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THE GAlA HYPOTHESIS'

CAN IT BE TESTED?

James w. Kirchner

Energyand ResourcesGroup Universityof California Berkeley

Abstract. The Gaia hypothesis' centralthemeis that between biotic and physical processesshould create biological processeshomeostatically maintain, on a feedback loops. The speculative formsof Gaia assert that planetary scale, geochemical and climatic conditions biologicalprocesses regulatethe physicalenvironment, favorable life. A number distinct for of hypotheses have keeping Earth'sclimateand surface geochemistry stable beenproposed, spanning rangefrom the self-evident a to and favorablefor life. As metaphors, theseversions of the highly speculative. The self-evidentforms of Gaia Gaiaare intriguing, untestable, if takenliterallyas a and, reiterate well-documented thatbiological the fact processes basis research, for potentially misleading. hypotheses, As are critical to biogeochemicalcycles, adding the theyareill-defined,unparsimonious, unfalsifiable. and straightforward (though importan0 pointthatthecoupling

INTRODUCTION

important and widely recognized biologically mediated

processes photosynthesis; aerobicand anaerobicmetabo"Geology is a particularlyalluring field for premature lism; denitrification; nitrogenfixation; bacterial sulfate attemptsat the explanationof imperfectly understood reduction and iron oxidation; mineraldissolution by data." (J. D. Dana) CO2-enriched groundwater; sedimentation organic of carbon, carbonates,silicates, phosphates,and other "Geologyhas to choose between rashness using materials; changes surface the of and in albedo produced by imperfect evidence or the sterility of uncorrelated, growthanddecayof vegetation.Theseprocesses create unexplained facts." (J. W. Gregory) important feedbackloops linking natural selection and

evolutionwithin the biota and biologicallyinduced changesin the physical environment[Cloud, 1968; Schneider Londer, and 1984]. Perhaps because study the It hasbeen clearfor some timethatbiological processes of these linkages inherently is interdisciplinary, have they are crucialfactorsin the Earth's surface geochemistry. not receivedthe emphasis that their importance in When Huxley [1877] wrote his textbookin physical geochemical and geophysical processes would seem to
(quoted VanHouten[1961,p. 89]) by

geography, could describe role of greenplantsin he the

demand.

A persistent undercurrent some related work is the in limitingatmospheric 2 concentrations a matterof CO as established fact. By the time Hutchinson[1954] reviewed observation that the physicalenvironment seemsremark-

thebiogeochemistry theterrestrial of atmosphere, could ably well suitedto the needsof terrestrial he life [e.g., add 4, N2,NH3, +, N20 NO2, 8042-, CH NH4 , H2, and Henderson, 1913] and that this fact might evenreflect formaldehyde the list of biogenicor biologically organisms to manipulating their environment meet their to controlled atmospheric substances. Harvey[1957]devoted needs [e.g., Redfield, 1958]. Themost recent, comprehentwochapters biological to influences oceanic on chemistry. sive, and controversialform of this idea is Lovelock and Sincethe 1950s, efforts two fronts on (unraveling contem- Margulis' Gaia hypothesis, which states [Lovelockand porary biogeochemical cycles andtracing geophysical, Watson,1982, p. 795] that "the climateand the chemical the geochemical, and biological evolutionof paleoclimate) composition theEarth'ssurface keptin homeostasis of are have produced explosion new insightstoo vast to at an optimumby and for the biosphere"(see also an of adequatelydocumenthere; reviews can be found by Margulisand Lovelock [1974],Lovelock and Margulls Holland [1978, 1984], Schopf [1983], and others. [1974a,b] andLovelock [1979a, 1986b,1988]). In this Following Bergeret al. [1984]we canlist among most viewthebiota the andthephysical environment sotightly are

Copyright 1989 by the American Geophysical Union


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224 Kirchner: THE GAlA HYPOTHESIS

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coupledthat they can be considered singlevastorganism a [Lovelock, 1986a, b, 1988] called Gaia (after the Greek goddess the Earth), and the atmosphere be considof can ered to be [Lovelock,1979b,p. 716] "like the fur of a cat and the shell of a snail, not living but madeby living cells soas to protectthemagainst unfavorable an environment." The Gaia hypothesis receiveda greatdeal of public has attention. I can only speculate why: it is a hypothesis of great generalityand vast scope;it has been presented in nontechnical language; it addressesmatters of both academicand practical concern; it suggests colorful a metaphor that many find intriguing;and its semantic and conceptual plasticity allow it to be virtuallyall things all to people. Two groups thatimmediately embraced Gaia were environmentalists and, paradoxically,industrialists. The former argued that harming any part of the planetary "organism"couldhavefar-reaching consequences, while thelatterargued thatGaia'scapacity homeostasis for made pollutioncontrolunnecessary [Schneider, 1987]. If verified, the Gaia hypothesiswould representa revolution in biogeochemistry, for it represents funa damentallydifferent view of the role of life on Earth. Some have spontaneously embracedit as obvious, and manyhavespontaneously dismissed as bizarre. Others, it representing diversearrayof specialities, a havewondered how the hypothesiscould be empirically tested; this questionwas the focus of the recent AGU Chapman
Conference.

them all into a singlecoherentstatement would be at best unwiseand at worst impossible. Instead,I have tried to compile a taxonomy of the central themes of different statements the Gaia hypothesis order to state each of in precisely to clarify the distinctions and between them.

Coevolutionary Gaia
The biota influences its abiotic environment, and the
environment in turn influences the evolution of the biota

by Darwinianprocesses. Watsonand Lovelock[1983, p.


284] state,

The biota have effected profound changeson the


environment of the surface of the earth. At the same

time, that environment has imposedconstraints the on biota, so that life and the environment may be consideredas two partsof a coupledsystem... perturbations of one will affect the other and this may in turn feed back on the originalchange. The feedback may tend eitherto enhance to diminishthe initial perturbation, or depending whether signis positive negative. on its or
Homeostatic Gaia

The biotainfluences abioticworld in a way that is the stabilizing. The dominantlinkagesbetweenthe biota and thephysical worldarenegative feedback loops.
Lovelock andMargulis [1974a,p. 93]
From the fossil record it can be deduced that stable

optimalconditions the biosphere for haveprevailedfor My primarypurpose here is not to catalogue and weigh thousands millions of years. We believethat these of the evidence and againstthe Gaia hypothesis. for Instead, properties of the terrestrial atmosphereare best I want to address more basicissue: Is the Gaia hypothea interpreted evidence homeostasis a planetary as of on sis a scientifically testable propositionat all? This scalemaintained life on the surface. by approachmay disappointthose who would prefer an introduction to the relevant biogeochemical and Lovelock Margulis [1974b, p. 3] and paleoclimatictheory and data. Nevertheless, first the The notion of the biosphere an active adaptive as question askof a theoryis not whether is trueor false, to it controlsystem able to maintainthe earthin homeostasis but what it means and whether it can be tested. we arecallingthe "Gaia" Hypothesis. I suspect that muchof the ongoingdebatesurrounding Gaia has arisen because different parties defined the Lovelock [1988, p. 13] hypothesis differentways,assumed their interpretain that Through Gaia theory,I seetheEarth andthe life it bears tionsof the hypothesis were scientifically meaningful and as a system, systemthat has the capacityto regulate a empiricallytestable, and thenproceeded argueover the to the temperatureand the compositionof the Earth's data. My premiseis that much of the debateover Gala surface to keepit comfortable livingorganisms. and for doesnot resultfrom the ambiguityof the geologicrecord or the gaps in our current understanding global of biogeochemical cycles,but instead derivesfrom a lack of Geophysiological Gaia clarity concerning what the Gaia hypothesis meansand The biosphere be compared can with a singleimmense
how (or whether) it can be tested.

organism which, like other organisms, may exhibit both


homeostatic and unstable behavior.

A TAXONOMY

OF GAlA

HYPOTHESES

Lovelock [1986b,pp. 12 and 19]

Defining the Gaia hypothesis difficult. So many is logically distinct theorieshave been put forth under the singlebannerof "the Gaia hypothesis"that compressing

Gala theorysuggests we inhabitand are part of a that quasi-livingentity that has the capacityfor global homeostasis. is thebasis geophysiology...a This for

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THE GAlA

HYPOTHESIS'225

argumentmay result from a simple systems approach Earth science. It is the essential deal of unnecessary to theoretical basisfor the putativeprofession planetary misunderstanding which hypothesis on the table at of of is medicine. any given time. The wide variety of Gaia hypotheses createsa risk of Lovelock[1988, p. 13] misinterpreting evidencefor one versionas somehow also proving another form of Gaia as well. For example, has it Gaia theory predictsthat the climate and chemical is composition the Earth are kept in homeostasis of for longbeenknownthat the atmosphere in greaterchemical long periods until some internal contradiction or disequilibrium than would be produced extraterrestrial by external forcecauses jump to a new stable a state. radiation alone[e.g.,Lewis and Randall, 1923]. This is, as

OptimizingGaia

The biota manipulates physicalenvironment ways its in minants of Earth's atmosphericchemistry. Lovelock thatcreate biologically favorable conditions. [1965] andHitchcock andLovelock[1967] pointedout that spectroscopic data shoulddetect chemicaldisequilibrium Lovelock andMargulis [1974a,p. 93] in the atmospheres other planets if life is present. of Lovelock's[1975] predictionthat life would not be found We arguethat it is unlikelythat chance aloneaccounts supportedby data from the for the fact that temperature, and the presence pH of on Mars was subsequently chemicaldisequilibrium is compoundsof nutrient elements have been, for Viking program. Atmospheric immense periodsof time,just thoseoptimalfor surface not, however, evidence that the biota manipulatesthe life. Ratherwe present "Gaia hypothesis" idea atmosphere any particular way or for any particular the the in that energy is expendedby the biota to actively reason. maintaintheseoptima. The many Gaia hypotheses make it easyto createfalse dichotomies.Rejectingone null hypothesis (e.g., that the Lovelock and Watson[1982, p. 795] biota has no effect on its physicalenvironment)does not The Gaia hypothesis... postulates the climateand provean alternativehypothesis that (e.g., that the biota and the chemical composition the Earth'ssurface kept in physical environmentform a goal-seeking,homeostatic of are homeostasis an optimumby andfor thebiosphere. at cybernetic control system)unlessthe two hypotheses are mutuallyexclusiveand jointly exhaustive (which, in this Lovelock[1988, p. 8] example, theyare not). Nonetheless, Lovelockand Watson [1982, p. 799] (see alsoLovelock[1979a], Lovelockand Planetarylife must be able to regulateits climate and chemical state... the greaterpart of our own environ- Margulis [1974a, b], and Margulis and Lovelock[1974]) menton earthis alwaysperfectandcomfortable life. proposethat "a test for Gaia is to considerwhat would for The energyof sunlight so well shared is that regulation happenif life were now deletedfrom the Earth." This is, is, effectively,free of charge. of course,a test for life, not a test for Gaia. A test for Gaia
Lovelock[1979a, p. 127]
should look for evidence that a lifeless Earth would be

Hutchinson [1954], Sillen [1966], and others have noted, evidence that biological processesare important deter-

The mostimportant property Gaia is the tendency of to optimizeconditions all terrestrial for life.
Some may take issuewith this taxonomy. This is not the only way to classifythe Gaia hypotheses, perhaps or eventhe best. My point is simplythatit needsto be done, because singleterm "Gaia" hasbeenusedto referboth the to observations manyfind self-evident to proposithat and tionsthatmanyfind extremely speculative. This wide rangeof possibilities, alongwith theengaging plasticityof the terminology, meansthat individuals can make of Gaia whateverthey wish. The unsympathetic can ridiculethe notionof globaloptimalcontrol. The sympatheticcan point out that the biota and the abioticenvironment are obviouslyinterrelatedin any numberof ways. The uncritical(particularlyamongthe publicandthe press) can take data detailing these interrelationships and misinterpret them as scientificvalidationof their fanciful extrapolations Gaia. And in scientific of discourse great a

differentin the way that Gaia saysa lifelessEarth should be different (that it should be less homeostatic,for

example),not just differentsomehow. Many arguments for Gaia are basedon rejectingthe hypothesis that biotic processes are completely irrelevant to the physical environment; hypothesis alreadybeenrejected this has by manyinvestigators quitesometime. for In order to proceedI must briefly summarizesome relevantepistemology scientificmethodology.These and pointsare familiar to manyresearchers, it will aid the but present discussion theyare stated if explicitly.

HYPOTHESES: TRUE, FALSE,AND UNTEsTABLE

The day-to-daybusiness scienceconsists testing of of hypotheses, somehypotheses but cannotbe tested. Some hypotheses untestable practice,thoughthe practical are in impossibility performingan exact test is not crucialif of surrogate canbe devised.For example, may hope tests one

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Somecriteriafor judging the usefulness hypotheses of can also be stated. Some hypotheses, while testableand be observed other scales,suchas in large forest fires. perhapseven correct, are simply not very useful. The at predictions derivedfrom a Other hypotheses (for example,the "hypothesis"that in more specificthe confirmatory 4000 B.C. a deviousgod assembled geological the record hypothesis, the moregeneralthe falsifyingpredictions, and in a way that gives the impression the Earth is much the morelogicalcontent hypothesis that the contains [Popper, older) are unrestable principle;thesecan obstructthe 1959, p. 113]. Hypotheses usefulto the degreethat in are progress science.In thispaper,unless of otherwise noted, theyare logicallydistinctfrom othertheories; a hypotheif the terms "testable" and "unrestable" will refer to sis simplyrestates widely accepted principles,or can be testabilityin principle. directlyderivedfrom them,a search confirmatory for data The minimal criteria of testability can be stated would be an inefficient use of scarce resources. are to concisely. In order to be testablea hypothesis must be Hypotheses alsousefulin proportion the phenomena clear, and its terms must be unambiguous. It must be that they alone can predictor explain and, perhaps more intelligiblein termsof observable phenomena.And most importantly, in inverse proportion to what they must importantly,it must generatepredictions two kinds: assume.This is the well-knownLaw of Parsimony:if two of confirmatory predictions (phenomenathat should be hypotheses explainthe samebody of data equally well, observed the hypothesis true and that would not be prefer the theory that imposesthe lightest burden of if is [Poincare, 1905, pp. 146 and 151]. While predictedby the existingbody of accepted theory) and assumptions falsifyingpredictions (phenomena should observed this principle cannot be precisely codified (and in a that be if the hypothesis false). is borderline casemay reveal the choiceamongcompeting to taste),it remainsa basic A hypothesis is unrestable muchlessusefulthan theories be a matterof aesthetic that is one that is merely false. A false hypothesis, onceknown tenetof the scientificenterprise. to be false, at least helps to restrictthe sphereof posnever be carried out, but some of its basic mechanismscan sibilities. Unrestable theories, on the other hand, are at

that a full-scale test of the nuclearwinter hypothesis will

best empty and at worst misleading, and in the mindsof the unwarythey can be entrancing.A few examples will sufficeto makethispoint. 1. Ill-defined hypotheses unrestable are becausethey canbe endlessly reinterpreted fit almostany data,but for to the same reasonthey cannot containspecificempirical information; theyexcludeno possibilities. 2. Tautological hypotheses unrestable are because they are true by definition, but as such,they containno new information; their conclusions are entirely containedin theirpremises. 3. Unfalsifiablehypotheses unrestable are because they makeno falsifyingpredictions, as such,they haveno but empiricalcontent; confirmation the hypothesis of doesnot restrict the sphere of possibilitiesbecausethe set of excluded data(datathat wouldhavebeeninconsistent with the hypothesis) empty. is Note thathypotheses eachof these in classes (whichare notlogically distinct, arestated way in theinterests but this of simplicity) will usually show good agreement with experimental data. That is why theyare misleading.One believes they fit the data so well because theycapture the essence the problem,whenin fact they fit because of they are independent the empirical facts. For the same of reasonthey are entrancing; one thinksit remarkable that the predictions are always confirmed. Attemptsto test such theoriesare ultimately futile, not becausethey are doomed to failure, but because they are doomed to inevitable andinconsequential success.

GAlA

AS METAPHOR

Somehave suggested Gaia is simply a metaphor that rather than a hypothesis. Metaphorsare unrestable, but they can nevertheless scientificallyuseful; they can be suggestnew lines of analysis, such as the search for biogenic sources cloudcondensation of nuclei[Charlson et al., 1987]. Indeed,muchof science, relyingas it doeson reasoning by analogy, is deeply metaphorical. As scientificmetaphors Gaia is unusuallycolorful, rich, go, andevocative, I hopethatit will spurmanyinteresting and and fruitful speculations. But metaphors hypotheses and are two differentthings,and it is importantnot to confuse
them.

Why is it misleading treata metaphor an empirito as cally testable proposition?As hypotheses, metaphors are ill-definedbecause they can be reinterpreted explain to almostany observed behavior; they fail to specifyin what sense the metaphor is true. One can agree with Shakespeare all the world is a stage,in somesense that (e.g., its inhabitants can be viewed as playing out their roles),but it is not a stagein all senses (e.g., it is not made of flooring and does not have a row of footlightsat its edge). There is, doubtless,some sensein which the biosphere can be considered an organism[Lovelock, as 1986a, b, 1988], but this analogy is not scientifically verifiablewithoutsomestipulation how it doesor does of not apply. Proposalsof specific ways in which the biosphere mightact like a globalorganism (e.g., it might

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Kirchner: THE GAlAHYPOTHESIS.227

exhibit homeostasis) of course, are, hypotheses may be and testable. The metaphor itself, however,is untestable.It hasno empiricalcontentbeyondthe data that suggest the specific waysin whichit doesor doesnotapply. Boston[1989] hasnotedthat Gaia can be interpreted as illustratingthe principle of emergentproperties,which holds that a complex system can have propertiesthat would not be predicted from studies of its isolated components. As a cautionarynote to those who view systems simplistically, this principleis well taken. As a hypothesis se, however,it is ill-defined. Even simple per abiotic systemscan exhibit emergentproperties;it is precisely which properties are predicted and which mechanisms createthemthatconstitutes hypothesis. a If Gaia is meantto be simplya metaphor a restateor mentof the principleof emergent properties, is puzzling it that its proponents make so manyreferences the Gaia to hypothesis, tests for Gaia, and proofs of Gaia [e.g.,

are interrelated (that is, eachhaseffectson the other), there are no otherpossibilities exceptthat feedback, positiveor negative, characterizes relationship. this

HOMEOSTATIC

GAlA

Gaia could be taken simply to mean that someof the interactions betweenthe biota and the physicalenvironment are stabilizing (i.e., characterizedby negative feedback). If this is so, it is not surprising. Given that thesefeedbackloopsare numerous and that each mustbe either stabilizingor destabilizing, is highly likely that it someare stabilizing. Homeostatic Gaia can, however,be stated in two nontrivial versions: a weak form (the
dominant interactions between the biotic and the abiotic

worlds are stabilizing) and a somewhatstrongerclaim (these interactionsmake Earth's physical environment Lovelock, 1983, 1988; Lovelock and Watson, 1982; significantlymore stablethan it would have been without Lovelockand Margulis, 1974b; Margulis and Lovelock, life). 1974]. If Gaia is not intended a scientific as proposition, it Note that the simplefact of climatichomeostasis the (to extent that climatic and paleoclimaticdata do or do not is misleading referto it as if it wereone. to indicate stable conditions, topicwell beyondthispaper's a scope) not at issue. Gaia concerns is mechanisms through COEVOLUTIONARY GAlA whichhomeostasis thought be maintained, just the is to not
fact that it exists. Evidence of homeostasis is therefore not

The idea that the biota and the physicalenvironment coevolve is not original or unique to Gaia. To my knowledge,it was first statedby Spencer[1844, p. 93], who held that the biogenicincrease atmospheric in oxygen concentration made possible "more perfectmental and a bodily development"in the biota. While Spencer's accountis simultaneously grander and cruder than the contemporary view,few woulddispute general its premise; bioticprocesses shapethe physicalenvironment, which in turnguides bioticevolution.Indeed,thefact thatthebiotic and physicalworlds are interrelatedis by now so well
documented [Hutchinson, 1954; Berkner and Marshall, 1964; Cloud, 1968; Holland, 1964, 1978, 1984; Schneider and Londer, 1984] that it would seem odd to call it a

compelling evidence Gaia; how canwe tell whetherthe for climate has been stablebecause biotic processes in of or spiteof them? The point has been made repeatedly(for example,by
Lovelock and Watson [1982], Lovelock [1979a], and

Margulis andLovelock[1974]) thatcrudephysicalmodels of climatecannot explainwhy theEarthwaswarmenough to supportlife during the Precambrian, when the Sun is thought havebeenapproximately to 25% dimmerthanit is now [Saganand Mullen, 1972;NewmanandRood, 1977].
This in itself is not a test of Gaia. The limitations of

hypothesisat all. An observationthat is so widely recognized lacks the tentative character of a true hypothesis. Coevolutionary Gaia asserts just that the biota and not the physical environment interrelated alsothat this are but relationshipis characterized by negative and positive
feedback [Lovelock and Watson, 1982; Watson and Lovelock, 1983]. While the observation that interrelated

systemcomponents exhibit feedbackis useful for those unfamiliar with feedback processes,it also follows analytically from elementary theorems systems of analysis andhasno specificempiricalcontent. An empiricaltestis therefore unnecessary. the biota and the physicalworld If

simple models (which assume that the Precambrian atmospherehad the same compositionas at present, despite clearevidence higherpartialpressures oneor for of more greenhouse gases)do not necessarily imply that biological control mechanisms [e.g., Lovelock and Whifi'eld, 1982] are neededto resolvethe "faint young Sun" paradox.The simplest solution the paradox to may be refined geophysical climate models,such as thoseof Berneret al. [1983]andWalkeret al. [1981]. It is easyto speculateabout hypotheticalbiological mechanisms that resolve the faint young Sun paradox and other incongruitiesin the climatichistoryof the Earth. The hardwork

liesin specifying plausible, a parsimonious, quantitatively realisticmechanism sufficientto dispose the paradox of andproposing completing testof that theory. One and a attempt to include quantitatively realistic biological

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feedbacksin models of global temperatureregulation resulting increased 2 release) in CO wouldamplifythe


[Volk, 1987] concludedthat their effect would be small. A temperature expected rise froma greenhouse warming. wide varietyof hypothetical controlmechanisms couldbe It is generally agreed that oxygen productionby speculatively invoked to explain any given discrepancy blue-green"algae" (recently reclassifiedas bacteria) between existing theory and data, but they should be changed redox the balance thePrecambrian of atmosphere testable against morethanjust thegapstheywereselected fromreducing oxidizing, a demonstration biologito in of to fill. cal controlthat was clearly not homeostatic (and which .Lovelock Margulis and [1974a, 100-101] pp. and virtually exterminated microbesresponsible it). the for MarguliandLovelock [1974,pp. 479-486] proposed Lovelock[1979a, p. 31] and Lovelockand Margulis a life wide rangeof biological feedback mechanisms might [1974b,p. 8] cite the fact that terrestrial survivedthis that controltheclimate. Conspicuously absent from thebulk of event evidence Gaia'sresilient as for adaptation change. to theensuing research, however, any mention processes If themost is of destabilizing bioticeventin Earth'shistory can that might destabilizethe climate. For example, in a beconstrued evidence Gaia,andtherelative as for stability review of the role of oceanicphytoplankton producing since then can also be cited as evidencefor Gaia, one in cloudcondensation nucleiand alteringthe globalradiation wonders what conceivable events couldnot be interpreted budget(the Gaian mechanism climate regulationthat as supporting Gaia hypothesis. thereare none,Gaia of the If hasreceivedthe greatest attention date),Charlson al. cannot tested to et be against geologic the record. If Gaia stabilizes and Gaia alestabilizes (sinceGaia is a [1987] note in passingthat it is unclearwhetherthe resultingfeedback loop is stabilizing or destabilizing. feedback system, thoseare the only two possibilities), is They thendiscuss lengthhow stabilizing at feedback might there possible any behavior is notGaian?Some that might arise and how it might be instrumental regulating object that the biota's destabilizing effect in the in climate they do not addressthe possibleorigins and Precambrian :, indicates Gaia hadnot fully matured, that but such objection openlytautological; defines an is it Gaia in consequenc.destabilizing esof feedback. Without knowing what biologicalmechanisms undermine may homeostasis, terms of homeostasis and then asserts that Gaia is
it is difficult to make a balanced assessment of the role of
the biota.

homeostatic.

In the absence of clear evidence that biotic mechanisms

An interesting outgrowth the Gaia hypothesis of has beena mathematical modeldemonstrating homeostatic that control of climate. behavior be achieved a biotic system can in withoutany Lashof[1989] hasestimated climaticandbiogeochemi- form of conscious control. The Daisyworld model cal feedbackresponses an anthropogenic to greenhouse [Watson Lovelock, and 1983; Lovelock, 1988],designed to warming. Of the five biologically mediated feedbacks that be heuristic rather than realistic,describes planet on a he examines,only one (an increasein photosynthesis which temperature controlled the albedo,which is is by by resulting from directCO2 fertilization) offsets effects determined the color of the daisiesgrowingon the the is how the of an anthropogenic doubling atmospheric 2. The surface. This model'spurpose not to describe of CO other four biologicallymediatedprocesses decrease (a in Earth'stemperature mightactually havebeenregulated in vegetation albedoas the tree line movesnorth,an increase thefaceof changing solarluminosity; instead, Daisyworld in plant respirationand metabolismof soil carbon, an is intended explore consequencesstrong to the of coupling increasein methanogenesis, and shifts in ocean biota between biotaandthephysical the environment.

Recently, ice core data have shown that variationsin biogenic cloud condensation nuclei and carbon dioxide serve to amplify, not damp, the temperature excursions during glacial periods. Levels of nonmarinesulfate (a proxy for biogenicdimethylsulfide,a precursor cloud of condensation nuclei [Charlson et al., 1987]) are much higher during glacial periods than during interglacials [Legrandet al., 1988]. Similarly, carbondioxide levels are significantlyhigher during interglacials than glacials [Barnola et al., 1987; Jouzel et al., 1987]. These are the two most climaticallyinfluentialbiogeniccompounds for which long-term recordsare available. Those records indicatethat both of thesebiologicallymediated feedback relationshipsare either (1) ineffective in offsetting nonbiologicalpositive feedback processescontrolling these compounds [Schwartz,1988] or (2) actively destabilizingin their own right. Neitherof thesepossibilities supportsthe notion of homeostasis through biological

do in fact stabilize globalenvironment, seems the it wiseto refrainfrom assuming theyoughtto do so. It is wiser that still, I think, to avoid the assumption biotic mechathat nismsshould generally be eitherstabilizing destabilizor ing, or thatthe stability properties these of mechanisms are theirmostinteresting important or features. Before addressing hypotheses the that I have termed geophysiological and optimizing Gaia Gaia, I wouldlike to pausehere to commenton mathematical modelsof Gaia and whetherthey demonstrate biologicalhomeostasis of
the environment.

MODELS

OF GAlA

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Kirchner' THEG^I^HYPOTHESlS

229

The theory behind Daisyworldcan be outlined as [Watson and Lovelock, 1983, p. 288] (italics in the a of follows. Bioticgrowthcurves a function environ- original)indicates selectivity emphasis. There is a (as of of for mental variables such temperature) characteristicallyrange luminosities whichthe temperature/luminosity as are peaked.If the variable interest a function biotic curve is flatter than it would have been without daisies,but of is of populations, thereis somerangeof changes external there are also rangeswhere it is much steeper(as simple in conditions for which the effect of the induced shift in continuity between the endpointsrequires). If the pointis neareitherendof the stable range,small bioticpopulations offsets imposed the change (Figure1). operating shifts in luminosity can have drastic consequences.
8O T1

Perturbations could also cause the system to suddenly jump fromonepathof thehysteresis loopsto another.The sense which suchbehaviorindicates"greater stability" in
is unclear.
T2

60
>,, 40

e
.--.

20

Likewise, the claim [Lovelock,1988, p. 46] that "in no way is the stability of Daisyworld dependenton an idiosyncratic choice of initial values, or rate constants" needsto be qualified. Daisyworld exhibitsstabilitynear the optimal temperaturefor daisies becausethe black daisies, whichwarm the surface, thrive in coolertempera30 40 50

10

20

tures than the white daisies, which cool it. But what if,

Mean surface temperature (C)

instead,the optimal temperature black daisieswere for

Figure1. Theeffectof temperature thepopulation daisies on of


(B) andtheeffectof daisycoveron themeantemperature the in Daisyworld model(seetext)for twovalues solarluminosity of (A andA1). This example assumes only white daisies that are present; increasing daisycoverraises albedo and lowersmean temperature. solarluminosity If increases from A to A1, the resultingtemperature rise increases daisy cover and raises albedo, limitingthetemperature fromT to T1. If daisy rise cover did not respond temperature, shift in luminosity to the would result a temperature in increase T2. Parameter to values used are

15C higher that white than for daisies? higher (A optimal


temperature might explain why the black daisiesevolved

to be 10C"warmer"in colorwhilesharing same the


environmentas white daisies.) If this were true, black daisieswould thrive in warmer temperatures and would tend to keep the surfacewarm (and vice versa for white

daisies).The resultis a tug-of-warbetween two types the of daisies. As Figure 3 illustrates, Daisyworldregulated a loop, where each value of solar thoseof Watsonand Lovelock[1983]. Thesecurvesdiffer from in this way is a hysteresis thoseof Lovelock[1986b, p. 15] because latter are mis- luminosity gives two equilibrium temperatures(each the labeled. What Lovelockrefers to as "daisy population"is representing complete extinction oneor theothertypeof of actually onefactorin the growth equation , rate [Watson and daisies)that lie well outsidethe optimal temperatures for Lovelock,1983, equation(1)]. What Lovelockrefers to as wouldlikely result "meantemperature" actually temperature the daisies. eitherspecies.Climaticperturbations is the of Mean temperature a function the daisytemperature, is of the in unstable transitions betweenthe upperequilibrium,in temperaturebareground, thefractional of and daisy cover.The whicha too hot Daisyworldtriesto keep itselftoo hot, and modelis verystable because selected the parameters makedaisy in tries covera nearlyverticalfunction temperature the operating thelowerequilibrium, whicha toocoldDaisyworld of at pointT. Daisycovershiftsfromzeroto 45% in response a to keepitself too cold. to temperature change only IC. of What is most interestingabout this "pathological" Daisyworldthat I have described that the availabledata is suggest that the biotic feedbacksregulating climate Thisresult general holds is and whether daisies the warm or coolthe surface.Figure2 shows behavior the actually work this way. As pointed out above, ice core the of model whenbothlightanddarkdaisies assumed be analyses are to indicatethat during glacial periods,biological produce less carbon dioxide and more cloud present; surface the temperature held nearlyconstant processes is over wide excursions solarluminosity. in condensation nuclei. In otherwords,biological processes appear to make the planet colder when it is cold and It is easyto seehow the resultin Figure2 is obtained. Black and white daisiesare assumed have exactly the warmer when it is warm. Dickinson and Hanson [1984] to same growth response temperature, blackdaisies to but are and Hansen et al. [1984] have estimated how Earth's assumed be 10Cwarmer to thanwhitedaisies equiva- vegetationpatternswould shift in responseto global in changes; they foundthat planetaryvegetation lent ambient conditions. Consequently, black daisies temperature thrive in cooler ambient conditions,lower the albedo, and albedo would shift in ways that exaggerate, rather than change. Of course,the warm the surface,while the oppositeis true of white offset, the imposedtemperature daisies. Daisyworld modelis heuristic and is not meantto imply Watsonand Lovelock'sconclusion that "regardlessof that the Earth's temperature actually regulatedby is changes vegetationalbedo. Nevertheless, the extent in to the directions thefeedbacks,the model always shows of albedoactuallydoesinfluenceclimate,the greaterstabilitywith daisiesthan it doeswithoutthem" that vegetation

230'

Kirchner: THE GAIA HYPOTHESIS


80

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60

, 40

' 0.5

0.5

1.0

1.5

Solar luminosity (arbitrary units)

and cooler,owingto their Figure 2. Mean surface temperature daisypopulations and for the ambienttemperature, whitedaisies black daisiesthrive in ambienttemperatures around theDaisyworld modelincorporating blackandwhitedaisies coloration; both over a range of solar luminosity(adaptedfrom Watsonand 15 to 25C and warm the surface, while white daisies thrive in Lovelock [1983]). Watson and Lovelock do not show the ambienttemperatures around20 to 30C and cool the surface. hysteresis loopsthat arisewhensolarluminosity decreases; they As a result,over a wide rangeof solarluminositythe surfaceis at favorablefor oneor bothdaisies. are shownhereto illustratethe existence multipleequilibriain maintained temperatures of somerangesof luminosity. Black daisieswill be warmer than

availabledata indicatethat it doesso in the way that the pathological Daisyworld (instead of Lovelock and Watson's Daisyworld) saysit should. Do models such as Daisyworld constitute[Lovelock, 1983,p. 66] "a cybemetic proofof theGaia hypothesis"?
Of course, clever modelers can make a model do almost

theory(in words)with itself (in mathematical terms). Such models can be usedto deriveand illuminatethe implications of the Gaia hypothesis, but not to establishits validity.

whatever they want,but the pointI will try to makehereis slightlymore subtle. Modelsare oftenusedto derivethe consequences hypotheses. of When the model'scomponentscan be independently verified or have been derived from well-groundedmechanistictheory (as in climate modeling), the results can be particularly useful in comparing theoryand observations. However,the results of heuristicmodelslike Daisyworldcannotbe directly tested against real world dam. Consequently, theoryis the tested,not by comparing model and real world behavior, but by comparing modelresultsto the behaviorpredicted by the Gaia hypothesis.That kind of testwill inevitably succeed (barringlogicalor mathematical errors)because it is not comparinga theory with data, but comparinga

GEOPHYSIOLOGICAL

GAlA

Lovelock [1986b, 1988] has recently proposed "geophysiology," representation the biosphere a the of as singleorganism,as a systems approach Earth science. to In thisview thebiosphere, an organism, generally like will maintaininternalhomeostasis the face of changing in external forces. Justas physiology addresses functionthe ing of whole organisms,geophysiology intended to is unify the constituentparts of the biosphere and the disciplinesthat addressthem. According to Lovelock [1988, preface], "specialties, like biogeochemistry, theoretical ecology,andevolutionary biology,all exist,but they have no more to offer the concerned environmental

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80

Kirchner: THE GAIA HYPOTHESIS o 231

60

, 4o- 20

o
0.5
80

1.0

1.5

0
0.5

'

,
1.0

....

--
1.5

, 40

0.5

1.0

1.5

Solar luminosity (arbitrary units)

that Figure 3. Daisyworld exhibiting "pathological" behavior a equilibria,both outsidethe rangesof ambienttemperatures as result fixingtheoptimal of temperature blackdaisies 15C are favorablefor either daisy color (20-30C for white, and of at one of the daisy above thatof whitedaisies.Doingsoreverses biasimplicitin 30-40C for black). At either equilibrium, the

Watsonand Lovelock's[1983] assumption black daisiesare that


10C "blacker" than white daisies. Note the existence of two

species drivento extinction. is

physician the patientthancouldthe analogous or science of biochemistry or microbiology in the nineteenth century." Some systemslevel approachto Earth scienceis desperately needed, should be itsbasis?It maybe but this misleadingto use the same languagewe apply to organismswhen discussing biosphere a planetary the as organism.To borrowa pointfrom evolutionary biologist E. Mayr, theremay be no suchthingas a healthyherdof deer,but only a herd of healthydeer. Now, a herd (or a planet)maybe saidto be healthy, not in the samesense but

naturalworld as if it were an organism.But the question of whetherthe Earth actually is an organism neither is scientifically meaningful scientifically nor answerable. As a hypothesis, biosphere/organism the metaphoris, like all metaphors, ill-defined. In whichsenses theEarth is like an organism, and how can we tell how far to take the analogy?The vastdiversity living organisms, of from men to microbesand plants to pachyderms, meansthat any conceivable phenomenon could be comparedto some behavior some of organism.For example, geophysiology explains sudden the redoxshiftat the endof theArcbean, a thatoneof its members healthy. is dramatic exampleof biologicaldestabilization, termsof in It is important to distinguishgeophysiological Gaia puberty [Lovelock, 1988,p. 99]. Similarly,geophysiology from an Earth-as-organism metaphor. Consider, for reinterprets ice core data (which suggest, described the as feedback reinforces temperature the comparison, the Earth-as-mechanism metaphor that above,that biological rather than underlies contemporary naturalscience. The success of shiftsbetweenglacial ages and interglacials, this mechanistic framework has demonstrated the effechomeostatically opposingthem) in the following way 1988,pp. 136 and 150]: "the glacialcool is the tiveness thinkingaboutthe naturalworld as if it werea [Lovelock, of machine(whetheror not it actually is, in any sense,a preferredstate of Gaia, [and] the interglacials like the one sometemporary failureof regulation, mechanism).I would agreewith the proponents Gaia present represent of that it may be useful to attemptto speculate about the a feveredstateof the planet... [duringglacials]cloud

232 Kirchner: THE GAIA HYPOTHESIS

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cover and low carbondioxide operatedin synchrony as

partof a geophysiological process keeptheEarthcool." to Geophysiology be endlessly can reinterpreted, for and that reasonit is unfalsifiable. Any conceivable behavior could be taken to be physiological, particularlyif one defines physiologically the "preferred"state will andif at one considers physiological breakdown (e.g., "feverish" interglacials) be evidence physiological to for behavior. Geophysiology predicts [Lovelock, 1988,p. 13] "homeostasis longperiods for untilsome internal contradiction or externalforce causes jump to a new stablestate." But a

there is some "objective function" that thoseconditions optimize. Every conceivable environment can be "favorable," if one hascomplete freedomto specifywhat
it is favorable for.

What could possiblybe optimal for the whole biosphere? One can define an optimal environment a for givenorganism a number ways. But whatconditions in of are optimal, or even simply favorable, for a diverse collection organisms haveconflicting of that requirements? One response that currentconditions optimal,but is are thiscreates obvious an tautology definingoptimalityin by external forces and internal contradictionsare always terms of prevalent conditionsand then assertingthat prevalent; instability any could (perhaps erroneously, since prevalent conditions (assumed to be maintained by control) are,in fact,optimal. a balanceof "internal contradictions" be stabilizing) biological can Stability and optimality(for the agent supplyingthe be attributed any numberof suchcauses. Moreover, to are of giventhat stasis defined the absence jumps,and homeostasis) mutuallyexclusive.The population an is as of that makes its environment more suitablefor jumpsaredefined theabsence stasis, as of geophysiologyorganism seems predictsimplythat the biosphere to will exhibit itself will grow, whereuponit dominatesmore of the homeostasis exceptwhen it doesnot. It is futile to try to environmentand grows still further; this behavior is Stabilityis achieved organisms by "fouling falsify sucha hypothesis, to try to derivemeaningful destabilizing. or their own nests" and thereby limiting their population predictions fromit. To some,an endlessly interpretable theoryis appealing growth. Consider the Daisyworldmodel in a world with only because can be usedto explainvirtuallyanything. But it pointthe daisies supplyhomeosfor thesame reason predicts it nothing, itsexplanations whitedaisies.At a stable and rises, more daisiesgrow are empty. If any conceivable phenomena be ex- tasisbecauseif the temperature can plained geophysiology, geophysiology simply and the albedo increases. This implies that a higher by then says would be more suitablefor white daisies,so that "anythingis possible." The value of a scientific temperature the temperature the stablepoint is coolerthan optimal, at theory, contrast, precisely limitingthesphere by lies in of preventsa warmingthat thepossible, thereby allowing to separate plausible and the daisies'growthresponse one the would be favorablefor them. At the peak of the daisy from the implausible. if If the purpose geophysiology of were simply to spur growthcurve,thereis no homeostasis; the temperature increases, daisiesdie, and the temperature increases still interesting speculations aboutthe climatichistoryof the further, and more daisiesdie. The optimum is reached Earth,onemightnot object. ButLovelock [1986b,p. 12] transition between stablesuboptimum a (see also Lovelock [1988, pp. 152-182]) expressly onlyin an unstable advocates geophysiology "the essential as theoretical basis and total extinction. A commonobjection Gaian optimization that it is to is for the putative profession planetary of medicine" cure to it and Earth's environmental ills. With no meansof testingthe teleological; would requiresomesort of foresight [Doolittle,1981]. It is important notethatone to underlying theory, how can onetell whether curesit planning the recommends theplanetary are equivalent penicillin of can form a mechanistic scenario for the evolution of of or without any "engineering,"just as bloodletting? The risk of creating iatrogenicglobal favorableconditions antsand termites haveevolvedstrategies regulating for the maladies throughtreatments that have no testable basis should not be underestimated. It is hard to see how temperature theirnests of withoutbeingtold how to do it: geophysiology could help us to distinguish between "planetary medicine" planetary and malpractice. Lovelock Margulis [1974a,p. 99] and
Although the environmentalcontrol mechanisms are likely to be subtle and complex, we believe their evolutioncan be comprehended broadly in terms of Neodarwinianthought .... Analogouswith the
evolution of local environmentalor internal control, in

OPTIMIZING

GAlA

The most speculative versionof the Gaia hypothesis holds that biotic control of the physical environment creates conditions that are favorable,or even optimal,for the biosphere.This hypothesis ill-definedunless is one can specifywhat constitutes favorableor optimalconditions. It canbe demonstrated for any setof conditions, that

the evolutionof atmospheric homeostasis thosespecies of organisms retainor alter conditions that optimizing their timess (i.e., proportionof offspring left to the subsequent generation) leavemoreof the same. In this way conditions retained alteredto theirbenefit. are or

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Kirchner: THEG^I^HYPOTHESlS

o233

favorable penguins, was met with the response for I that Arctic animalsare eccentric, exotic,fringe species.That, It is a recognized fact of biologythat environments are of course,is exactly the point; on a much colder Earth, regulated a much more local scale... We merely on penguins (and latter-daymastodons, perhaps)would be suggestextrapolationof these ideas to the entire moreprominent, scientists and wouldpeerout fromtheir atmosphere-biosphere system. fur coatsto observe that the environment seemed ideally suited to their needs. Lovelock[ 1986b,p. 13]

MargulisandLovelock[1974,p. 486]

Life and the environment evolvetogether a single as


system that not only doesthe species leavesthe so that most progenytend to inherit the environment but also
CONCLUSIONS

the environment favorsthe mostprogeny itself that is


sustained.

The proponents the Gaia hypothesis of havemadethree contributions. First,theyhavereiterated point the The problem with these arguments not that theyare valuable is originally made by those investigating biogeochemical demonstrably wrong,but that they are radicallyunparplay a vital role in simonious and, given ordinaryDarwinianremovalof the cycles,namelythat biologicalagents unfit,theyareunfalsifiable. global For control arise to by creatingthe Earth's physicaland chemicalenvironment in chemistry and evolution, an organism's effects on "the entire and that thoseinterested the surface of arbitrarily ignorebiological atmosphere-biosphere system"wouldneedto be signifi- physics the Earthcannot Second, theyhavesuggested somemechanisms cantenough affectits ownreproductive to success. This is processes. highlyimplausible, Doolittle [1981] haspointedout: by which the biota may be particularlyimportantin as determining Earth'sclimate. Third, theyhavecreated the "It is difficult to acceptthat behaviors whoseeffects... and that somehaveusedas will notbe felt for thousands generations be selected an engaging colorfulmetaphor of can for for, especially when the first beneficiaries thoseeffects a unifyingthemeand a motivator research.In my of view, however, this metaphor(if taken literally) is may be organisms which are not themselves responsible color one's for them." Recently, Lovelock[1988, pp. 126-151] has ultimately misleadingand will needlessly of environment emphasized Gaianmechanisms arisingthrough behaviors impressions howthebiotaandthephysical as thatbenefit organisms directly, otherthanGaianways. interact. Attemptsto test this metaphor a scientific in proposition be, in my opinion, will ultimately futile. But if the connection betweenindividualreproductive Some have viewed Gaia as a dating but viable advantage and Gaian altruismis only coincidental, we would expectGaian and non-Gaianbehaviorto arise with hypothesis. My view is that each of the different Gaia is equal probability. Darwin said it all much more hypotheses either daring or viable, but not both. Gaia but it straightforwardly: environment the biotaare well Coevolutionary is testable not daring; simply the and recasts, different in language, long-standing widely the and matched because organisms arepoorly that adapteA be will accepted observation that biologicalprocesses and the underrepresented nextgeneration's pool. in the gene Given that the unfit are selected against, claimsthat physicalenvironmentinfluence one another. The claim Gaia creates a favorable environment are unfalsifiable. thatthese interrelationships generally, evenuniverare or (homeostatic Gaia) may be testable and Whether notGaia optimizes, wouldstill observe or we that sally,stabilizing may well spursome interesting researchinto thefeedback the Earth's organismsmatch its environment, because controlling climate.The scanty dataavailable organisms that do not suit the environment not thrive mechanisms do itself. andwill notbenoticed. Holland[1984, 539]putit quite to date, however,tend to refute the hypothesis p. Geophysiological by contrast, Gaia, cannot refuted be by simply: data,notbecause is inescapably it correct, because is but it The geologicrecordseemsmuchmore in accordwith ill-defined canbe reinterpreted explain and to almost any the view that the organisms that are better able to imaginable phenomena.OptimizingGaia is similarly competehave come to dominate,and that the Earth's ill-defined, andgivensimpleDarwinianelimination the of near-surface environments processes and have accom- unfit,it is bothunparsimonious unfalsifiable. and modatedthemselves the changeswrought by to Gaiais crippled its greatgenerality; searches a by it for biologicalevolution... We live on an Earth thatis the description therole of life on Earth. The of bestof all possible worldsonly for thosewho are well simplecapsule key fact, that the biota and the physicalworlds are adapted its currentstate. to interconnected, realizedlong ago. The day-to-day was When,at the recentAGU Chapman Conference the task of figuring out "how the world works" consists on of Gaiahypothesis, I raised objection Gaia,arguing an to that documentingthese interconnections and their consethe Earth's physicalenvironment was obviouslynot quences. Thereis a lot of difficult,important science be to

234 Kirchner'

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done here, and we shouldget on with it. An untestable belief that theserelationships oughtto have somegeneral form or functionmay only blind us to what they actually are and actually do. What is "out there," what can be observed, measured, hypothesized, verified, and falsified, is biogeochemistry, natural selection,evolution,and so forth. Gaia may be a grandvision,but it is not the kind of visionthatcanbe scientifically validated. Some may objectthat to abandon empiricalsearch the for Gaia is to forgo the chanceof discovering grand, a ultimate truth. With Poincare [1905, p. 163] I would respond,
To those who feel that we are going too far in our
limitations of the domain accessible to the scientist, I

Hansen,J., A. Lacis, D. Rind, G. Russell,P. Stone,I. Fung, R. Ruedy, and J. Lerner, Climate sensitivity: Analysis of
feedback mechanisms, in Climate Processes and Climate

Sensitivity, Geophys. Monogr. Ser., vol. 29, editedby J. E. Hansenand T. Takahashi,pp. 130-163, AGU, Washington,
D.C., 1984.

Harvey,H. W., The Chemistry Fertility of Sea Waters,2rid and ed.,240 pp.,Cambridge University Press, New York, 1957.
Henderson,L. J., The Fitness of the Enviromnent,Macmillan, New York, 1913. (Reprinted BeaconPress, by Boston, Mass., 1958.) Hitchcock, D. R., and J. E. Lovelock, Life detection by atmospheric analysis, Icarus,7, 149, 1967.
Holland, H. D., On the chemical evolution of the terrestrial and

reply: These questionswhich we forbid you to investigate,and which you so regret, are not only insoluble, theyare illusoryanddevoidof meaning.
ACKNOWLEDGMENTS. J. Harte first suggestedthat a taxonomyof the Gaia hypotheses would be useful and made many vital comments.I am gratefulto J. E. Lovelock,H. D. Holland, S. H. Schneider, E. Power, and B. A. Roy for their M. suggestions, the anonymous to reviewers theircomments, for and
to the William and Flora Hewlett Foundation for financial

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Hutchinson, G. E., The biogeochemistryof the terrestrial atmosphere, The Earth as a Planet,edited G. P. Kuiper, in by pp.371-433, University Chicago of Press, Chicago, 1954. Ill., Huxley, T. H., Physiography, Introduction the Studyof An to Nature,384 pp.,MacMillan, London,1877. Jouzel, J., C. Lorius, J. Petit, C. Genthon, N. Barkov, V. Kotlyakov,and V. Petrov,Vostok ice core: A continuous isotope temperaturerecord over the last climatic cycle (160,000years), Nature,329, 403-408, 1987. Lashof,D., The dynamicgreenhouse, Clim. Change,14, in press,

1989. Ann Henderson-Sellers was the editor responsible for this paper. Shethanks G. CogleyandP. Boston theirassistance Legrand,M. R., R. J. Delmas, and R. J. Charlson,Climate J. for forcing implicationsfrom vostok ice-core sulphate data, in evaluating paper. this

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James W. Kirchner, Energyand Resources Group, Universityof California, Berkeley, CA 94720.

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