Académique Documents
Professionnel Documents
Culture Documents
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/
info/about/policies/terms.jsp
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content
in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship.
For more information about JSTOR, please contact support@jstor.org.
The Royal Society is collaborating with JSTOR to digitize, preserve and extend access to Philosophical Transactions: Biological
Sciences.
http://www.jstor.org
This content downloaded from 146.155.94.33 on Thu, 16 Jul 2015 15:06:21 UTC
All use subject to JSTOR Terms and Conditions
PHILOSOPHICAL
TRANSACTIONS
-OF
THE ROYAL
SOCIETY
Introduction
in
entropy production
and ecological
systems
Maximum
environmental
Axel
Yadvinder
Kleidon1'*,
Malhi2
and Peter M.
Cox3
The
interactions; Earth
thermodynamics;
Keywords:
Lovelock
1944;
in 1886 that:
1965).
Boltzmann
... the
of animate
general
beings
struggle for existence
a struggle
for organ
for raw materials?these,
is not
are
isms,
air, water
and
soil,
all abundantly
available,
available
the hot
sun
through
to the cold
the
transition
of
energy
Earth.
for correspondence
contribution
of
in ecological
production
and implications'.
17
(akleidon@bgc-jena.mpg.de).
to a Theme
and
Issue
environmental
'Maximum
systems:
entropy
applications
system science;
ecosystems
Taken
suggest that in
together, these examples
order to better understand
Earth's
environmental
and ecological systems and their couplings, we need
to view these as coupled thermodynamic systems that
are organized in a state far from thermodynamic equi
librium. Central to thermodynamics is the concept of
'entropy'
as
a measure
of
'disorder'
or
'randomness'.
While
1298
A. Kleidon
et al.
Introduction.Maximum
entropyproduction
power
systems (Dewar
2003)?maximum
1988) or exergy (Jorgensen &
(Lotka 1922a,?; Odum
Svirezhev 2004), depletion of gradients (Schneider &
a proposed
constructal
law
and
2005)
Sagan
of these hypotheses
(Bejan & Lorente 2006). Many
have in common that at least in part they have been
the
motivated
specifically
by
thermodynamics,
these
law of thermodynamics.
second
However,
at a highly
hypotheses have mostly been proposed
Seleznev
&
Whitfield
2006;
2005; Martyushev
Kleidon
2009) and indications for its general validity
of heat transport
in simple energy balance models
as well as complex climate
et al. 2001)
(Lorenz
models
(Kleidon et al 2003, 2006).
This special issue resulted from a workshop on the
in the Earth
topic of 'Maximum Entropy Production
at
in
2008
the
Max-Planck-Institute
held
May
system',
in Jena, Germany. The contri
for Biogeochemistry
butions aim to shed light on how Earth systems are
and organized far from thermodynamic
on how these
equilibrium, with special emphasis
system states can be described by using the proposed
maintained
MEP
principle.
Before we provide an overview of the contributions,
we will briefly elaborate on the nature of the MEP
principle, and the ambiguities that surround its appli
cations and interpretation. We first note that at a
fundamental
no
cesses,
ecological,
energy,
mass,
momentum
and
other
properties.
Trans. R. Soc.
sufficient degrees
of freedom, that is, sufficient
choice among steady states that all meet the funda
mental
conservation
be
laws, the system would
of entropy pro
characterized
by a maximization
state would
duction. Roughly
speaking, this MEP
to a state at which maximum
correspond
physical
power is generated, gradients are dissipated at the fast
rate and hence the associated
est possible
entropy
production ismaximized.
This sounds relatively straightforward, but in prac
tice many questions come up. Before we can test the
how do we describe environ
application of MEP,
mental
and
in
processes
purely
ecological
are
con
terms?
Which
the
relevant
thermodynamic
are the degrees of freedom? How do
straints? What
these degrees of freedom develop and how are they
maintained? Which
flux should be optimized,
and
not
in
this
flux
and
another?
The
contributions
why
this special
issue aim to resolve some of these
questions.
ofMEP
(2010)
This content downloaded from 146.155.94.33 on Thu, 16 Jul 2015 15:06:21 UTC
All use subject to JSTOR Terms and Conditions
Introduction.Maximum
fromMEP
shows high variability in its instantaneous
fluxes and rates, as found for instance in turbulent
fluid flow, heat convection and ecological
systems.
(2010) provides a brief account on the
Martyushev
reasonability ofMEP within the context of non-equili
brium
(2010)
Bejan & Lorente
thermodynamics.
an
of
overview
the
so-called constructal law,
provide
as a possibly complementary approach to understand
how optimized structures develop, such as the organiz
in living
ation of river systems or blood vessels
the atmosphere.
The following two papers deal with hydrological
processes on land. Zehe et al. (2010) evaluate the
effect of preferential flow associated with biogenic
soil structures on hydrological
fluxes using non
equilibrium thermodynamics. They show that these
structures act to maximize
of chemical
dissipation
soil.
within
the
Paik
& Kumar
potential gradients
a
review
of
(2010) provide
optimality approaches
to characterize
that have been proposed
fluvial
patterns in hydrology and geomorphology. Several of
these are related to energetic properties,
such as
minimum stream power, minimum energy dissipation,
maximum
friction and minimum energy expenditure.
In order to evaluate the validity of these apparently
Phil.
Trans. R. Soc.
entropyproduction
A. Kleidon
et al.
1299
in entropy production.
outperform abiotic processes
Vallino argues that ecosystems should be viewed as
that function to
self-organizing molecular machines
maximize
long-term entropy production at the levels
of ecosystem. Dewar
(2010) reviews different optimiz
ation theories in plant physiology and shows how MEP
can unify these different theories. By setting the system
boundary at different scales (leaf, plant and ecosys
associated
with
the
tem), he shows that MEP
the
of
chemical
created
energy
dissipation
by photo
of different
synthesis can result in the maximization
plant fluxes, such as net primary productivity, net
growth rate and canopy photosynthesis, respectively.
mation
in vegetation in semiarid regions. They show
that the patterned state can be adequately represented
if the vegetated fraction is
by a simple two-box model
that MEP
optimized to yield MEP.
They conclude
to better aggregate
could be used
subgrid scale
heterogeneity in numerical simulation models of the
land-atmosphere
system.
(2010)
This content downloaded from 146.155.94.33 on Thu, 16 Jul 2015 15:06:21 UTC
All use subject to JSTOR Terms and Conditions
1300
A. Kleidon
et al.
Introduction.Maximum
entropyproduction
time
IO1
resistance,
IO2
IO3
IO4
/?, (% /?,)
boundary
(c)
1)
of entropy production:
that can lead to confusion
1,
regarding maximization
(a) confusion
examples
versus
states of a
behaviour
the selection
transient
a difference
in focus on understanding
among
steady
or even
of one flux that can simultaneously
result in a decrease
from the maximization
2, originating
system;
(b) confusion
of another flux, here illustrated with a simple electric circuit diagram with a fixed resistance
minimization
Ri and the sensitivity
1. Three
Figure
originating
of the resistances
of dissipation
boundary
different
from
(see
3. A NOTE OF CAUTION
Ri
and R2
to the value
of R2
Trans. R. Soc.
(c) confusion
3, originating
from
the drawing
of the system's
explanations).
and
(2010)
This content downloaded from 146.155.94.33 on Thu, 16 Jul 2015 15:06:21 UTC
All use subject to JSTOR Terms and Conditions
Introduction.Maximum
MEP
Rim
the theoretical
the conditions
principles are
applicable.
IfMEP
is indeed valid and its range of applicability
is understood
inmore detail, itwould be of tremen
us
It would
to formulate
dous
allow
help.
fundamental evolutionary directions for both environ
mental as well as ecological systems. Given that natural
processes operate and interact at vastly different time
scales (e.g. weather versus plate tectonics, leaf gas
exchange versus speciation), the tendency of slow pro
cesses to evolve to anMEP
state should manifest itself
in an evolutionary
trend towards higher rates of
at
shorter time scales. At a practi
entropy production
cal level, the MEP
principle could pave the way for
better and more theoretically grounded parameteriza
tions
in
Earth
of different
system models
a
MEP
would
critical
role in
complexities. Here,
play
and
variabil
scaling spatial heterogeneities
temporal
ities to the inevitable coarser grid and time steps that
are unavoidable
in numerical
of
representations
environmental and ecological systems.
Phil.
Trans. R. Soc.
and planetary
complex biological
A.K.
the Helmholtz
thanks
systems.
as his
Gemeinschaft
research
alliance
and
Evolution
'Planetary
has
through the
Life'.
is
Y.M.
REFERENCES
A.
Bejan,
ation
& Lorente,
of
2006
S.
Constructal
theory of gener
and
engineering.
nature
in
configuration
design
L.
Boltzmann,
ischen
in nature.
evolution
365, 1335-1347.
Phil.
Trans.
R.
Soc.
(doi:10.1098/rstb.2009.0302)
1886
Der
zweite
der mechan
Hauptsatz
Almanach
der
kaiserlichen
W?rmetheorie.
maximum
theorem,
self-organized
C.
R.
Dewar,
and
entropy production,
in non-equilibrium
stationary
criticality
states. J. Phys. A
4470/36/3/303)
2005a
36, 631-641.
Maximum
statistical
non-equilibrium
(doi: 10.1088/0305
entropy
mechanics.
and
production
In Non-equilibrium
Germany:
Springer
Verlag.
plant
1301
Dewar,
et al.
A. Kleidon
entropyproduction
C.
2010
Maximum
1429-1435.
365,
(doi:10.1098/rstb.2009.0293)
R.
Holdaway,
and
entropy
production
Phil.
Trans. R. Soc.
theories.
optimization
A.
J., Sparrow,
D.
&
A.
D.
Coomes,
2010
ment
in the Amazon
1437-1447.
Jorgensen,
Basin.
Phil.
Trans.
R.
Soc.
365,
(doi: 10.1098/rstb.2009.0298)
S. E.
&
Svirezhev,
Y. M.
2004
Towards
thermo
Elsevier.
from
a two-box
dynamics.
Phil.
2010 MEP
climate
Trans.
R.
model
Soc.
atmos
containing
1355-1365.
365,
(doi:10.1098/rstb.2009.0297)
A.
2009 Non-equilibrium
Kleidon,
thermodynamics
and maximum entropy production in the Earth system:
applications
and
653-677.
Kleidon, A.
implications.
96,
Naturwissenschaften
maximum
entropy
production.
Phil.
Trans.
R.
Soc.
circulation
and
states
of maximum
entropy
entropy
layer exchange
(2010)
This content downloaded from 146.155.94.33 on Thu, 16 Jul 2015 15:06:21 UTC
All use subject to JSTOR Terms and Conditions
1302
et al.
A. Kleidon
Introduction.Maximum
model.
Lorenz,
R.
D.
2010
The
(doi:10.1029/
R. SOC.B365,
Lorenz,
R. D.3
entropy
heat
& McKay,
P. G.
P.
C.
Trans.
(doi:10.1098/rstb.2009.0312)
J. L, Withers,
Lunine,
Phil.
studies.
production
1349-1354.
transport.
Lett.
Res.
Geophys.
28,
Proc.
ution.
Sci.
Acad.
Nati
USA
8,
147-151.
10.
(doi:
Lovelock,
J. E.
experiments.
207,
568-570.
(doi:10.1038/
two basic
1333-1334.
maximum
questions.
Phil.
entropy
Trans.
R.
production
Soc.
365,
(doi:10.1098/rstb.2009.0295)
L.
Martyushev,
The
2010
&
M.
Seleznev,
V.
D.
2006
(doi:10.1098/rstb.2009.0300)
R.
potential
K.
for non-equilibrium
of
flow
free-energy-like
systems at steady state.
365, 1323-1331.
(doi:10.1098/rstb.
2009.0296)
. T.
Odum,
1988
Self-organization,
transformity,
and
1132-1139.
(doi: 10.1126/
R. D.
T.
science.242.4882.1132)
Ozawa,
H.,
Ohmura,
A.,
Lorenz,
& Pujol,
2003
The
system?a
review
principle. Rev.
2002RG000113)
Phil
Trans. R. Soc.
of
the maximum
Geophys.
41,
entropy
1018.
production
(doi:10.1029/
&
D.
Sagan,
2005
Into
the cool:
of irrevers
energy flow,
IL: University
life. Chicago,
in semiarid
of heterogeneity
representation
of
ecosystems.
365, 1449-1455.
(doi: 10.1098/
rstb.2009.0309)
Swenson, R. 1997 Autocatakinetics, evolution and the law of
maximum entropy production: a principled foundation
of human
the study
ecology.
Adv.
Ecol.
Hum.
6,
1-47.
Phil.
production.
Trans.
R.
Soc.
how
rather,
you
it. Phil.
produce
Trans.
R.
Soc.
Minimization
2010
(doi:10.1098/
Press.
maximum
F. J. R. & Bruers,
S. 2010 Ecosystem
functioning
Meysman,
a quantitative
test of
and maximum
entropy production:
1405-1416.
Soc.
Trans.
R.
Phil.
365,
hypotheses.
landscapes.
Maximum
365, 1387-1395.
thermodynamicsand
towards
L. M.
principle:
Niven,
E. D.
Schneider,
207568a0)
Martyushev,
on
patterns
to thermodynamics
I. 1955 Introduction
Prigogine,
IL: Thournes.
ible processes. Chicago,
Chicago
1073/pnas.8.6.147)
fluvial
rstb.2009.0303)
415-418.
(doi: 10.1029/2000GL012336)
Lotka, A. J. 1922a Contribution to the energetics of evol
to
characteristic
limit
maximum
. & Kumar,
Paik,
describe
of climate:
model
two-box
entropyproduction
W.
&
Y.
Liu,
2010
new
one-dimensional
radiative
entropy
Phil.
Trans.
R.
Soc.
(doi:10.1098/rstb.2009.0301)
Zehe, E., Blume, T. & Bl?schl, G.
'maximum
perspective
tures. Phil.
Trans.
energy dissipation':
thermodynamic
on rapid water
flow in connected
soil struc
1377-1386.
Trans. R. Soc.
365,
(doi: 10.
Soc.
Chemotaxis
R.
1367-1376.
a novel
1098/rstb.2009.0308)
Zupanovic, P., Brumen, M.,
Bacterial
365,
365,
entropy
1397-1403.
2009.0307)
(2010)
This content downloaded from 146.155.94.33 on Thu, 16 Jul 2015 15:06:21 UTC
All use subject to JSTOR Terms and Conditions
production.
Phil.
(doi:10.1098/rstb.