoul, we can estimate a theoretical boundary for the
Korea, or the Jesse Owens-Adolf Hitler confron- speed of man.
tation in the pre-WWII Berlin Games of 1936?
The winner is usually labeled the “world’s fastest Historical perspectives
human,” and many have become celebrities.
The race itself is brimming with tension and Predicting the limits of human running per-
excitement as modern-day athletes match speed, formance is not a new endeavor. Models con- “= . 1s fhere real/Y a
l

strutted based on trends of past record-breaking human limit to this

strength, and cunning reminiscent of Wild West
performances have been developed by Rumball fastest of all races?”
gunfighters. In the past, there have been many
experts who have speculated on the “human” and Coleman (12) and Ryder et al. (13) using
limits of the race. At one time, it was felt that no fitted logarithmic or polynomial expressions.
one could ever break the magical 1 O-s mark just Such models include many assumptions and are
as 4 min was the perceived physiological barrier not very interesting from a physiological per-
of the mile before Roger Bannister’s famous run spective. Others such as Perronnet and Thibault
for glory. However, as the athletes become (IO), Morton (9), and Ward-Smith (15) have
bigger and stronger and with better conditioning developed models using a more elegant ap-
and faster tracks, this barrier is now commonly proach. These models are concerned primarily
broken as we slowly chip away at the remainder with metabolic factors and thermodynamic prin-
of the IO s with record after new record. The ciples and are rich in content. However, these
models are designed to emphasize aerobic fac-
tors and races of longer distances. They also
R. L. Summers is Assistant Professor and Research Director,
Dept. of Emergency Medicine, University of Mississippi
neglect some important biomechanical Iimita-
Medical Center, 2500 North State St., Jackson, MS 392 7 6, tions that may be seen in sprinting. Most of these
USA. models have predicted ultimate speeds for the

0886-l 71497 $5.00 0 1997 Int. Union Physiol. Sci./Am. Physiol. Sot. News Physiol. Sci. l Volume 12 l June 1997 131
 100-m sprint at just a few tenths above 9 s. This The sprinter also faces atmospheric resistance
is still considerably faster than the present-day during the race (11). Indeed, it is well known that
records. sprinters are often faster in the thin air of higher
altitudes. This is one reason that so many sprint
Physics of sprinting and field records were broken at the Mexico City
Games. The same does not apply for the more
The Newtonian laws can be applied to sprint
aerobically dependent middle- and long-dis-
racing just as they can be used to analyze any
tance races. The equation for the resistive force
body in motion (2, 11, 14). In simple terms, the
of the air at a given Vfor the sprinter has been
amount of energy (K) required to move an object
determined to be
is related to the mass (M) and velocity (V) of the
body by the equation air resistance = l/2 CpAV2 (4

where C is drag coefficient, p is density of the air,

If a known amount of power is released from and A is cross-sectional area of the runner.
ATP splitting in the initial phases of maximal Mechanical efficiency (4) is defined as the
exertion, this can be integrated to give the ability to transform energy into mechanical work
amount of energy released at any point in time and takes the mathematical form of
during the race
mechanical efficiency = work/energy (5)

K= power (2) Physiology of sprinting

i
Before a model of the 100-m sprint is con-
This expression can then be used with Eq. I to structed, it is important to first consider what
calculate the velocity of the runner of known M physiological events occur during the first 10 s of
at any point during the race. maximal running exertion (7). It is well known
The sprinter meets with two main resistive that the initial energy used during extreme
“The anaerobic forces. Although very little energy is required to exertion comes primarily from anaerobic
sources can be maintain a velocity, a certain amount of energy sources. In fact, during the first 10 s of a sprint,
broken down into is required to keep the legs moving at the same -95% of all energy used is anaerobic. This is due
lactic and alactic speed as the rest of the body while they over- to the delay in the maximum activation of the
components. . . .” come the gravitational and frictional forces of the enzymes used in aerobic metabolism.
running surface (14). Such movements include The anaerobic sources can be broken down
both acceleration and deceleration forces of the into lactic and alactic components (6, 7). Al-
leg muscles through a complete stride. The though lactic acid is the end result of anaerobic
energy required during a single running stride glycolysis, it takes several seconds for this lactic
can again be determined by using Eq. I, with M system to become fully mobilized, and these
being the mass of the legs only. If the stride is energies are not available to the sprinter at the
lengthened so that fewer strides are needed to onset of the race. Alactic sources are available
cover the same distance in a given amount of immediately, but their rate of release of energy
time, then less energy is used to maintain the by the enzymatic splitting of ATP is a function of
velocity of the runner. Hence the amount of the available phosphagen stores. These restric-
power required for this resistive force can be tions do not represent a genetic or training
expressed in terms of the stride frequency by the limitation of the runner but rather represent the
equation physicochemical limitations of the enzymes in-
volved as determined by Michaelis-Menten ki-
netics. Hence the major energy supply in the
K resist = ‘I2 SM,,, V2 (3
initial phases of the 100 m comes from imme-
I
diately available ATP and phosphagen stores
where S is strides/s and MI,, is mass of the leg. available in the muscle and is termed the alactic
It is easy to see from Eq. 3 that an increase in process. These stores are used for the initial
the number of strides that a runner is required to power of acceleration and as a bridge between
take during the sprint will only serve to increase the power needed at the onset of running and the
the resistance to forward motion by the utiliza- initiation of the enzyme systems. All of this
tion of energy stores required for forward pro- alactic energy is expended rapidly and is ex-
pulsion (14). The longer the stride length, the hausted within -8 s. In an all-out sprint, max-
fewer strides that a runner will take to cover the imal power output is reached nearly instanta-
same amount of ground. neously and is sustained at a constant rate for

132 News Physiol. Sci. l Volume 12 l June 1997

 -6-7 s. This is why many sprinters reach
peak velocity
maximal velocity within the first 3-4 s and 1
thereafter begin to lose speed in the last third of
the 100 m when lactic acid-producing glycolysis
becomes the major energy source (7). Although -F-
lactic acid slows muscle contraction and re- deceleration phase
duces power, it is unlikely that there is enough
buildup of this acid within 10 s to be a significant
factor (7). Rather, it is the near exhaustion of the acceleration phase
phophagens as an energy source that leads to the
slowdown. - 0 2 4 6 8 10
At the 4- to 6-s mark of the sprint, glycolytic Time (seconds)
enzyme systems become significantly activated
FIGURE 1. Theoretical time-velocity profile for a 70-kg
and the power available from anaerobic lactic sprinter.
acid-producing sources rises rapidly. This en-
ergy source reaches a peak at w-30-40 s and
derived energy of the leg muscles and is
therefore is probably more important at ZOO and
resisted by the atmospheric drag and the “0
l l the sprinter Who
400 m (10). is able to reach maxi-
energy required to maintain the stride (5, 10,
Because there is little chance of running out of ma/ velocity the earli-
11, 14).
energy over such a short distance in a race lasting est has the greatest
2) Aerobic metabolism contributes essentially
only IO s, the sprinter who is able to reach likelihood of winning
none of the energy required in the 100-m
maximal velocity the earliest has the greatest the race.”
sprint (7).
likelihood of winning the race. Because the
3) The anaerobic metabolism is made up of both
energy of acceleration is from the alactic ATP-
an alactic portion from phosphagen sources
creatine phosphate process, the rate of release of
and a lactic acid-producing glycolytic path-
this energy (the slope of the power-time rela-
way. The peak rate of anaerobic energy
tionship for this system during maximal exertion)
release is directly related to the anaerobic
represents the most important factor in deter-
capacity, and at any point in time, only 50%
mining a runner’s ability to reach top speed
of the available 15,000 J of alactic energy can
quickly (6, 10).
be utilized until it is totally exhausted after
-7-8 s of all-out exertion (6-8).
The model
4) Glycolytic pathways become functional after
A mathematical computer model combining -3-6 s of maximal exertion and provide
many of the biological and physical processes energy at a rate of 500-I ,000 J/s until a peak
involved in running the 100-m sprint was con- contribution is realized -30-40 s into the
structed with the use of VisSim, an object- sprint (6).
oriented modeling software package. The model 5) The biomechanical efficiency of the muscles
was then solved with a Pentium-based personal used in running is -5O-60% in highly
computer using a second-order Runga Kutta trained sprinters. Although there have been
integration methodology. The configuration of reports of efficiency of running of up to 70%,
the model drew from several well-validated the typical individual maintains a biome-
models of running kinetics, bioenergetics, and chanical efficiency of -25%. However, it
metabolism while it centered on a physics-based would be expected that Olympic athletes
representation of running motion and biome- would increase this to -57% with optimal
chanics. The model uses parameter values that technique (3, 4, 8).
have been established in the literature from
observations made with modern world-class Computer simulations and discussion
sprinters. An assortment of both physical and
Computer-simulation studies of a 100-m
metabolic variables is utilized so that the model
sprint were performed using the model under a
can be tested over a wide range of anthropo-
variety of parametric conditions. In Fig. 1, the
morphic and atmospheric conditions. The rela-
model displays the theoretical time-velocity pro-
tive contributions of kinetic and resistive forces
file for a 7O-kg sprinter. This configuration of a
are brought together to determine the velocity of
rapid rise to peak velocity within the first 3 s with
a runner of a predetermined mass. The major
a gradual slowing during the latter third of the
assumptions of the model are as follows:
race is consistent with that typically seen in
I) Forward velocity of the spri rter is determined well-trained athletes (15). This slowing phenom-
from the anaerobically (a attic and lactic) ena is also well known in track circles and is the

News Physiol. Sci. l Volume 12 l June 1997 133

 faster acceleration
c -
important throughout the race
s/o wer finish

2 4 6 8 10 2 4 6 8 10
Time - seconds Time - seconds

increasing efficiency

small effect at the finish

faster acceleration

0 2 4 6 8 10 0 2 8 10
Time - seconds Time” secoids
FIGURE 2. Effects of various factors on time-velocity profile. A: muscle mass. B: stride length. C: mechanical efficiency.
D: altitude.

object of many race strategies. Carl Lewis has with the longer (and fewer) strides may not
notoriously poor starts out of the blocks and possess the power for quick acceleration, but
attributes much of his success at 100 m to his they are clearly at an advantage in the latter third
ability to relax and allow his momentum to carry of the race when the energy-consuming resistive
him through his alactic energy expenditure to the forces of accelerating and decelerating the legs
finish line (5). with each stride will erode the maximal velocity
The time-velocity curve is the signature profile obtainable. Sprinters with longer legs, such as
for a sprinter, depicting the subtleties of his or her Carl Lewis, often pull past the field in the last
strengths and physical characteristics. It can also 20 m of the race by relaxing and extending their
be used to describe the relative effects of differ- stride. On the other hand, it has also been shown
ent factors on all phases of the race and will be that for any given individual there is an optimum
used as such in the present analysis. The effects stride length that provides the lowest energy
of these varying factors on the time-velocity consumption (1, 5). In general, the overall en-
profile are combined in Fig. 2. ergy consumption decreases with stride length
By varying the weight (muscle only) of the until the increase in stride becomes unnatural
athlete over 1 O-kg increments, a pattern emerges and requires effort. At this point, there is an
that can be analyzed (Fig. 2A). Even though increase in energy expenditure in running.
increasing the muscle mass of the legs provides Hence naturally long strides are highly depen-
“The mechanical effi-
more power and readily available ATP stores dent on the physique of the athlete.
ciency of muscle is
during the initial phases of the race (faster The mechanical efficiency of muscle is some-
something that can
acceleration), the additional body weight re- thing that can be modified with training (3).
be modified with
quires more kinetic energy for motion and po- Increasing the efficiency from the low range of
training. . . . ”
tentiates the resistance effects of the atmosphere 0.25 to a high of 0.75 would have a significant
and stride biomechanics as they dominate in the effect on the time-velocity curve. Figure 2C
later stages of the race (terminal velocity). demonstrates how this change in muscle effi-
Shorter, stocky sprinters with powerful leg mus- ciency can significantly influence sprint times.
cles are often the best starters. Indeed, the Because training and technique have been dem-
steroid-built muscle mass of Ben Johnson made onstrated to have a significant effect on the
him one of the fastest starters in the history of the mechanical efficiency of running, this simula-
event. tion demonstrates the importance of coaching
If the stride length is varied, as in Fig. 2B, an for the track athlete rather than relying on innate
interesting result is noted. The tall lanky sprinters skill alone. Many sprinters have dramatically

134 News Physiol. Sci. l Volume 12 l June 1997

 TABLE 1. Comparison of predicted limits
* 100 for 100-m sprint
k
z 80 Summers Morton Ryder et Peronnet and
is (this study) (9) al. (13) Thibault (IO)
I 60
Time, s 9.3 9.15 9.34 9.37
z
5 40
Y
in
5 20
when compared with coaching and training in
0 the success of the sprint athletes. By observing
the simulation studies as a whole, a pattern
0 1 3 4 6 7 9
Time - seconds emerges.
Size and relative muscle mass do not neces-
FIGURE 3. Time limits for optimum sprinter.
sarily appear to be beneficial as a factor alone.
Hence the contributions of the extremes of bulk
changed their times for better or worse by a
weight training and steroid abuse are of ques-
change in their running style.
tionable benefit for the sprinter. However, this
There is often considerable controversy sur-
does assume that the ratio of fast- to slow-twitch
rounding records that were established at loca-
fibers (a primarily genetically determined char-
tions well above sea level. There is little doubt
acteristic) is the same for all the sprinters. Be- “Conserving energy is
that the reduced air density and smaller resistive
cause the ratio can be changed to a certain not usually an impor-
force of the higher altitudes are advantageous for
degree by training, this could change the relative tan t consideration in
the sprinter. The computer simulation study in
rate of release of the energy from the muscle for the short sprints. . . . N
Fig. 20 illustrates the effect of altitude on the
use in the early phases of the sprint (3).
outcome of the 100-m race when all other
The study suggests that stride length is of major
factors remain the same. The relative lack of
importance in determining the speed of the
oxygen in mountainous areas makes the task of
sprinter. A runner’s stride has both natural and
middle- and long-distance running more diffi-
learned components. Even though the body
cult. However, the sprinter relies solely on
inherently seeks a gait that will optimize the
anaerobic metabolism. This atmospheric effect
body energetics and conserve energy, this may
was obvious in the 1968 Mexico Olympics
not necessarily produce the longest possible
where many sprint and field records were bro-
stride (1). Conserving energy is not usually an
ken. It should be noted from Eq. 4 that this
important consideration in the short sprints
phenomenon would be potentiated in the
where all energy is used maximally anyway.
sprinter with less body mass. In fact, the 100-m
Training that stretches muscles and tendons
world record at altitude was set in the mountains
and lengthens the natural stride may be more
of Colorado by Calvin Smith, one of the physi-
advantageous.
cal ly smaller world-class sprinters. Although our
Mechanical efficiency of the runner involves
study agrees with the conclusion that altitude
both bioenergetics and biomechanics and ap-
does affect sprint times, this difference appears to
pears from our study to be very important in
be relatively small when compared with the
determining the overall speed. Of all the factors,
other factors involved.
this is the one where athletic training, diet, and
The current world record holder in the 100 m,
coaching may have the greatest influence. A
Donovan Bailey, possesses both a large muscle
smooth start out of the block matched with a
mass and a good stride length. One of the goals
continuing efficient concert of motion of arms
of the simulation was to determine the theoret-
and legs throughout the race is the mark of a
ical limits for the race based on our current
well-trained athlete.
knowledge of physiology and biophysics. The
ideological optimum sprinter, with a realistic
physique, working at a practical mechanical Conclusion
efficiency of 57%, would be expected to run the Although this theoretical boundary of 9.3 s for
100 m in 9.3 s at sea level using the current the 100 m may seem unobtainable by today’s
model (Fig. 3). This is 0.54 tenths of a second standards, it should be remembered that the
faster than the present-day record of 9.84 s. This present record would have seemed equally
result compares with the predictions of a number unlikely only 20 years ago. The results of the
of other modelers who have searched for this analysis are limited by the assumptions of the
limit (Table I). model and our current knowledge of running
The relative importance of natural physique physiology. As the modern athlete continues to
and genetic characteristics is often debated emerge as a specialized entity, the current con-

News Physiol. Sci. l Volume 12 l June 1997 135

 straints of anaerobic capacity and sprint tech- 5. Lewis, C., and J. Marx. inside Track. New York: Simon
nique as defined by mechanical efficiency may and Schuster, 1990.
6. Margaria, R., P. Cerretelli, and F. Mangili. Balance
also evolve. Clearly, the simulation studies sug-
and kinetics of anaerobic energy release during
gest that training could have a dominant effect strenuous exercise in man. J. Appl. Physiol. 19:
on sprint times. Hence the model would predict 623-628,1964.
an even faster time. Whatever the determined 7. McArdle, W. D., F. I. Katch, and V. L. Katch. Exercise
maximum speed of humans, reaching this limit Physiology(3rd ed.). Malvern, PA: Lea & Febiger, 1991.
8. Medbo, J. I., and S. Burgers. Effects of training on the
will ultimately require that a number of disparate
anaerobic capacity. Med. Sci. Sports Exercise 22: 501-
variables of genetics and training and environ- 507,1990.
mental factors come together at precisely the 9. Morton, R. H. A three component model of human
same time (<IO s) to create the optimal condi- bioenergetics. J. Math. Biol. 24: 451-466, 1986.
tions within a single individual sprinter. 10. Peronnet, F., and C. Thibault, Mathematical analysis of
running performance and world running records.
J. Appl. Physiol. 67: 453-465, 1989.
This work was supported by National Heart, Lung, and 11. Peronnet, F., G. Thibault, and D. L. Cousineau. A
Blood Institute Grant HL-5 1971. theoretical analysis of the effect of altitude on running
performance. J. Appl. Physiol. 70: 399-404, 1991.
12. Rumball, W. M., and C. E. Coleman. Analysis of running
References and the prediction of ultimate performance. Nature 228:
184-185,197O.
1. Cavanagh, P. R., and R. Kram. Stride length in distance 13. Ryder, W. H., H. J. Carr, and P. Herget. Future perfor-
running: velocity, body dimensions and added mass mance in foot racing. Sci. Am. 234: 109-I 19, 1976.
effects. Med. Sci. Sports Exercise 21 : 467-479, 1989. 14. Van lngen Schenau, G. J., J. J. de Koning, and G. de
2. Cromer, A. H. Physics for the Life Sciences (2nd ed.). Groot. Optimisation of sprinting performance in run-
New York: McGraw-Hill, 1977. ning, cycling and speed skating. Sports Med. 17: 259-
3. Di Prampero, P. E. Energetics of muscular exercise. Rev. 275, 1994.
Physiol. Biochem. Pharmacol. 89: 143-222, 1981. 15. Ward-Smith, A. J. A mathematical theory of running,
4. Kyrolainen, H., P. V. Komi, and A. Belli. Mechanical based on the first law of thermodynamics, and its
efficiency in athletes during running. Stand. /. Med. Sci. application to the performance of world-class athletes.
Sports 5: 200-208, 1995. J. Biomech. 18:337-349, 1985.

GABA, Receptors: A Novel Receptor

Family with Unusual Pharmacology
Enrico Cherubini and Fabrizio Strata

GA BA,-receptor channels constitute a novel class of structurally defined receptors

that are composed by the recently discovered p-subunits. They conduct chloride
ions, are insensitive to both bicuculline and baclofen, and are blocked
by picrotoxin. They are present in the hippocampus during development
and in the retina where they play a crucial role in regulating
visual processing.

“The GA BA, receptor

is competitively
T he main
aminobutyric
inhibitory

uted in the vertebrate

neurotransmitter,
acid (GABA), is widely distrib-
central nervous system
y- the GABA,
channels
receptor
(through
is coupled
GTP binding
to cationic
proteins and
second messengers) and belongs to the G pro-
antagonized by (CNS). It inhibits neuronal firing and stabilizes tein-linked receptor family. The GABA, receptor
bicuculline. . . .‘I the resting potential by activating two distinct is competitively antagonized by bicuculline and
classes of receptor types named GABA, and contains allosteric modulatory sites for a variety
GABA,. The CABA, receptor is a protein that of ligands; some such as barbiturates and ben-
gates chloride channels and belongs to the zodiazepines are of therapeutic interest. The
ligand-gated ion-channel superfamily, whereas GABA, receptor is activated by baclofen, antag-
onized by phaclofen, 2-hydroxysaclofen, and
E. Cherubini and F. Strata are in the Biophysics Laboratory
CGP-35348 or-55845A, and insensitive to drugs
and lstituto Nazionale Fisica della Materia Unit, Interna-
tional School for Advanced Studies (SISSA), Via Beirut 2-4, that modulate GABA, receptors. The GABA,
340 I4 Trieste, Italy. receptor is a heteropentameric structure com-

136 News Physiol. Sci. l Volume 12 l June 1997 0886-I 714/97 $5.00 0 1997 Int. Union Phvsiol. Sci./Am. Phvsiol. Sot.