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Atmospheric homeostasis by and for the biosphere:

the gaia hypothesis

By JAMES E. LOVELOCK, Bowerchalke, Nr. Salisbury, Wilts. England and


LYNN MARGULIS, Department of Biology, Boston University, 2 , Cummington Street,
Boston, Mass. U S A

(Manuscript received May 8; revised version August 20, 1973)

ABSTRACT

During the time, 3.2 x 109 years, that life has been present on Earth, the physical and
chemical conditions of most of the planetary surface have never varied from those
most favourable for life. The geological record reads that liquid water was always
present and that the pH was never far from neutral. During this same period, however,
the Earth’s radiation environment underwent large changes. As the sun moved along
the course set by the main sequence of stars its output will have increased at least
30 % and possibly 100 %. It may also have fluctuated in brightness over periods of a
few million years. At the same time hydrogen was escaping t o space from the Earth
and so causing progressive changes in the chemical environment. This in turn through
atmospheric compositional changes could have affected the Earth’s radiation balance.
It may have been that these physical and chemical changes always by blind chance
followed the path whose bounds are the conditions favouring the continued existence
of life. This paper offers an alternative explanation that, early after life began it
acquired control of the planetary environment and that this homeostasis by and for
the biosphere has persisted ever since. Historic and contemporary evidence and argu-
ments for this hypothesis will be presented.

It is widely believed that the abundance of organic equilibria are seen as more apparent
the principal gases N2 and 0, is determined by than real.
equilibrium chemistry. One of the larger pro- A starting point is a consideration of the
blems in the atmospheric sciences is t h a t of profoundly anomalous composition of the
reconciling this belief with the uncomfortable Earth’s atmosphere when it is compared with
fact that these same gases are cycled by the that of the expected atmosphere of a planet
Biosphere with a geometric mean residence time interpolated between Mars and Venus. Thus on
measured in thousands of years. The more Earth the simultaneous presence of 0, and CH,
thoroughly the inventory of a n individual gas is at the present concentrations is a violation of
audited the more certain it seems t h a t inorganic the rules of equilibrium chemistry of no less
equilibrium or steady state processes determine than 30 orders of magnitude. Indeed so great
its atmospheric concentration but the same is the disequilibrium among the gases of the
audit frequently further reveals the extent of Earth‘s atmosphere t h a t i t tends towards a
its biological involvement. A lucid account of combustible mixture, whereas the gases of Mars
contemporary information on the problem of and Venus are close t o chemical equilibrium and
the cycle of gases is in the paper of Junge are more like combustion products.
(1972). The anomalous nature of the atmosphere has
This paper presents a new view of the atmos- been known since Lewis & Randall (1923) first
phere, one in which it is seen as a component commented t h a t at the pE and p H of the Earth
part of the biosphere rather than as a mere the stable compound of nitrogen is the NOB ion
environment for life. I n this new context the in the oceans; gaseous nitrogen should not be
incompatibilities of biological cycles and in- present. I n spite of reminders b y Hutchinson

Tellus XXVI (1974),1-2


ATMOSPHERIC HOMEOSTASIS BY AND FOR THE BIOSPHERE 3

(1954) and Sill& (1966) this anomaly has covered. With the total ecosystem, Gaia, apart
remained unnoticed in the debate on atmosphe- from the shadowy evidence of those primitive
ric cycles. beliefs in her reality we are dependent upon
Given the chemical composition of a planetary physical rather than biological evidence. There
atmosphere it is possible to infer the presence is little doubt that living things are elaborate
or absence of life, Hitchcock & Lovelock (1967). contrivances. Life as a phenomenon might there-
To do this the entire ensemble of reactive gases fore be considered in the context of those app-
constituting the atmosphere needs be con- lied physical sciences which grew up to explain
sidered and when this is done information is inventions and contrivances, namely thermo-
made available which is otherwise inaccessible dynamics, cybernetics and information theory.
when each gas is considered separately in isola- The first cautious approach to a classification
tion. This approach applied to the present of life, reached general agreement as follows
problem of the anomaly of the chemical distribu- ‘Life is one member of the class of phenomena
tion of the gases of the atmosphere, offers a which are open or continuous reaction systems
strong suggestion that the Earth’s atmosphere able to decrease their entropy a t the expense of
is more than merely anomalous; it appears to be free energy taken from the environment and
a contrivance specifically constituted for a set of subsequently rejected in a degraded form’ (Ber-
purposes. nal, 1951; Wigner, 1961).
This paper examines the hypothesis that the This may also be expressed in the form of the
total ensemble of living organisms which consti- equation of continuity for entropy (Denbigh,
tute the biosphere can act as a single entity to 1951).
regulate chemical composition, surface p H and dS
possibly also climate. The notion of the bios- e -+divS=8
dt
phere as an active adaptive control system able
to maintain the Earth in homeostasis we are Where 8 is the rate of internal creation of
calling the ‘Gaia’ hypothesis, Lovelock (1972). entropy, e the density and S the entropy; div S
Hence forward the word Gaia will be used to is the outflow of entropy and e(dS/dt) the rate
describe the biosphere and all of those parts of of change of entropy in the enclosed region; 0
the Earth with which it actively interacts to must by second law be zero or positive; the
form the hypothetical new entity with proper- possibility that div S can be large and positive
ties that could not be predicted from the sum makes possible a negative trend for e(dS/dt).
of its parts. This classification is broad and includes also
phenomena such as vortices and flames and
many others. Life differs from such primitive
Theoretical basis processes of the abiological steady state in the
The fundamental problem underlying the for- singularity, persistence and size of the entropy
mal recognition of an unfamiliar living associa- reduction it sustains. Although limited, this
tion, such as Gaia, is that of recognising life phenomenological description of the class of
itself. We are so preprogrammed to recognise process, which includes life, is helpful in our
life instinctively that the logical basis of the search for proof of the existence of Gaia in two
recognition is rarely questioned. Thus the dis- ways. Firstly by serving to define the boundary
covery of a member of new and bizarre animal of the internal region where entropy is reduced
species raises the question, what is it, never the and secondly by suggesting that the recognition
question, is it life? This instinctive recognition of a living entity can be based upon the extent
passes beyond organisms to systems provided of its physical and chemical disequilibrium from
that they are not too big to be seen. Thus a the background environment.
bees nest is recognised as a purposeful structure, On the matter of boundaries, it is obvious that
quite different from a cluster of non-social a man, as an example of a living entity, takes in
insects. The intricate organisation of an eco- free energy in the form of the chemical potential
system such as a tropical rain forest, however, difference between food and oxygen and sus-
was not recognised until the evidence of the stains a low internal entropy through excretion
interdependence of its parts and the economy of waste chemicals and heat. To a man, the
of the cycling of essential elements was dis- environment to which entropy is discarded in-
Tellus XXVI (1974). 1-2
4 JAMES E. LOVELOCK ET AL.

eludes the atmosphere and his boundary is Table 1. The partial pressures’ in millibars of
therefore his skin. It might seem pointless there- CO,, N , and 0 , on Mars, Earth and Venus and
fore on Earth to seek the existence of a general on two model abiological Earth‘s
living system, Gaia, in terms of entropy reduc- Model ( D )after life had been deleted and Model (I)
tions within the atmosphere which clearly for a geometric mean interpolation between Mars and
some species is a sink for degraded products and Venus
energy but this neglects the fact that photo-
synthetic life uses premium radiation direct Abiological
earth
from the sun to sustain a high chemical potential models
gradient within the atmosphere on a planetary
scale. For a tree, the boundary within which Gas Venus Earth Mars D I
entropy is reduced is not its surface in contact
with the atmosphere but rather the interface CO, 90000 0.3 5 0.3-1 000 300
between the sun with the atmosphere as an N, 1000 780 0.05 1 30
extension of the tree. The tree produces not 0, 0 210 0.1 1 0.3
only food for consumers but also the equally
important gas, oxygen which does not accumu-
late within the tree waiting to be eaten. where the right hand side of the equation
When the whole assembly of life is so seen it expresses information in terms of temperature
is clear that the true boundary is space. The ( T ) pressure ( P ) internal energy ( E ) , volume
outgoing entropy flux from the Earth indeed ( V ) ,entropy (S),and the chemical potential ( N )
from Gaia ‘if she exists’, is long wavelength of the molecules present; it follows that informa-
infra red radiation to space. This then, is the tion is a measure of disequilibrium in the classic
physical justification for delineating the boun- sense and recognisability in the information
dary of life as the outer reaches of the atmos- theoretic sense.
phere. There is also to a lesser extent an inner By examining the extent to which the atmos-
boundary represented by the interface with phere is in chemical and physical disequilibrium
those inner parts of the Earth as yet unaffected both within itself and with the surface of the
by surface processes. We may now consider all Earth we have a measure of the extent to
that is encompassed by the bounds as putative which it is recognisable as a separate identity
life. Whether or not Gaia is real will depend against a neutral background equilibrium state.
upon the extent to which the entropy reduction Whether or not it is seen to be a component
within a compartment such as the atmosphere is part of Gaia will depend upon the size of the
recognisably different from the abiologicalsteady disequilibrium revealed.
state background.
On the matter of recognition a debt is owed
t o the fertile concept of information theory,
Thermodynamic evidence
Shannon & Weaver (1963). It has been demon- We can determine the degree of departure
strated, for example by Evans (1969) that the from a conceivable abiological steady state
classical properties, entropy and free energy, atmosphere of the Earth by two means; firstly,
have exact information theoretic equivalents. by comparing the Earth’s atmosphere with Mars
Thus the information (I)of a system can be and Venus which may be taken as representa-
defined as tive of lifeless planets and secondly by compar-
I =so-s ing the present atmosphere with the atmos-
phere to be expected if life were deleted from
where Sois the entropy of the components of the Earth.
system a t thermodynamic equilibrium and S the Table 1 shows the atmospheric composition
entropy of the system assembled. This relation- of Mars and Venus compared with that of a
ship can be transferred directly from informa- hypothetical abiological Earth interpolated be-
tion theoretic to classical thermodynamic terms tween them. Also shown is the atmospheric
as follows: composition to be expected of the Earth if
life were deleted. This was derived as follows:
I = ( E +PV - T S - Z N , N , ) / X nitrogen reacts under the influence of solar
Tellus XXVI (1974), 1-2
ATMOSPHERIC HOMEOSTASIS BY AND FOR THE BIOSPHERE 5
Table 2. Gases in the Earth’s atmosphere with origins in the Biosphere
The source strengths, residence times, concentrations,and concentrations expected if the atmosphere was a
thermodynamic equilibrium

Anticipated Departure Residence output


Atmospheric equilibrium from time megatons
Gas abundance concentration equilibrium years YR-1 Source

N, 0.8 E-10 E 10 3E6 E3 Denitrifying bacteria


CH, 1.5E-6 E-36 E 29 7 2E3 Anaerobic fermenting
bacteria
N,O 3E-7 E-20 E 13 10 632 Denitrifying bacteria
NH3 1E-8 E-35 E 21 E-2 1.6E3 Nearly all organisms
CH,I 1E-12 E-35 E 23 E-3 30 Marine algae

UV, electrical discharges, ionisation radiation nitrogen rather than NO, as the stable com-
and combustion with both 0, and CO, to give pound. That nitrogen has not yet been detected
oxides of nitrogen. By such processes in time on Mars is in agreement with the expectation
all nitrogen would be converted to the stable of equilibrium chemistry. A lifeless Earth
NO, ion dissolved in the oceans. At the pE would similarly be without atmospheric nitro-
even of Mars the reverse reactions converting gen. These arguments were used successfully t o
NO, to N, do not proceed. I n the course of the predict that Mars would lack N, before this
removal of N, most of the 0, also would go fact was discovered (Lovelock & Giffen, 1969).
from the atmosphere. The continuation of The anomalies of abundance are more marked
water and CO, photolysis would ensure the with the lesser gases &s is shown in Table 2
production of some oxygen, so that nitrogen which lists their concentrations, residence
removal could go to completion. I n the end the times, sources and expected concentrations,
atmosphere would apart from argon be domin- were chemical equilibrium determining.
ated by CO, like those of Mars and Venus. The simultaneous and large fluxes of N, CHI,
The model of a lifeless Earth coincides well NH,, N,O and 0, are all inconsistent with an
with that of a planet interpolated between Mars abiological model. The degree of departure from
and Venus, the slight chemical disequilibrium equilibrium expectations is to be measured in
is due to dissociative effect of sunlight and the 10’s of orders of magnitude. I n these circum-
escape of hydrogen. The present Earth is stances there is almost a certainty that the
clearly recognisable against the background of atmosphere is a part of the biosphere, for now-
the lifeless model. here, but in living systems is so intensive and
The terrestrial planets orbit in the outer constant a disequilibrium revealed. Conversely
reaches of the sun’s hydrogen atmosphere and there is a neglible possibility that the atmos-
their interiors are reducing hence any con- phere is a neutral background source of materials
centration of oxygen in a planetary atmosphere which life merely recycles.
is indicative of disequilibrium. Mars has 0.01 %
of oxygen in its atmosphere and Venus less
than this. Such small oxygen concentrations are
The atmosphere as a contrivance
quite consistent with the abiological steady I f we assume the Gaia hypothesis, and regard
state consequent upon the upper atmosphere the atmosphere as a contrivance, then it is
photolysis of water vapour and CO, on Mars reasonable to ask what is the function of its
and perhaps to a lesser extent on Venus. It is various component gases. Outside the Gaia
well to consider that Venus has probably lost hypothesis such a question would rightly be
oceans of water by photolysis and other reac- condemned as circular and illogical but in its
tions, yet still has no 0,. The presence of context such questions are no more unrea-
nitrogen on Venus does not indicate disequili- sonable than asking, for example, what is the
brium since Venus is both hot and acid and function of fibrinogen in blood. The function of
bereft of water, these are all conditions favouring oxygen and nitrogen in sustaining the cycle of
Tellus XXVI (1974), 1-2
6 JAMES E . LOVELOCK ET AL.

raw material and energy is evident. Less obvious the biosphere as is the shell to a snail or the fur
is the function for example of CHI. This gas is to a mink.
made on a scale requiring the shunting of as
much as 5 % of the photosynthetic energy of the
entire biosphere. I n the Gaia context it must
Systems analysis
surely have an important purpose of biology is One set of tests which can be applied t o
normally efficient and parsimonious over prove the existence of a control system are
waste. Such a possible important function for those which make changes in the variables
methane has been proposed (Lovelock & Lodge, thought to be controlled. It is possible to show
1972). It is seen as a kind of molecular hydrogen for example that the core temperature of a man
balloon whose purpose is to carry excess hydro- remains constant when the environmental
gen to the upper atmosphere where it can temperature ranges between 0 and 45°C. With
escape and thus sustain the present oxygen the Earth it is reasonable to suspect that a t
tension of the Earth. The details of this notion least the following variables may be controlled:
are given in the reference quoted. Wofsy et al., (1) Atmospheric, oceanic and soil chemical
(1972) suggest that such is the magnitude of the composition. (2) Surface temperatures in the
upward transfer of methane to the stratosphere 'core' region i.e. between 45" N and 45" S lati-
where it is oxidised to water vapour that the tudes. ( 3 ) Surface and ocean pH.
stratosphere is a source of water vapour to the It is not usually possible to vary the global
troposphere rather than a sink. It follows that environment as a test but perturbations affect-
much of the water vapour in the upper atmos- ing these variables have accompanied the evolu-
phere which is photolysed to produce oxygen is tion of the solar system. Since life began, 3.5 x
a biological product resulting from the oxidation 109 years ago, the solar output has probably
of biological methane. increased between 30 and 1 0 0 % (Sagan&
As with CH, the biosphere uses a great Mullen, 1972). The most recent estimates in
amount of energy for NH, production. When which the current low rate of neutrino emission
the total production of acids by the oxidation from the sun is taken into account is 70 %
of nitrogen and sulphur is taken into account (Dilke & Gough, 1972). It is even suggested that
the ammonia production by the biosphere is in addition to the slow exponential increase in
found to be just sufficient to sustain a rainfall solar output, fluctuations of 10 % may occur
p H near 8, the optimum for life. One penalty of over short periods i.e. 10 M years. The other
oxidising environments is the tendency t o low major environmental change is that of redox
p H as the elements such as carbon, nitrogen and potential. At its formation the Earth was
sulphur oxidise. The huge biological ammonia highly reducing with hydrogen as a predominant
production conveniently answers this need. Can atmospheric gas. It is suggested that 3.5 x lo9
it be accidental and have always been so? years ago the Earth's p E was - 5 a t which
Other atmospheric gases and vapours for gases such as CH,, NH,, H,S etc. would be
example dimethyl sulphide, dimethyl selenide stable in their own chemical right. Hydrogen
and methyl iodide may serve in the mass trans- would be escaping t o space but for some time
fer of essential elements between the land and there would be sufficient to regenerate the
the sea. hydrides dissociated by solar UV. Such an
A gas wholly in disequilibrium, a biological atmosphere could according to Sagan & Mullen
product on a grand scale, is nitrous oxide. It is (1972) effectively retain heat. It would permit
made by soil microorganisms and some pene- surface temperatures similar to those of present
trates to the stratosphere where it reacts to in spite of a much reduced solar output. This is
give NO,. By the Gaia hypothesis it must have due principally t o the radiative properties of
an important atmospheric purpose, could this gases such as ammonia and the others men-
be concerned with the regulation of the position tioned above. These were probably the condi-
or density of the ozone layer? I n the summary tions under which life evolved.
there is overwhelming evidence that the atmos- Soon after the evolution of life and possibly
phere apart from its content of noble gases is a because of it, Yeas (1972) the pace of hydrogen
biological product. It may also be a biological loss accelerated. By 3.2 x los years ago the pE
contrivance; not living but as essential a part of had risen above -4 a t which level NH, is no

Tellus XXVI (1974), 1-2


ATMOSPHERIC HOMEOSTASIS BY AND FOR THE BIOSPHERE 7
20

10
-
-w
0

o
r
3
%
5 -10
a
5
w

reducing oxidizing
F-20\ atmosphere atmosphere

T ~ o,Surface
, temperature predicted with
-301 solar luminosity increase af 309.
T70, ,Surface temperature piedccted with
solor luminosily increase of 709.
IW

Fig. 1. The mean surface temperature of the Earth t o 3.5 x lo8 years B. P. Shaded area, temperature range
deduced from the geological record and the persistence of life. Solid and dashed lines, the temperatures
expected for a lifeless Earth assuming a 70 % (solid line) or a 30 % (dashed line) increase in solar output.

longer stable in its own right (Sill&, 1966). Had tration of ammonia sufficient to sustain by its
the laws of chemistry and physics operated greenhouse effect equable and tolerable tempera-
soon after this time NH, and other polyatomic tures. This is supported by the fact that the
reducing gases would have vanished leaving an current biological ammonia production, which
atmosphere of nitrogen and CO, a t a concentra- is 2 x 10° tons per year, might in a neutral or
tion not substantially different from the reducing atmosphere have sustained the con-
present. If we assume that the solar output was centration of ammonia said to be needed to
in fact lower a t this time such an atmosphere provide the desired radiative properties namely
could not possibly have sustained present day by volume. However, if we assume that
temperatures the short term equilibrium tem- the solar output was indeed lower in remote
perature would have been -5 and - 16°C. If times, the mere blind production of a gas such
the large albedo of ice and snow cover is addi- as NH, by the biosphere would not have been
tionally taken into account the equilibrium likely to suffice in the control of surface tempera-
temperature could even have been lower. tures by its radiative properties. With the
Chemical changes of the evolving earth and reduced solar output and the possibility of posi-
the physical changes of its radiation environ- tive feedback on cooling, any failure of the
ment are shown in Fig. 1. The geological record ammonia harvest or the evolution of a vigorous
and the fact of the persistence of life provide NH, consumer, of which there are many today,
strong evidence that at no time in the past would have set in train a course for catastrophic
3.5 x lo8 years did the oceans freeze or the irreversible cooling. A mechanism which plau-
mean temperature rise above 50". Indeed the sibly can oppose an unfavourable trend is not
evidence suggests that apart from glacial enough, there must also be a system able t o
episodes, the temperature has always been sense the trend and actively control the means
constant. Glaciations do not usually affect for its opposition. It must be observed that
regions between latitudes 45" N and 45" S there are other 'greenhouse' gases than NH,.
where 70 % of the surface resides. CO, a t high concentrations could have per-
Sagan & Mullen (1972) propose that the con- formed a similar function as could other poly-
tinuous biological synthesis of ammonia before atomic gases. To sustain a constant temperature
oxygen appeared gave a n atmospheric concen- for 3 x loo years in the face of chemical and
Tellus XXVI (1974), 1-2
8 JAMES E . LOVELOCK ET AL.

physical environmental changes strongly implies


the presence of an active process for thermo-
stasis. Life a t its origin can be considered to have
fed on the blanket of gas which kept it warm.
I t s continued survival required the early devel-
opment of the capacity to recognise potentially
adverse changes and of processes which could
oppose such changes. It is not the purpose of
this article to consider in detail the exact
mechanism which could have been used to this
end. The change of atmospheric composition is
a powerful technique but other changes such
as in the surface albedo the rate of evaporation
of water, are a t least conceivable. A more
detailed account of speculations on possible
mechanisms is given in a forthcoming paper I I I I I
(Margulis & Lovelock, 1973). Our purpose is to
stress that the present knowledge of the early
environment suggests strongly that a first task
of life was to secure the environment against
adverse physical and chemical change. Such
security could only come from the active
are still not known although it is conceivable
process of homeostasis in which unfavourable
that the CO, concentration rose as the p H fell
tendencies could be sensed and counter mea-
and to some extent offset the effects of the loss
sures operated before irreversible damage had
of NH,. It is unlikely however that this alone
been done.
would have been sufficient to sustain tolerable
I n addition to the possibility of remorseless
temperatures a t this period when the solar out-
environmental change set by the evolution of
put may have been a t least 10 and possibly 40 %
the sun, there were also major changes in the
less than now. Nevertheless sufficiently constant
chemical environment. These were caused in
temperatures were sustained and life persisted.
part by the evolution of life itself but they also
Even today the regulation of oxygen concentra-
reflected upon the physical environment through
tion may represent a n important task. Fig. 2
the change in atmosphere composition. Promin-
shows how drastically the probability of the
ent amongst these changes must have been the
ignition of inflammable material rises with
first appearance of free oxygen in the atmosphere
increasing oxygen concentration. Above 25 %
which is thought to have taken place between
even trees in rain forests might be in hazard
1 and 2 x 109 years ago. It is worth digressing a
from fire. With these crises as with those
moment to consider what a catastrophic air
earlier the blind adaptation t o change seems
pollutant oxygen must have been. Its presence
unlikely to have been enough. ,
in the air must have destroyed a vast range of
species and driven underground into anaerobic
muds others from which prison they have never Conclusions
returned. Not only was the appearance of
oxygen itself a biological crisis but also the We have presented some of the evidence con-
rapid reduction of p H which inevitably would cerning the Gaia hypothesis. The hard core geo-
accompany oxidation of nitrogen and sulphur chemists will still no doubt argue that the
compounds and the increased rate of removal of cycling of gases through the biosphere is a
ammonia must have posed other serious pro- passive process and does not determine atmos-
blems. Firstly in the maintenance of p H itself pheric composition. He may compare it with
and secondly in sustaining a sufficient concen- the cycling of water from the oceans to the land.
tration of ammonia as this gas was depleted by The delivery of rain to the land surfaces involves
its new role of neutralising acidity. Again the a disequilibrium of gravitational and osmotic
details of the atmospheric changes of this time potential and is driven by solar energy. The
Tellus XXVI (1974), 1-2
ATMOSPHERIC HOMEOSTASiS BY AND FOR THE BIOSPHERE 9

vegetation merely borrows the rain and repays The purpose of this paper is t o introduce the
at the rate of borrowing. We think t h a t it is Gaia hypothesis at least for entertainment and
all a matter of degree. Sunlight may distil for the induction of new questions about the
water from the sea later t o fall as rain on the Earth. Proof of Gaia’s existence may never
land but sunlight does not spontaneously at the approach certainty but further evidence is more
Earth’s surface split oxygen from water and likely t o come from the study of the contempo-
drive reactions leading t o the synthesis of rary Earth. Astronomical evidence is notori-
intricate compounds and structures. The dis- ously fickle and although geological evidence is
equilibrium characteristic of water cycling is rather more certain one learns less about a
minor when compared with t h a t of cycling of person from the study of his grandfather’s
the atmospheric gases. bones than from talking t o him face t o face.
To those who are convinced that the atmos-
pheric gases are biological products but are
reluctant t o accept the notion of homeostasis Acknowledgement
we say: if life has merely a passive role in cycling
the gases of the air then the concentrations will We acknowledge the unstinted help and ad-
be set by equilibrium chemistry; in fact they vise of our colleagues s. R. Epton, P. B. Fell-
most certainly are not. If life actively cycles the gett, J. P. Lodge and P. G. Simmonds. We
gases then we ask how could such a system be thank Shell Research Ltd., for financial support
stable in the long run without homeostasis? t o one of us (J.E. L.)

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Tellus XXVI (1974), 1-2
10 JAMES E. LOVELOCK ET AL.

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Tellus XXVI (1974), 1-2

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