Ecology, Economy and Society–the INSEE Journal 6(1): 31-51, January 2023

RESEARCH PAPER

Exergy Analysis: A Guide to Sustainability?

Tejasvi Chauhan and Vinod K Gaur

Abstract: This paper argues for a continuing exploration of Nature’s organizing

principles that sustain prolonged homeostasis of the earth’s ecosystems punctuated
by forceful transitions to new emergent states. Ecosystems develop and maintain a
dynamically stable state by transacting energy and materials with the surrounding
flows to keep reversing their continual fall to the ground state. Conversely, the
elevation of any component of the ecosystem above the ground level may be
regarded as a measure of its functional efficiency. This measure, called exergy, can
be calculated for an eco-subsystem based on knowledge of the energy and material
fluxes that thread it and, most importantly, of where the ground level happens to
be. Admittedly, it is not straightforward to quantify these figures, and the departure
of assumptions from reality will inevitably translate into errors in the calculated
exergy figures. However, the variance may be estimated by analysing the results of
an ensemble of calculations with randomly perturbed input values. Even with these
limitations, however, a map of exergy losses characterizing different parts of an
ecosystem has the potential to reveal relative thermodynamic efficiencies for
appropriate ameliorative interventions.

Keywords: Exergy, Ecosystem, Thermodynamic limits

1. INTRODUCTION
Sustainability is not a value-neutral concern, begging the question as to what
is it aimed at. But at its heart lies the ambition to prolong the ambient state
of the earth’s ecosystems, whose future trajectories are inalienably
conditioned by the current state. Insightful hints to design a path towards
sustainability, irrespective of the focus of its concern, can perhaps be
gleaned by looking more incisively at the way ecosystems work.

Indian Institute of Technology, Bombay, India. [email protected]

CSIR - Fourth Paradigm Institute, Bangalore, India. [email protected]
Copyright © Chauhan and Gaur 2023. Released under Creative Commons Attribution ©
NonCommercial 4.0 International licence (CC BY-NC 4.0) by the author.
Published by Indian Society for Ecological Economics (INSEE), c/o Institute of Economic
Growth, University Enclave, North Campus, Delhi 110007.
ISSN: 2581–6152 (print); 2581–6101 (web).
DOI: https://doi.org/10.37773/ees.v6i1.914
 Ecology, Economy and Society–the INSEE Journal [32]

Planet earth’s ecosystems are unique in the solar system. They evolved from
rudimentary forms of self - replicating molecules, feeding off solar radiation
or energy from the deep earth, growing slowly at first and then more
deliberately by dispersal and structure formation. The first of these is a
piece of the universal process that inexorably enlarges the degrees of
freedom available to a system’s condition, a natural consequence of
unbiased treatment of all aspects of a situation. The second process, also
seen at work in inanimate systems fed by constant energy (such as
atmospheric convection) has been extensively studied through laboratory
experiments (for example, the Bénard convection). The results suggest that
structure formation in constantly powered systems (like ecosystems) greatly
enhances the rate of energy dissipation, perhaps to hasten the descent to
equilibrium. The significant fact is that they develop and sustain low-
entropy organized entities in the very act of falling along a gradient.
Every habitat on earth, including that of humans, belongs to some
ecosystem of the planet (Figure 1). Ecosystems are a society of living and
breathing beings much like our own, sustained by throughflows of matter,
energy, and information exchanged with the surroundings. They grow,
develop and stay alive by abstracting energy from other sources, such as
solar radiation and solar-driven cycles of wind and water, to constantly
restore the driving gradients of their arterial flows. Their potential energies
keep falling inexorably toward the dead state of equilibrium through the
very act of flowing. This is the spontaneous process forever at work in the
universe — a progressive flattening of all gradients. Highlands erode to
peneplains, gurgling mountain streams slow down to staid meanders,
concentrated materials disperse into environmental wastes, and energy
molecules oxidize to inert ones.
Unless injected constantly with fresh energy, these downhill processes
would eventually lay all systems to eternal rest (Figure 2). Even when a
source such as the daily inflow of solar radiation exists, and the landscape
can sustain thermal, topographic, and material flows (as seen in other
planets in the solar system) these do not result in a 'Goldilocks world’
where the conditions for life are just right. For that to happen, the solar-
driven flows need to be channelled through entities that can transform their
energy into useful work; entities that would keep reversing the downhill
processes by assembling inert matter into a chain of energy molecules, and
transforming wastes to clean the environment and mitigate hazards. This
functional machinery developed on earth as its ecosystems, made possible
by the planet’s special endowments: just the right mass to hold an
atmosphere in its gravitational cage and the right distance from the sun to
let liquid water exist. In turn, these conditions orchestrated the evolution of
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a blue planet, dynamically driven within by plate tectonics and on its surface
by the hydrological cycle. In time, the latter greened and flowered its land,
turning it into one of the most spectacular objects in space.
Figure 1: A Visual Representation of an Ecosystem

Source: Authors

Figure 1 presents an iconic view of an ecosystem. Its several distinctly

recognizable units are, in fact, mutually supported through invisible streams
of energy, materials, and information flows. These flows, primarily driven
by the thermal and electrochemical gradients created by solar radiation, are,
in turn, channelled through a vast network of energy, material, and
information transforming devices or ‘eco-engines’. The latter use the work
potential of their throughflows to maintain their live state and resist the
universal tendency towards levelling. In the process, they perform critical
ecosystem services, most notably, they help other eco-engines down the line
do the same.

2. ECOSYSTEMS AS A HIERARCHICAL NETWORK OF ECO-

ENGINES
Since energy can neither be created nor destroyed, but only transformed
through flows of heat, work, materials, or information, ecosystems are
essentially constituted as energy transforming devices called ‘engines’. In the
manner of all dynamical systems with continual energy inputs, which are
known to maximize the efficiency of energy transformation by developing
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an orderly structure (for example, Bénard convection cells, as examined in

Demirel, 2002), ecosystems too develop an optimal network of energy-
transforming flows. They exploit all possible configurations allowable by
their landforms and the daily and seasonally renewable stocks of energy and
materials, to maximize their flow density pathways, especially of the miracle
substance, water. The extraordinary properties of water as a universal
solvent and its large thermal inertia, further enhance an ecosystem’s
opportunity space for differentiation and co-development. The result is a
hierarchically ordered web of material and energy flows and their storages
that nourish and create the foundational structure of the food chain: soils,
forests, lakes, wetlands, groundwater, and biodiversity (Allen & Starr, n.d.;
Müller, 1992; O’Neill et al., 1989; Pahl-Wostl, 1993).
The invention of steam engines in the seventeenth century was inspired by
the experiential learning of early civilizations about transforming heat to
mechanical energy (for example, fire pistons used by Neolithic
communities) and creation of several ingenious devices that used steam
power to move shafts. In the nineteenth century, an analytical inquiry in the
efficiency of energy transformation processes led to an understanding of
energy quality as distinct from energy quantity. Analyzing the results of a
thought experiment, Sadi Carnot (Kelvin et al., 1924), showed that whilst
steam flowing from a high temperature source can be made to perform
work by moving a piston to drive a wheel, the amount of work (W)
performed by it was only a certain fraction () of the input steam energy
(Q). As the energy of flowing steam is progressively consumed in the
process of performing work, its thermal potential, that is, its temperature,
gradually reaches equilibrium with the surroundings. Thereupon, the
residual heat, unable to flow and be transformed any further, simply joins
the vast and unusable stock of environmental heat. The part of a system’s
total energy capable of being turned completely into work can thus be
measured by the elevation of its ambient state above the equilibrium level.
This is called ‘exergy’, and it is measured in the same unit as energy on a
scale defined by the zero potential energy of the equilibrium state or a state
of complete stasis.
Equivalently, the exergy of a particular system state is the energy required to
create it from the chemically inert, thoroughly dispersed, zero gradient
states of its components. For example, the exergy of one mole (a pack of a
given number of molecules) of biogas (methane) being 832 kJ, means that
a mole of the gas will produce 832 kJ of energy when allowed to unpack
itself. This is in fact, equal to the energy stored in every mole of methane
through the work performed by bio-organisms to i) extract carbon dioxide
(20 kJ) and water (2.6 kJ) from the environment, ii) bond the extracted
[35] Chauhan and Gaur

carbon and hydrogen molecules into one of methane (818 kJ), and iii)
return the residual oxygen (–7.8 kJ) to space. Unlike energy, exergy is not
conserved, and is, therefore, a more sensible qualifier of the work potential
of material and energy flows.
Figure 2: Flow of Energy within an Ecosystem

Source: Authors

Figure 2 illustrates the ability of systems to perform work through certain

processes by virtue of the fact that their elevated states are lifted above the
flat equilibrium state: raised highlands erode to renew soil fertility,
evaporated sea water is blown by wind to drive the hydrological cycle, inert
carbon dioxide molecules hydrogenated through photosynthesis form high-
energy glucose molecules, and the osmotic flow of water to plant roots is
driven by the concentration gradient. Left to their own devices, systems
possessing potential energies tend to move downhill to the zero gradient
equilibrium state, consuming the stored exergy that can be utilized to
perform useful work if channelled through an engine, such as the
production of electricity from falling water or metabolic energy from the
oxidation of carbohydrates in the guts of living beings. Ecosystems
integrate these processes into a biogeochemical cycle by abstracting the
exergy of inflowing solar energy and outflowing heat energy from the
earth’s depths to build gravitational and chemical potential energies in the
form of landscapes and biomass.
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Ecosystems maintain their health and build reserves for the future by
channelling the surrounding flows of materials, energy, and information
into a succession of lower-order streams thereby harvesting the maximum
of the available exergy (Silow & Mokry, 2010). Exergy extracted from the
daily and seasonally restored natural flow gradients that drive the
atmosphere and oceans, and proton flows in plant cells, is consumed to re-
build other less consumptive gradients of the ecosystem. Fertile flood plains
develop in the wake of diminishing stream power, as do wetlands, where an
incredible diversity of bio-organisms reduce environmental wastes to
organic carbon, thus preparing the base of the food chain. As some exergy
is irretrievably lost along each flow path as dead heat (because of the
ubiquitous presence of irreversibilities), the hierarchically organized chain of
eco-engines is driven by a progressively diminishing sequence of residual
exergies delivered by a preceding one (Figure 3). The entire exergy of solar
radiation extracted by ecosystems is, thus, used up by a succession of their
eco-engines to sustain the vital environmental flows and store some of it as
chemical exergy in the food and fuel molecules for use in the lean season
(Silow et al., 2011).

Analogously, one can visualize the same process at work when considering
a mountain stream that drives large rock masses down a steep valley and
deposits them on the flattening valley floor on entering the plains (Figure
3). This happens because the reduced gradient and approaching closeness
to equilibrium reduces the potential for energy exchange, reducing the
stream’s power even as the quantity of its flow remains the same. This
progressive reduction in stream’s power continues apace, depositing finer
and finer sediments into the biogeochemical marvel of a delta as it levels
with the sea. Thus, all of the earth’s energy requirements, including those of
its biosphere and human civilization, are met from the exergy or quality of
solar energy (Figure 3), not its quantity as in case of a hydroelectric plant
(which draws from the gravitational exergy of a flowing stream without
consuming a single drop of water).
Figure 3 shows how the exergy of solar energy is used by the two principal
thermodynamic engines of the earth system: the atmosphere-ocean and the
ecosystem, by harnessing the electromagnetic (thermal and electronic)
gradients created by the high exergy solar radiation. One is created by the
earth’s non-uniform heating, and the other by excitation of its otherwise
inert molecules (photosynthesis). The first converts solar exergy to move
the atmosphere & oceans, which in turn, drive the hydrological cycle and a
cascade of progressively smaller engines, notably the nutrient cycle and the
longer time-scale rock cycle (not shown). The hydrological & nutrient cycles
collaborate with the photosynthesizing engine to grow and develop the
[37] Chauhan and Gaur

living world, which continually reverses the various potential gradients to

maintain its low entropy islands against their ineluctable attrition. Like all
dynamic systems that evolve into an organized structure, the earth’s primary
engines too, harness the daily supply of solar radiation to structure the
earth’s climate and ecosystems. The figure (left middle) shows how the
exergy of solar radiation is used up in maintaining and developing the living
earth through a progressively diminishing cascade of energy and material
transforming Carnot engines, losing some exergy at every stage as heat
without using any of the energy received from the sun, which at the end of
the day, is radiated back to space with all its exergy reduced to zero.
Figure 3: Uses of the Exergy of Solar Energy

Source: Authors

At the top of the earth’s exergy flow pyramid are two major flow–field
gradients forged directly by the sun (Figure 3). The first one is the thermal
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gradient resulting from the non-uniform distribution of temperature on

earth because of its asymmetry with respect to the sun – earth axis and the
varying reflectivity of its variegated surface features. The inexorable
levelling of the thermal gradient mediated, in turn, by convective flows of
the earth’s fluid spheres keeps the planet at a moderate temperature and
creates humidity gradients, thereby driving a hydrological cycle. The latter,
interacting with solid earths near surface texture and topographic gradients,
creates surface flows of freshwater and its delayed release storages in
mountain glaciers, soils and in the ground below.
The second major gradient, collaboratively created with hydrological flows
and the entrained nutrients, is the electrochemical gradient created by the
absorption of solar photonic energy by electrons in the green molecules of
plants and algae (called chlorophyll). The exergy yielded by the flow of
protons down this gradient hydrogenates the carbon dioxide (CO2)
molecules plucked from the air, packing part of the flow energy in the
molecular bonds of the new glucose molecule (C6 H12 O6), which is the
basic building block of all life. An economically viable replication of the
process may indeed deliver us one day from the conundrum of global
warming.
The earth’s internal thermodynamic engine too builds topographic and
geochemical gradients through episodic upheavals, subsidence, and volcanic
effusions, but this involves much longer time scales than the diurnal and
seasonal flows of energy powered by the sun and the solar energy–driven
hydrological cycle. However, the former is responsible for sculpting the
longer-lasting surface features of the earth into smaller biogeophysical units
in which the annual and seasonal scale solar-powered environmental
processes operate.
Ecosystems flourish by flowing their share of fresh water and materials
along their landscape gradients (principally the gravitational and the
climatic) and generating a cascade of new ones using the exergy extracted
from the former. The latter gradients, which are primarily biogeochemical,
tend to organize themselves into a network of productive subsystems to
maximize the extraction of all available free energy for conversion into
biomass and ecological services, which is required by dependent
communities. The efficiency with which the seasonally replenished natural
resources of an ecosystem can be harnessed to sustain its well-being and
productivity is largely determined by the efficiency with which available free
energy in its various flow systems is harvested. Haphazard anthropogenic
interventions across the globe have disrupted many ecosystems with self-
organized flows. However, these may yet be rewired to maximize the
[39] Chauhan and Gaur

productivity of their eco-engines using creative engineering designs

informed by thermodynamic limits. What are these limits?

3. THERMODYNAMIC LIMITS
As explained previously, living systems (Figure A1) survive by being open,
that is, by exchanging materials and energy with the environment, and by
using up the latter’s exergies to maintain their well-being, and, thereby the
functioning of their subsystems. Healthy ecosystems unceasingly deliver
vital eco-services in the form of energy and material storages for use in
regular and lean seasons and ensure the availability of clean water and a
clean environment by transforming wastes. These processes, however,
proceed in strict accordance with the laws of thermodynamics. The first of
these laws states that energy is always conserved in any process. This is a
corollary of the fundamental principle of symmetry established by Emmy
Noether in 1915, a principle which helps explain the evolving universe.
Since energy transformation can only be mediated through the exchange of
heat, matter, information, or work, and the total energy must be conserved,
we can represent this fact symbolically as:
Qnet + Enet (matter) + Enet (Info.) − Wnet = (Einternal)syst. (1)
The RHS of (1) denotes net additions to a system’s energy stock accrued
from transformations of flowing streams of heat (Q), materials, and
information minus the work (W) delivered by the system. Work (W), in
particular, denotes the energy associated with the displacement of materials
mediated by force or pressure (such as moving a shaft). The statement
asserts that the net sum can only appear as a change in the system’s
molecular structure and its kinetic energy, collectively understood as its
internal energy (Einternal).
However, this statement gives no indication of the quality or potential of
the resulting exergy in terms of performing work. For example, the heat
energy of a stable atmosphere, which is known to be very large, cannot be
made to perform any work unless it is made to flow along a temperature
gradient, which is done by artificially creating a colder reservoir.
Quantifying energy quality is a concept that is explored in the second law of
thermodynamics, which was distilled from the analysis of a deeply insightful
thought experiment by Sadi Carnot in 1824. Assuming the flow to be
unhurried to allow reversing the process at any instant without any losses
(such as pushing a bicycle pump infinitely slowly), he proved that the
maximum useful work (W) extractable from a heat flux (QH) flowing from a
hot reservoir at temperature TH to a colder one at temperature T0 (to drive
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a motor, for example), is only a certain fraction () of the total inflow (QH).
The unhurried, reversible condition specified in his thought experiment was
used to ensure that no part of the input energy leaked out as waste heat (as
would otherwise happen, for example, when a bicycle pump is pushed
energetically). The efficiency () with which heat energy can be transformed
to work is equal to {1 – (T0 / TH)}, which is always less than 100%, even in
a slowly transforming reversible process, because, when flowing down a
thermal gradient with concomitant cooling, a part of it (= Q0) cannot
produce any work when it reaches equilibrium with the surrounding air at
T0. He further proved that this limitation was universally true irrespective of
the nature of the material flowing through the engine, thereby making the
result applicable to a wide range and variety of energy transformation
processes. Of equal significance is the fact that he proved that there was no
way by which this theoretical limit of efficiency could be breached, and
further, that in a reversible process of energy transformation from state A
to state B, the quantity, SB = (Q/T)B, remained constant irrespective of the
path of transformation. Claussius (1824) recognized this quantity as a
characteristic of the state of a system and called it ‘entropy’—a Greek word
related to transformation. As explained previously, entropy is inevitably
generated in any process that transforms heat to work when a part of the
input energy is reduced to impotence upon reaching equilibrium with its
surroundings. Indeed, entropy is generated in the transformation of even
those energies that possess 100% exergy, such as electric energy or
mechanical work, because of the various dissipative losses involved in all
physical operations. The ubiquitous presence of irreversibility in the real
world, such as friction-generated heat, thus makes the entropy of a
subsequent state of any physical system (SB) greater than the entropy of the
preceding state (SA). Biological systems and refrigerators transform energy
to create lower entropy islands locally at the expense of a larger quantum
being added to the environment. This one-way street, where a system
constantly evolves into a state of higher entropy, is thus an unexceptional
rule of energy transformation in the universe.
Or, dSA→B = (SB – SA) = (QB /TB – QA /TA ) = A →B d(Q/T) = A →B dS  0 (2)
The equality holds strictly true for a reversible process where Q = TS,
which, however, provides a reference for how far removed the efficiency of
a non-reversible physical system is from the ideal, and how imaginative
interventions may enable minimizing the gap. Given the thermodynamic
arbitrage imposed by the second law, we can restate the conservation law
for reversible processes as:
TS + Enet (matter) + Enet (Info.) − Wnet = (Einternal)syst. (3)
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Drawing logical corollaries from the above equation, we can show (vide
Appendix) that under reversible, ideal conditions, the rate of performing
useful work, or the power delivery of an open system exchanging heat,
matter, and information, is provided by (4) below.
Ẇcv = − d/dt(Ecv − T0Scv) + i=1→n Q̇i (1 − T0 /Ti) + ṁ (h + h* − T0s) (4)
Here, Wcv is the work potential of the system, or its exergy, expressed as the
net sum of various inflowing exergies (of its internal energy, heat, mass, and
information).
However, irreversibilities in real-world systems invariably destroy some of
the available exergy depending on the degree of their imperfections,
reducing the actual delivery to less than what was computed from equation
(3). The quantum of exergy destruction by an open system distilled from (3)
is given by (5):
(Exergy destruction rate)cv = T0{d/dt(Sgen)} (5)
The exergy destruction figures of an ecosystem or of its subsystems (which
can be calculated by applying (5) to the observed data) are an illuminating
indicator of their respective health. By mapping the exergy destruction
figures of sub-systems across an ecosystem, we can identify the ones whose
performance is short of the theoretical limit for reversible processes, and
thus warrant design interventions.

4. EXERGY FLOW THROUGH THE EARTH SYSTEM

The earth system receives a daily supply of highly concentrated radiation
energy, Qs  1.4 × 1022 joules, or 1.74 × 1017 watts (Kleidon 2012), shone
by a 6000* K hot sun. Since this radiation is non-uniformly distributed on
earth, a thermal gradient develops between the equator and the poles,
driving atmospheric and oceanic flows that keep the earth at an average
temperature of 300  27* K. Despite this prodigious supply of daily
radiation, the earth system, on average, is not heating or cooling
significantly during the day, as it radiates back virtually the entire Qs, after
abstracting its exergy  Qs = (1 – 300/6000) Qs = 0.95 × 1.74 × 1017 =
1.65 × 10 17 Watts, to fuel its works and the living world.
Additionally, the moderate thermal state of the earth, mediated by its
circulating fluid spheres and constantly hydrated by the wind-driven
hydrological cycle, promotes photosynthesis. The latter process can
produce biomass exergy equal to 7.0 × 1020 joules per day at 5% theoretical
efficiency (Zhu et al. 2008), if every ray of sunlight were to be captured by a
green plant. Actual biomass production on earth is, however, limited by
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non-ideal conditions to 70 giga-tonnes/yr. (Popp et al. 2014), providing a

daily budget of 3.5 x 1018 joules of chemical potential energy that fuels all
life forms on earth and most of their eco-service requirements.
The significant point to note here is that in any energy transformation
process, which is only possible through the exchange of heat, matter,
information, or work, mediated by a system (or engine), while energy itself
is conserved, the part available for performing work, that is, its exergy, even
under ideal conditions, is always less than the original: exergy   energy.
Thus, while the exergies of various forms of energy have some maximum
theoretical limits denoted by , the values actually realised are less than 
because of the ubiquitous presence of irreversible processes in far from
ideal, real-world systems. The latter are determined by the texture of
specific systems, such as friction-generated heat losses in an electric motor
despite the 100% theoretical exergy of electrical energy or pollutants-
induced biogeochemical debilitation in the performance of ecosystem
services. The deficit of an actual state, compared with the maximum
attainable energy (represented by ) of various system components, may
thus be regarded as a measure of exergy loss that could be minimized by
better design.
The numerical values of the available or designed exergies of eco-
subsystems, can, therefore, prove quite useful, not only in ranking their
relative contributions to an ecosystem’s well-being, but also in targeting
potentially ameliorative ones. Furthermore, exergy, being a system’s work
potential, is a dynamic quantity that increases when energy is stored in it
and decreases as the system moves downhill in the course of performing
some work. Its value at any given stage thus measures both its economic
worth and health. Also, as work potential, it acts as a universal metric (in
joules) for quantifying the state of any component of a system, whether
physical, chemical, or biological.

5. EXERGY OF ECOSYSTEMS
The exergy of ecosystems is primarily driven by their biomass, which
includes all life forms. It can be explicitly calculated by unpacking the
second term, Enet (matter), in (3) above. This can be shown (vide A6f of
Appendix) to be equal to the sum of the fractional concentration densities
xq of the endemic chemical and biological species present in a given
ecosystem, multiplied by their respective weighting factors, q, that is:
Execo = k=0→n ( k xk) (6)
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The  values for a fair number of commonly occurring species have been
calculated based on data from various ecosystems, which, in turn,
determines the efficiency of their exergy use. These values have been
updated accordingly in newer models (Jørgensen 2002; Jørgensen et al.
2005). Some interesting applications in the study of the exergy of
ecosystems can be found in Silow et al. (2011).
The wide range of ecological services that sustain us are rarely
acknowledged. What receives even lesser recognition, however, is the role
played by the immense diversity of life forms in sustaining the perennial
wheeling of energy and material cycles, which are essentially mediated by
the biochemical functions ordained by their embodied genetic information.
As explained above, the growth space available to ecosystems for
maximizing the harvestable exergy of solar radiation is dependent on the
number and variety of their eco-engines, that is, their biodiversity. The
latter also reinforces the resilience of ecosystems when it comes to
withstanding newly emerging stressors. Eco-engines do this by increasing
their bio-geographical spreads by creating new niches that suit a
proportionately larger pool of differentiated genetic traits. Indeed, some
studies on exergy analysis of ecosystems (Silow et al., 2011) confirm that the
movement away from thermodynamic equilibrium during ecosystem
growth and development coincides with higher levels of organization,
system configurations, and maximization of exergy use and storage of both
its chemical and information potential.
In essence, the pursuit of rational inquiry looks for an organizing principle
underlying masses of data. The intriguing work of living organisms in
relentlessly reversing downhill universal processes to maintain a dynamically
stable state has long engendered the conjecture that life processes are
governed by some overarching principles. Based on acute observations of
how the living world sustains itself, Lotka (1921−22), in his papers,
proposed that in the event of there being higher available exergy than
utilized, “an opportunity is furnished for suitably constituted organisms to
enlarge the total energy flux through the system”. This prescient conjecture
implies a self-directed orientation towards a higher range and level of
organization and complexity (information potential) in ecosystems that
would maximize the harvesting of available exergy, given the existence of a
suitably constituted genetic order. An interesting experiment designed by
Horowitz and England (2017) showed that a special configuration of atoms
(Lotka’s suitably constituted genetic order), will start tapping into those
energy sources, aligning and rearranging to better absorb energy and
dissipate it as heat. Prigogine and Stengers (1984) state that the growth of
organized structures, which characterize the growth of all living systems,
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can only be maintained by exporting a proportionately higher entropy

energy, produced by higher dissipation. This could, therefore, be a directive
principle leading the evolutionary trajectory of all dynamical systems.
Further, the implied principle, in order to be specific, must in some way be
unique, such as an extremal principle (Kelidon and Lorenz, 2005). Indeed,
several other indicators of ecosystem health have been proposed, notably
emergy (Odum 1991), total system throughflow (Patten 1995), and ascendency
(Robert E Ulanowicz 2000). However, as Fath et al. (2001) argue, they are
either implied by each other or are complementary. Thus, persuasive as
these ideas are, they remain to be tested, even though a substantial amount
of data may now be available in traditional archives of ecological research.
Exergy analysis of eco-subsystems (by providing a direct measure of their
efficiency) lights a path to maximize exergy storage for a given set of inputs,
thereby also maximize the total system throughflow (considered by some as
an independent goal function). In our view, therefore, exergy analysis by
itself or, wherever possible, complementarily with other indicators, offers
obvious advantages in designing ecosystems and subsystems at various
scales to serve urgent contemporary objectives such as meeting the decadal
goals of ecosystem rehabilitation. It also holds high promise of leading to a
robust theory of ecosystems, further enriching the methodology and
approach to ecosystem design that would streamline its human dimension.

APPENDIX
Exergy Analysis of Ecosystems
Ecosystems constitute a web of energy-transforming sub-systems (Figure 3)
that constantly exchange energy, materials, and information with their
surroundings to maintain their evolving dynamic states above the
thermodynamically dead state of equilibrium. Their ability to keep
producing ecosystem goods and services is accordingly measured by their
dynamically maintained distance (potential) from the downhill flat
equilibrium state. This equals the total useful work or exergy that a system
will yield if it were allowed to drift towards the equilibrium state of zero
exergy when it ceases to exchange any energy or materials with its
surroundings (Figure 2). Similarly, this potential is equal to the energy
required to create this system out of the zero gradients everywhere of all
physical and chemical potentials.
[45] Chauhan and Gaur

Figure A1: A Schematic of an Open System

Source: Authors

Figure A1 provides a schematic of an open system—an energy and

material-transforming engine— lying in the field of flowing streams of heat,
information, and work from which it abstracts exergies depending on its
specific structure. The application of the principles of thermodynamics to
the ambient dynamical parameters of the open system enables us to
calculate the gap between their actual functioning and a possible one,
thereby providing a diagnostic tool to identify their state of viability and
design engineering interventions to improve their condition.
To calculate the exergy produced by such a system, we visualize a
representative element of it, called the control volume (Figure A1), with
flowing streams of heat, materials, information, and work. Let Qin (t), min
(t), Iin (t), and Win (t) denote the heat, mass, information, and work,
respectively, contained in an imaginary wafer of thickness dx, ready to be
pushed in such a control volume at the time instant t, by pressure pin. A
similar wafer containing the respective quantities of Qout, mout, Iout, and Wout
leaves the control volume, t, seconds later at the time (t + t), pushed out
by pressure pout. Unlike the heat and work streams, however, the material
stream carries at least five forms of energy: i) internal energy, ii) the work
performed by the difference of pressures moving the wafers through the
system, iii) kinetic energy, iv) gravitational potential, v) and chemical
potential released by chemical reactions such as the digestion of food.
Material streams also contain much of the information flowing through the
system in the form of biodiversity (gene codes) and learned knowledge that
guides the various organisms to seek food and mates and protect
 Ecology, Economy and Society–the INSEE Journal [46]

themselves from predators. Thus, the second and third terms in (1) can be
written as:
Enet (matter) + Enet (Info.) = [j min{(u + pvsp.) + (cin2 /2 + gz + q0j + I)}in − k
mout {(u + pvsp.) +(cout2 /2 + gz + q0k + I)}out] = 1→j m(h + h*)in – 1→k m(h
+h*)out, (A1a)
Here, the summation over j and k denotes the number of inlet and outlet
portals an open system may have, c is the velocity term in kinetic energy, h
is the specific enthalpy per unit mass of internal energy u, pvsp is the work
performed by the inlet and outlet pressures, and h* denotes the sum of the
kinetic and potential energies of gravitation, information, and chemical
bonds of the qth chemical species.
Accordingly, it is instructive to rewrite the conservation equation (1) more
explicitly by including all forms of energy:
Qnet + Enet (material) + Enet (Info.) − Wnet = (Einternal)cv. = [1→n Qnet + 1→j
m(h+h*)in − 1→k m(h+h*)out ] – Q0 – Wcv , (A1a)
Or, Wcv = –Ecv + 1→n Qnet + 1→j m(h+h*)in − 1→k m(h+h*)out – Q0
(A1b)
Here, Wcv is the output work of the system and Q0 is the heat dumped in
the surroundings, adding entropy equal to Srev. = Q0/ T0 at its equilibrium
temperature, T0. T0 remains largely unaffected by the various exchanges
with the system because of its substantially larger thermal reservoir. We use
this fact to temper the conservation law, otherwise devoid of any sense of
energy quality, with the limits imposed by the second law.
According to the second law, the change in entropy Scv within the control
volume over a time interval t is equal to the net sum of entropies
associated with the inflowing and outflowing streams of heat and materials
[{(SQ)in –(SQ)out +  (ms)in − (ms)out], regarded as ideal reversible
processes, in addition to the entropy Sgen generated within the system due to
all departures from the assumed reversibility. Accordingly,
Scv = [{Sgen – (Q0/T0)} + {1→n{(Qi/Ti) + 1→j (ms)in − 1→k (ms)out}] (A2a),
where capital S refers to the residual entropies of the system as well as those
generated within, and its lower case to specific entropy (per unit mass).
Or, Q0 = T0[–Scv + Sgen + [1→n {(Qi/Ti) + (ms)in – (ms)out] (A2b)
Substituting the above value of Q0 into (A1), one obtains the work output
of the system as:
[47] Chauhan and Gaur

Wcv = −(Ecv − T0Scv +T0Sgen) + 1→n Qnet (1 − T0 /Ti) + 1→j m (h + h* − T0s) –

1→k m (h + h* − T0s) (A3a)
The first term on the RHS is the sum of i) exergy lost from the internal
energy of the system due to various irreversibilities of energy
transformation over the time interval t, while the second and third terms
are the exergies gained by the system from the streams of heat and materials
through a reversible transformation. Dividing the various difference terms
in (A3a) by t, the time interval over which the change in the flow regime
through the system is defined by the entry of an infinitesimally thin wafer of
their contents and the exit of another, one can express the work output of
the system by an equivalent power output as:
Ẇcv = − d/dt(Ecv − T0Scv +T0Sgen) +  Q̇i (1 − T0 /Ti) + ṁ (h + h* − T0s) (A3b)
Since exergy is the work potential of a system assuming reversibility of
transformations, it is obtained by deleting the T0Sgen term from (A3b), being
the only one caused by irreversibilities. Thus:
(Exergy Power)cv = −d/dt(Ecv − T0Scv)+ Q̇i (1 − T0 /Ti) + ṁ (h + h* − T0s)
(A3c)
The difference between (A3b) and (A3c) yields the amount (Ex)Dest of
exergy lost or destroyed by a given transformer.
(Ex)Dest rate = d/dt(T0Sgen) (A4)
Since (Ex)Dest should ideally be zero for a perfect system or only marginally
positive for a healthy one, its value above zero indicates the health of an
ecosystem. Mapping the numerical values of the exergy destruction of eco-
subsystems, which can be calculated using (A4), provides a powerful tool
for auditing the functioning of extant ecosystems as well as for testing the
efficacy of those ecosystems designed for engineering interventions.
As an example, consider the case of a flowing stream in a steady state, that
is, d/dt(Ecv) = 0, and all heat exchanges taking place at the environmental
temperature T0. With these conditions the exergy/mass of a steadily flowing
stream is given by:
(Specific Exergy)steady flowing stream = (h + h* − T0s)in − (h + h* − T0s)out (A5)
Exergy of Ecosystems
The work potential of biomass resides in i) energy stored in the chemical
bonds that hold the molecules of a substance together, ii) information
contained in the genetic code of bio-organisms that ordain their
functioning, and iii) the network of biological society that catalyses their
 Ecology, Economy and Society–the INSEE Journal [48]

collective synergy. Thermodynamic processes involving material

transformations either perform work to raise the exergy of lower exergy
inert molecules or extract the energy stored in their chemical bonds by
breaking them. In either case, the energy transaction alters the chemical
potential of the molecular assemblage, which is accordingly defined as the
energy required to add or remove an atom or molecule from a given mass.
The chemical potential q of a particular species q in a mixture is
accordingly defined as the partial derivative of the energy of the substance
with respect to the number of that species, all other species remaining
constant q = E/nq. The chemical exergy of an ecosystem is hidden in
the last term of (A3c).
Therefore, when rewriting this expression in a differential form for a unit
molar mass and dropping the suffixes, we get:
dE = d(TS – PV) +  q = 1→n dq (A6a)
Or, (dE – TdS – SdT – Pdv + VdP) =  dq,
which, for isothermal and constant volume processes largely true for
ecological systems, may be reduced to:
VdP = dq = (RT/P)dP (A6b)
Integrating (A6b), one obtains:
q = RT[ log (Pq/Pq0) q0] = RT  q = 0→n xq [log (xq /xq0) – q0 ] (A6c)
where Pq is the partial pressure associated with the flow of the qth chemical
species = xqP0, xq being the molar fractional concentration of the qth species
such that xq = 1, and xq0, its value in the zero gradient equilibrium state.
The summation includes species of inorganic (q = 0) and organic molecules
(q = 1) and bio-organisms (q  2). The latter, viewed as information exergy
engines, are driven by both coded information in the genes of different
communities and network information.
The biochemical exergy of an ecosystem, which is defined as the work
potential above the equilibrium state xq q0, is therefore given by:
Execosystem = RT  q = 0→n xq log(xq /xq0) (A6d)
Values of xq for the inorganic as well as organic molecules in the ecosystem
can be determined by measuring their fractional values in the laboratory. To
obtain these with respect to bio-organisms, we must interpret them from
the perspective of the probability of their occurrence in the environment: xq
= pq X, where
X =  q = 1→n xq, and X0 =  q = 1→n xq0 (A6e)
[49] Chauhan and Gaur

where X0 is the total organic matter per mole, at the equilibrium state,
which, for an evolving ecosystem, represents a previous state. Assuming,
however, that there is no substantial transfer of matter during the period,
we see:
Execosystem = XRT  q = 0→n pq log(pq /pq0) =  q = 0→ n qxq, (A6f)
Jørgensen (2002) indicates how the quantity RT log(pq /pq0) = q, may be
calculated by measuring the concentration density of the respective species
in the environment. This calculation is based on available knowledge of the
genetic structure of endemic communities, and the model assumed for
transforming information into eco-products. New research on the structure
of information-controlling genes, for example, was used by Jørgensen et al.
(2005) to revise Jorgensen’s earlier determinations of q values (Jørgensen
2002). Researchers continue to update these values for better evaluation of
ecosystem exergy as new understanding is gained about the microstructure
of the living world.

ACKNOWLEDGEMENTS
The authors wish to thank Sophie Gaur for designing the figures. TC
acknowledges the support of the Prime Minister’s Research Fellowship
(PMRF), the Ministry of Education, and the Government of India—
RSPMRF0262.

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