## TL;DR Because density, muscle stress, and metabolic power per...
A beautiful multi-scale connection: The scaling depends ultimate...
#### Mass Comes from Volume Mass is essentially **density × volu...
Muscles produce force by contracting fibers. Imagine a muscle sh...
Galileo noticed that: - **Strength ∝ cross-sectional area** - *...
One beautiful consequence is that the paper predicts a common locom...
#### Why Large Animals Break the Law For large animals another...
How fast do living organisms move: Maximum speeds from bacteria
to elephants and whales
Nicole Meyer-Vernet
a)
LESIA, Observatoire de Paris, CNRS, UPMC, Universit
e Paris Diderot, 92195 Cedex Meudon, France
Jean-Pierre Rospars
b)
INRA, UMR 1392, Institut d’Ecologie et des Sciences de l’Environnement de Paris, 78000 Versailles, France
(Received 30 October 2014; accepted 29 March 2015)
Despite their variety and complexity, living organisms obey simple scaling laws due to the
universality of the laws of physics. In the present paper, we study the scaling between maximum
speed and size, from bacteria to the largest mammals. While the preferred speed has been widely
studied in the framework of Newtonian mechanics, the maximum speed has rarely attracted the
interest of physicists, despite its remarkable scaling property; it is roughly proportional to length
throughout nearly the whole range of running and swimming organisms. We propose a simple
order-of-magnitude interpretation of this ubiquitous relationship, based on physical properties
shared by life forms of very different body structure and varying by more than 20 orders of
magnitude in body mass.
V
C
2015 American Association of Physics Teachers.
[http://dx.doi.org/10.1119/1.4917310]
I. INTRODUCTION
A fundamental property of many living organisms is au-
tonomous locomotion. On Earth, which is a rocky planet of
surface temperature T 300 K with liquid water and an
atmosphere, animals can run, swim, and/or fly. Each individ-
ual has a preferred speed of locomotion which is determined
by its size and by dynamical constraints,
1,2
depending on the
means of locomotion and the ambient conditions. Newtonian
mechanics tells us that a walker of size L approximated as an
inverted pendulum in the Earth’s gravitational field g, moves
at angular frequency ðg=LÞ
1=2
; hence its preferred speed V is
of order of magnitude the step length L times the fre-
quency: V ðLgÞ
1=2
=2p. More elaborate arguments have
been used to propose scalings for various means of locomo-
tion, depending on such factors as the gravity and the density
of the surrounding medium.
37
However, when living organisms are driven by circum-
stances to move as fast as possible, they may increase their
speed above the preferred speed. As human beings, we can
increase our speed by one order of magnitude by running.
The fastest human sprint speed on record is 12.2 m/s for
Usain Bolt (size L ¼ 1:96 m and mass M ¼ 86 kg in 2009),
8
whereas the recent 50 m swimming record of Florent
Manaudou (L ¼ 1:99 m, M ¼ 99 kg) corresponds to an aver-
age speed of 2.5 m/s. For comparing organisms of widely
different sizes, it is more appropriate to express the speed in
terms of the body length. The above values yield 6.1 and 1.2
lengths per second, respectively. Near the lower extreme of
the size range, the 2.5-lm-long bacterium Bacillus subtilis
can swim at 15 lm/s, or 6 lengths/s, strikingly close to Usain
Bolt’s running performance. A 4-mm-long ant runs at
60 mm/s (15 lengths/s), whereas a 2.1-m ostrich runs at
23 m/s (11 lengths/s).
4
These examples reflect a ubiquitous
property of living organisms: the maximum speed of running
and swimming lies between 1 and 100 lengths per second, in
an overall mass range covering nearly 20 orders of magni-
tude, as first noted by Bonner in a classic book.
9
Indeed, while maximum speeds V
max
and lengths L vary
by nearly 7 orders of magnitude (and mass by three times
more), maximum relative speeds V
max
=L remain constant at
ten per second with an accuracy of a factor of ten,
4,911
even
though different scalings hold over narrower mass
ranges.
12,13
This large-scale pattern, which holds for organ-
isms as diverse in overall structure, means of locomotion,
and surrounding medium as eight-legged mites and two-
legged ostriches, from small bacteria to large mammals, has
apparently not been appreciated by the physics community
and requires a basic explanation. A solution was proposed by
Dusenbery,
11
but being based on viscous drag it only applies
to the small range of organisms swimming at low Reynolds
numbers.
In the present paper, we illustrate the approximate linear
relation between maximum speed and body size for a large
number of running and swimming species from micro-
organisms to the largest mammals (Sec. II), and we propose a
simple order-of-magnitude interpretation, based on three ba-
sic properties of living beings that constrain their performan-
ces: their density q, the applied force per unit cross-sectional
area r, and the maximum rate of energy consumption per unit
mass b
M
. These three quantities are known to be roughly in-
dependent on size over virtually the full mass range of mov-
ing species, as summarized in Sec. III. We shall briefly
discuss in Sec. IV two exceptions to the large-scale relation
V
max
=L 10 s
1
: very large organisms, whose maximum
speed tends to level off,
4,12
and flyers.
The huge mass range studied enables us to estimate orders
of magnitude by neglecting specific details that are important
in scaling studies over narrower mass ranges. In this sense,
the present paper can be viewed as an exercise at the bound-
ary between comparative zoology and order-of-magnitude
physics inspired by Victor Weisskopf’s physics courses.
14,15
As usual, the symbol means that two quantities are equal
to within one order of magnitude or so, whereas means
equal within a factor of two or so. Unless otherwise stated,
units are SI.
II. EMPIRICAL RESULTS
Figure 1 shows the measured maximum relative speed
V
max
=L as a function of mass for running and swimming,
from micro-organisms to the largest mammals. The mass
719 Am. J. Phys. 83 (8), August 2015 http://aapt.org/ajp
V
C
2015 American Association of Physics Teachers 719
This article is copyrighted as indicated in the article. Reuse of AAPT content is subject to the terms at: http://scitation.aip.org/termsconditions. Downloaded to IP:
145.238.153.34 On: Wed, 22 Jul 2015 11:49:06
range goes from mites to the African bush elephant for run-
ning and from micrometer-sized bacteria to whales for swim-
ming. Almost all the data lie in the range 1 < V
max
=L < 100.
This range is remarkably narrow compared to the 10
20
-fold
variation in body mass and confirms the striking constancy
of the maximum relative speed first noted by Bonner.
9
The
human world records for running and swimming are plo tted
as asterisks (red and blue, respectively). Both lie in the lower
range of animal running and swimming relative speeds,
respectively, illustrating the low rank of human beings in the
animal world for sprinting and swimming. Nevertheless,
these records still lie within an order of magnitude of the
scaling V
max
=L ¼ 10 s
1
.
Figure 1 also suggests that the maximum speed tends to
level-off for large masses,
4
a question that we shall discuss
in Sec. IV. We have not plotted flying speeds, which follow
a different scaling law (see Sec. IV).
4
III. ESTIMATION OF MAXIMUM SPEED
In order to propose a basic interpretation of the observed
scaling, let us consider the three universal properties of
living species which constrain their maximum speed of loco-
motion: mass density q, applied force per unit cross-
sectional area r, and maximum power per unit mass b
M
(maximum metabolic rate).
A. Three ubiquitous properties of living species
First, the mass density of organisms is roughly that of
liquid water, on which life on Earth is based
q 10
3
kg m
3
: (1)
Second, the applied force per unit cross-sectional area of
tissue
6,17
is of order of magnitude
r 2 10
5
Nm
2
; (2)
from micro-organisms to the largest animals.
18
This is an
example of the rule dating back to Galileo that the strength
of an object is proportional to its cross-section. Here, Eq. (2)
is not the resistance to fracture, the so-called tensile strength,
but the average active tension applied by organisms for their
locomotion. This tension has a similar value for all organ-
isms because it is based on biological molecular motors of
similar basic properties. Biological motors are molecules
converting chemical energy into mechanical energy via a
conformational change in their molecular structure.
19
This
3-dimensional structure is held together by non-covalent
bonds, with the typical free energy
W
0
10 k
B
T; (3)
which prevents their destruction by thermal agitation, and
their typical size is
20
a
0
e
2
=4p
0
W
0
6nm; (4)
despite the complexity of electrostatic interactions within
large molecules.
21
Basically, a molecular motor uses an
energy W
0
for moving by one “step” via a change in 3-D
structure, so that the “step” length is a
0
. The elementary
force is thus
F
0
W
0
=a
0
7 pN (5)
over an equivalent cross-section area whose order of magni-
tude is a
2
0
, so that the force per unit cross-section area is
r F
0
=a
2
0
W
0
=a
3
0
: (6)
Substituting Eqs. (3) and (4) into Eq. (6) yields Eq. (2).
This order of magnitude holds for muscles of animals,
which are made of filaments containing hundreds of elemen-
tary motors (myosin), as well as for the moving appendages
of micro-organisms.
18,22,23
Third, consider the power available. Transport of heat and
nutrients takes place across surfaces, which are expected to
scale as the square of size, and thus to vary with body mass
as M
2=3
; therefore, the energy consumption rate of living
beings (the so-called “metabolic rate”) per unit mass is
expected to scale as M
2=3
=M ¼ M
1=3
. Reality is more com-
plicated because body shape and structure change with size,
so that different scalings are observed
24
with an exponent
closer to 1=4 than to 1 =3. After decades-long controver-
sies,
25,26
it has been shown, albeit rarely appreciated in the
physics community, that the basal metabolic rate per unit
mass remains roughly constant across life forms.
27,28
More
precisely, for the vast majority of organisms it remains
within a 30-fold range,
29
which is remarkably narrow com-
pared with the 10
20
-fold body mass range concerned. Since
Fig. 1. Maximum relative speed versus body mass for 202 running species (157 mammals plotted in magenta and 45 non-mammals plotted in green), 127
swimming species and 91 micro-organisms (plotted in blue). The sources of the data are given in Ref. 16. The solid line is the maximum relative speed
[Eq. (13)] estimated in Sec. III. The human world records are plotted as asterisks (upper for running and lower for swimming). Some examples of organisms of
various masses are sketched in black (drawings by Franc¸ois Meyer).
720 Am. J. Phys., Vol. 83, No. 8, August 2015 N. Meyer-Vernet and J.-P. Rospars 720
This article is copyrighted as indicated in the article. Reuse of AAPT content is subject to the terms at: http://scitation.aip.org/termsconditions. Downloaded to IP:
145.238.153.34 On: Wed, 22 Jul 2015 11:49:06
we wish to estimate the maximum speed, the relevant prop-
erty is not the basal metabolic rate but rather the maximum
metabolic rate. The order of magnitude of this parameter has
been shown to be roughly constant, too, when scaled to the
mass, with the value
b
M
2 10
3
Wkg
1
(7)
per unit of working tissue.
27,30,31
B. Maximum relative speed
If the maximum relative speed V
max
=L only depends on
the parameters q, r, and b
M
, dimensional analysis can be
used to deduce its scaling. In terms of the three dimensions
½M , ½L; ½T, the density scales as
q M ½L
3
: (8)
Since r is a force (M½L½T
2
) per unit cross-section
(L
2
), it sca les as
r M ½L
1
½T
2
; (9)
and since b
M
is a power (M ½L
2
½T
3
) per unit mass, it
scales as
b
M
L
2
½T
3
: (10)
Therefore, since V
max
=L T
1
, we deduce
V
max
=L / b
M
q=r: (11)
In order to make a quantitative estimate, let us go a step
further than dimensional analysis. First, consider running and
swimming of animals beyond the micro-organism range. At
zero order, both means of locomotion can be considered as a
cyclic process (of frequency f) in which an organism of length
L moves by one “step” of length L during each cycle, by
contracting muscles. Consider an organism of cross-section S
and length L:
its mass is M qSL,
moving by one step of length L by applying the force
rS requires the energy per unit mass w rSL=M r=q,
since f steps per second consume the energy fw per unit
mass, which must be smaller than b
M
, the maximum step
rate is f
max
b
M
=w b
M
q=r.
The maximum speed equals the step length L times the
maximum step rate f
max
, whence
V
max
=L f
max
b
M
q=r: (12)
Substituting Eqs. (1), (2) , and (7) into Eq. (12) yields
V
max
=L 10 s
1
; (13)
which is the large-scale relation mentioned in the
Introduction.
Consider now micro-organisms. They move by rotating or
undulating flagella, cilia, or pili, which are operated by mo-
lecular motors as are the muscles of larger organisms, even
though the number of motors is much smaller for micro-
organisms. In this case, it is more enligh tening to consider
the microscopic level. During one period of rotation or undu-
lation,
23,32
a micro-organism of length L moves along a dis-
tance L using energy W
0
[given in Eq. (3)] per molecular
motor. With f cycles per second, the power spent is fW
0
.
For a motor of size a
0
given in Eq. (4) and mass qa
3
0
, the
power cannot exceed the maximum metabolic rate b
M
qa
3
0
.
This yields f b
M
qa
3
0
=W
0
, whence
V
max
=L b
M
qa
3
0
=W
0
: (14)
With n motors, both the numerator and the denominator of
Eq. (14) are multiplied by n, which does not change the
result. Since from Eq. (6) r W
0
=a
3
0
, Eq. (14) is equivalent
to Eq. (12).
Hence, both micro-organisms and larger animals should
have a similar maximum relative speed for running and
swimming, given by Eqs. (12)(13), in agreement with the
data plotted in Fig. 1.
IV. CONCLUDING REMARKS
There are two exceptions to the scaling derived above: fly-
ing species and ver y large organisms.
Flying is outside the scope of our simplified model
because in that case the muscles essentially govern wing
flapping, and this frequency does not yield the total relative
speed of the organism. In addition, air drag represents the
dominant constraint at large flying speeds.
4
Consider now large running and swimming organisms, for
which V
max
=L tends to decrease (Fig. 1), even though the
data do not lie below one order of magnitude of the scaling
(13) except for the largest animal. Several effects become
important at high speeds, such as friction and excess heat
production. However, Fig. 1 suggests a similar trend for run-
ning and swimming, which points to a more fundamental li-
mitation, independent of the surrounding medium.
Let us consider an organism of cross-section S and length
L, as in Sec. III B, and approximate the locomotion as a peri-
odic motion of legs (for running) or tail (for swimming) of
length L. The maximum frequency is constrained not only
by the power available, as considered in Sec. III B, but also
by the maximum angular acceleration that muscles can pro-
vide. With the torqu e C rSL and moment of inertia
I ML
2
qSL
3
, the angular acceleration d
2
h=dt
2
C=I is
constrained by
d
2
h=dt
2
r=ðqL
2
Þ: (15)
Integrating Eq. (15) twice yields the order of magnitude of
the time for the append age to be accelerated up to a fixed
angle h:
t Lðqh= rÞ
1=2
: (16)
Setting h 1 in Eq. (16) yields the frequency f
1=t ðr=qÞ
1=2
=L and therefore the upper limit of the maxi-
mum speed
V
max
ðr=qÞ
1=2
: (17)
Hence, the value of V
max
=L in Eq. (12) can only hold for
L ðr=qÞ
1=2
=ðb
M
q=rÞ¼ðr=qÞ
3=2
=b
M
: (18)
721 Am. J. Phys., Vol. 83, No. 8, August 2015 N. Meyer-Vernet and J.-P. Rospars 721
This article is copyrighted as indicated in the article. Reuse of AAPT content is subject to the terms at: http://scitation.aip.org/termsconditions. Downloaded to IP:
145.238.153.34 On: Wed, 22 Jul 2015 11:49:06
Substituting Eqs. (1), (2), and (7) into Eq. (18) yields L 1: 4
m. This limitation prevents larger organisms from following
Eq. (12), which suggests that the maximum speed should
increase linearly with body length onl y up to (approxi-
mately) meter-sized organisms, in agreement with Fig. 1.
Consider the blue whale (blue point at M 1:5 10
5
kg),
which lies below one order of magnitude of the scaling
(12)(13); with its length L 26 m, Eq. (17) yields
V
max
=L 0:5s
1
, a limit close to the observed value plotted
in Fig. 1.
Finally, one should be reminded that in the spirit of this
paper, Eqs. (12) and (17) are order-of-magnitude results.
Because of the huge diversity of organisms and sizes, we
have ignored the specific methods of locomotion, using dras-
tic approximations for the applied forces, cross-sections and
distances involved, as well as approximating by unity the ef-
ficiency of energy conversion and the proportion of active
tissue. The numerous correction factors tend to cancel out in
the final order-of-magnitude result.
In conclusion, we explain the ubiquity of the maximum
relative speed at about ten lengths per second for running or
swimming, from bacteria to large mammals, by the ubiquity
of the density, the applied force (per unit cross-sectional
area), and the maximum metabolic rate (per mass of active
tissue). The maximu m absolute speed is limited by the maxi-
mum acceleration that mu scles can provide, which may
explain why animals larger than the ostrich do not move
faster.
ACKNOWLEDGMENTS
The authors thank three anonymous reviewers for helpful
comments and suggestions. The authors are grateful to
Franc¸ois Meyer, who did the original drawings superimposed
on Fig. 1.
a)
Electronic mail: nicole.meyer@obspm.fr
b)
Electronic mail: jean-pierre.rospars@versailles.inra.fr
1
J. B. S. Haldane, “On being the right size,” in The world of mathematics,
edited by J. R. Newman (Simon and Schuster, London, 1956), Vol. 2,
Chap. 16. D. W. Thompson, “On magnitude,” ibid., Chap. 17.
2
A. V. Hill, “The dimensions of animals and their muscular dynamics,” Sci.
Prog. London 38, 209–230 (1950).
3
H. Lin, “Newtonian mechanics and the human body: Some estimates of
performance,” Am. J. Phys. 46, 15–18 (1978).
4
T. A. McMahon and J. T. Bonner, On Size and Life (Scientific American
Library, New York, 1983).
5
K. Schmidt-Nielsen, Scaling, Why is Animal Size so Important?
(Cambridge U.P., Cambridge, New York, 1984).
6
C. J. Pennycuick, Newton Rules Biology, A Physical Approach to
Biological Problems (Oxford U.P., Oxford, New York, 1992).
7
A. Bejan, “The constructal-law origin of the wheel, size, and skeleton in
animal design,” Am. J. Phys. 78, 692–699 (2010).
8
J. J. Hern
andez G
omez et al., “On the performance of Usain Bolt in the
100-m sprint,” Eur. J. Phys. 34, 1227–1233 (2013).
9
J. T. Bonner, Size and Cycle: An Essay on the Structure of Biology
(Princeton U.P., Princeton, New Jersey, 1965). See also Refs. 4 and 10.
10
J. T. Bonner, Why Size Matters, From Bacteria to Blue Whales (Princeton
U.P., Princeton & Oxford, 2006).
11
D. B. Dusenbery, Living at Microscale, The Unexpected Physics of Being
Small (Harvard U.P., Cambridge, Massachusetts & London, England,
2009).
12
T. Garland, Jr. “The relation between maximal running speed and body
mass in terrestrial mammals,” J. Zool., London 199, 157–170 (1983).
13
J. Iriarte-Diaz, “Differential scaling of locomotor performance in small
and large terrestrial mammals,” J. Exp. Biol. 205, 2897–2908 (2002).
14
V. F. Weisskopf, “Of atoms, mountains and stars: A study in qualitative
physics,” Science 187, 605–612 (1975).
15
V. F. Weisskopf, The Privilege of Being a Physicist (W. H. Freeman, New
York, 1989).
16
N. Meyer-Vernet and J.-P. Rospars, “Maximum relative speeds of living
organisms: Why do bacteria perform as fast as ostriches?” (in
preparation).
17
S. Medler “Comparative trends in shortening velocity and force production
in skeletal muscles,” Am. J. Physiol. Reg. Integr. Comp. Physiol. 283,
R368–R378 (2002).
18
J.-P. Rospars and N. Meyer-Vernet, “Force per unit area from molecules
to muscles: A general property of biological motors” (in preparation).
19
M. E. Fisher and A. B. Kolomeisky, “Molecular motors and the forces
they exert,” Physica A 274, 241–266 (1999).
20
See, for example, J. Zhang, “Protein-length distributions for the three
domains of life,” Trends Gen. 16(3), 107–109 (2000).
21
B. Honig and A. Nicholls, “Classical electrostatics in biology and chem-
istry,” Science 268, 1144–1149 (1995).
22
M. Schliwa and G. Woehlke, “Molecular motors,” Nature 422, 759–765
(2003).
23
H. C. Berg, “The rotary motor of bacterial flagella,” Annu. Rev. Biochem.
72, 19–54 (2003).
24
H. Hoppeler and E. R. Weibel, “Scaling functions to body size: Theories
and facts,” J. Exp. Biol. 208, 1573–1574 (2005).
25
P. S. Agutter and D. N. Wheatley, “Metabolic scaling: Consensus or con-
troversy,” Theor. Biol. Med. Modelli. 1, 13 (2004).
26
J. Kozlowski and M. Konarzewski, “Is West, Brown and Enquists model
of allometric scaling mathematically correct and biologically relevant?,”
Funct. Ecol. 18, 283–289 (2004).
27
A. M. Makarieva, V. G. Gorshkov, and B.-L. Li, “Energetics of the small-
est: Do bacteria breathe at the same rate as whales,” Proc. R. Soc. B 272,
2219–2224 (2005).
28
A. M. Makarieva, V. G. Gorshkov, and B.-L. Li, “Mean mass-specific
metabolic rates are strikingly similar across life’s major domains:
Evidence for life’s metabolic optimum,” Proc. Natl. Acad. Sci. USA 105,
16994–16999 (2008).
29
This range is compatible with the different scalings observed in narrow
mass ranges; see Ref. 28.
30
C. J. Pennycuick and M. A. Rezende, “The specific power output of aero-
bic muscle, related to the power density of mitochondria,” J. Exp. Biol.
108, 377–392 (1984).
31
E. R. Weibel and H. Hoppeler, “Exercise-induced maximal metabolic rate
scales with muscle aerobic capacity,” J. Exp. Biol. 208, 1635–1644
(2005).
32
B. de Lima Bernardo and F. Moraes, “Simplified model for the dynamics
of a helical flagellum,” Am. J. Phys. 79, 736–740 (2011).
722 Am. J. Phys., Vol. 83, No. 8, August 2015 N. Meyer-Vernet and J.-P. Rospars 722
This article is copyrighted as indicated in the article. Reuse of AAPT content is subject to the terms at: http://scitation.aip.org/termsconditions. Downloaded to IP:
145.238.153.34 On: Wed, 22 Jul 2015 11:49:06

Discussion

#### Mass Comes from Volume Mass is essentially **density × volume**: \[ M = \rho V \] where: - \(M\) = mass - \(\rho\) = density - \(V\) = volume For most organisms, density is roughly constant (close to the density of water). Therefore: \[ M \propto V \] If we model organisms as having a characteristic body length \(L\), then: \[ V \sim L^3 \] So mass scales as: \[ M \sim L^3 \] This implies: \[ L \sim M^{1/3} \] --- #### Rewrite Surface Area in Terms of Mass Surface area scales with the square of length: \[ A \sim L^2 \] Substitute the relation \(L \sim M^{1/3}\): \[ A \sim (M^{1/3})^2 \] \[ A \sim M^{2/3} \] Muscles produce force by contracting fibers. Imagine a muscle shaped like a cylinder. The force it can produce depends on its cross-sectional area, not its length. But force per area is nearly constant across animals, because all muscles use the same molecular machinery. Galileo noticed that: - **Strength ∝ cross-sectional area** - **Weight ∝ volume** Mathematically: \[ \text{Strength} \propto L^2 \] \[ \text{Weight} \propto L^3 \] where \(L\) is the characteristic body length. Because weight increases faster than strength as animals get larger, **larger animals are relatively weaker** (in terms of strength relative to their body weight). This is why large animals must compensate structurally — for example: - **Elephants have very thick legs** - **Large animals have more column-like limb structures** - **Small animals can appear much stronger relative to their size** One beautiful consequence is that the paper predicts a common locomotor clock of about 0.1 s per body length at max speed. The authors later extended this idea: their 2021 paper on response times argued that across organisms from bacteria to large vertebrates, most simple stimulus-response times cluster around 25 ms, only a few times smaller than this locomotor timescale. That is a lovely hint that biology may operate with a small set of broadly shared dynamical timescales. A beautiful multi-scale connection: The scaling depends ultimately on $k_BT$ via the molecular motor energy, so the speed of animals is ultimately limited by thermal physics at the nanometer scale! ## TL;DR Because density, muscle stress, and metabolic power per mass are nearly universal across life, organisms (from bacteria to mammals) naturally end up with a maximum speed of roughly 10 body lengths per second. \[ \frac{V_{\max}}{L} \approx 10 \ \text{s}^{-1} \] Meaning: most organisms can move at about **10 body lengths per second**, despite **~20 orders of magnitude variation in mass**. The authors argue that this universal scaling comes from three biological quantities that are roughly constant across life: 1. **Body density** \[ \rho \approx 10^3 \text{ kg/m}^3 \] ≈ density of water. 2. **Muscle stress (force per cross-section)** \[ \sigma \approx 2\times10^5 \text{ N/m}^2 \] Set by molecular motors like myosin. 3. **Maximum metabolic power per mass** \[ b_M \approx 2\times10^3 \text{ W/kg} \] Using dimensional reasoning and energy limits on locomotion cycles, they derive: \[ \frac{V_{\max}}{L} \sim \frac{b_M \rho}{\sigma} \] Plugging in numbers gives: \[ \frac{V_{\max}}{L} \approx 10 \text{ s}^{-1} \] which matches observations. #### Why Large Animals Break the Law For large animals another constraint appears: **inertia of limbs**. #### Torque applied \[ C = \sigma S L \] #### Moment of inertia \[ I = M L^2 \] Since \[ M = \rho S L \] then \[ I = \rho S L^3 \] ### Angular acceleration \[ \alpha = \frac{C}{I} \] Substituting: \[ \alpha = \frac{\sigma}{\rho L^2} \] Thus **limb angular acceleration decreases with body size**. #### Consequence This leads to a **maximum absolute running speed** \[ V_{max} \sim \sqrt{\frac{\sigma}{\rho}} \] which becomes **independent of body size**. #### This explains why animals like - cheetahs - horses - ostriches all reach similar **top speeds around 20–30 m/s** despite large differences in body mass.