10/05/2017 EntropyandlifeWikipedia

Entropyandlife
FromWikipedia,thefreeencyclopedia

Researchconcerningtherelationshipbetweenthethermodynamicquantityentropyandthe
evolutionoflifebeganaroundtheturnofthe20thcentury.In1910,AmericanhistorianHenry
AdamsprintedanddistributedtouniversitylibrariesandhistoryprofessorsthesmallvolumeA
LettertoAmericanTeachersofHistoryproposingatheoryofhistorybasedonthesecondlaw
ofthermodynamicsandontheprincipleofentropy.[1][2]The1944bookWhatisLife?byNobel
laureatephysicistErwinSchrdingerstimulatedresearchinthefield.Inhisbook,Schrdinger
originallystatedthatlifefeedsonnegativeentropy,ornegentropyasitissometimescalled,but
inalatereditioncorrectedhimselfinresponsetocomplaintsandstatedthetruesourceisfree
energy.MorerecentworkhasrestrictedthediscussiontoGibbsfreeenergybecausebiological
processesonEarthnormallyoccurataconstanttemperatureandpressure,suchasinthe
atmosphereoratthebottomofanocean,butnotacrossbothovershortperiodsoftimefor
individualorganisms.

Contents
1 Origin
2 Earlyviews
3 Negativeentropy
4 Gibbsfreeenergyandbiologicalevolution
5 Entropyandtheoriginoflife
6 EntropyandthesearchforlifeelsewhereintheUniverse
7 Otherterms
8 Objections
9 Seealso
10 References
11 Furtherreading
12 Externallinks

Origin
In1863,RudolfClausiuspublishedhisnotedmemoir"OntheConcentrationofRaysofHeat
andLight,andontheLimitsofitsAction"whereinheoutlinedapreliminaryrelationship,as
basedonhisownworkandthatofWilliamThomson,betweenhisnewlydevelopedconceptof
entropyandlife.Buildingonthis,oneofthefirsttospeculateonapossiblethermodynamic
perspectiveofevolutionwastheAustrianphysicistLudwigBoltzmann.In1875,buildingonthe
worksofClausiusandKelvin,Boltzmannreasoned:

Thegeneralstruggleforexistenceofanimatebeingsisnotastruggleforraw
materialsthese,fororganisms,areair,waterandsoil,allabundantlyavailable
norforenergywhichexistsinplentyinanybodyintheformofheat,butastruggle
for[negative]entropy,whichbecomesavailablethroughthetransitionofenergyfrom
thehotsuntothecoldearth.[3]

https://en.wikipedia.org/wiki/Entropy_and_life 1/8
10/05/2017 EntropyandlifeWikipedia

Earlyviews
In1876,AmericancivilengineerRichardSearsMcCulloh,inhisTreatiseontheMechanical
TheoryofHeatanditsApplicationtotheSteamEngine,whichwasanearlythermodynamics
textbook,states,afterspeakingaboutthelawsofthephysicalworld,that"therearenonethat
areestablishedonafirmerbasisthanthetwogeneralpropositionsofJouleandCarnotwhich
constitutethefundamentallawsofoursubject."McCullochthengoesontoshowthatthesetwo
lawsmaybecombinedinasingleexpressionasfollows:

where

=entropy
=adifferentialamountofheatpassedintoathermodynamicsystem
=absolutetemperature

McCullochthendeclaresthattheapplicationsofthesetwolaws,i.e.whatarecurrentlyknown
asthefirstlawofthermodynamicsandthesecondlawofthermodynamics,areinnumerable.
Hethenstates:

Whenwereflecthowgenerallyphysicalphenomenaareconnectedwiththermal
changesandrelations,itatoncebecomesobviousthattherearefew,ifany,
branchesofnaturalsciencewhicharenotmoreorlessdependentuponthegreat
truthsunderconsideration.Norshouldit,therefore,beamatterofsurprisethat
already,intheshortspaceoftime,notyetonegeneration,elapsedsincethe
mechanicaltheoryofheathasbeenfreelyadopted,wholebranchesofphysical
sciencehavebeenrevolutionizedbyit.[4]:p.267

McCullochthengivesafewofwhathecallsthemoreinterestingexamplesoftheapplication
oftheselawsinextentandutility.Thefirstexamplehegivesisphysiology,whereinhestates
thatthebodyofananimal,notlessthanasteamer,oralocomotive,istrulyaheatengine,and
theconsumptionoffoodintheoneispreciselyanalogoustotheburningoffuelintheotherin
both,thechemicalprocessisthesame:thatcalledcombustion.Hethenincorporatesa
discussionofLavoisierstheoryofrespirationwithcyclesofdigestionandexcretion,
perspiration,butthencontradictsLavoisierwithrecentfindings,suchasinternalheat
generatedbyfriction,accordingtothenewtheoryofheat,which,accordingtoMcCulloch,
statesthattheheatofthebodygenerallyanduniformlyisdiffusedinsteadofbeing
concentratedinthechest.McCullochthengivesanexampleofthesecondlaw,wherehe
statesthatfriction,especiallyinthesmallerbloodedvessels,mustdevelopheat.Without
doubt,animalheatisthusinpartproduced.Hethenasks:butwhencetheexpenditureof
energycausingthatfriction,andwhichmustbeitselfaccountedfor?"

Toanswerthisquestionheturnstothemechanicaltheoryofheatandgoesontoloosely
outlinehowtheheartiswhathecallsaforcepump,whichreceivesbloodandsendsitto
everypartofthebody,asdiscoveredbyWilliamHarvey,thatactslikethepistonofanengine
andisdependentuponandconsequentlyduetothecycleofnutritionandexcretionwhich
https://en.wikipedia.org/wiki/Entropy_and_life 2/8
10/05/2017 EntropyandlifeWikipedia

sustainsphysicalororganiclife.Itislikely,here,thatMcCullochwasmodelingpartsofthis
argumentonthatofthefamousCarnotcycle.Inconclusion,hesummarizeshisfirstand
secondlawargumentassuch:

Everythingphysicalbeingsubjecttothelawofconservationofenergy,itfollowsthat
nophysiologicalactioncantakeplaceexceptwithexpenditureofenergyderived
fromfoodalso,thatananimalperformingmechanicalworkmustfromthesame
quantityoffoodgeneratelessheatthanoneabstainingfromexertion,thedifference
beingpreciselytheheatequivalentofthatofwork.[4]:p.270

Negativeentropy
Later,buildingonthispremise,inthefamous1944bookWhatisLife?,Nobellaureatephysicist
ErwinSchrdingertheorizesthatlife,contrarytothegeneraltendencydictatedbytheSecond
lawofthermodynamics,decreasesormaintainsitsentropybyfeedingonnegativeentropy.[5]
InhisnotetoChapter6ofWhatisLife?,however,Schrdingerremarksonhisusageofthe
termnegativeentropy:

Letmesayfirst,thatifIhadbeencateringforthem[physicists]aloneIshouldhave
letthediscussionturnonfreeenergyinstead.Itisthemorefamiliarnotioninthis
context.Butthishighlytechnicaltermseemedlinguisticallytooneartoenergyfor
makingtheaveragereaderalivetothecontrastbetweenthetwothings.

Thisiswhatisarguedtodifferentiatelifefromotherformsofmatterorganization.Inthis
direction,althoughlife'sdynamicsmaybearguedtogoagainstthetendencyofsecondlaw,
whichstatesthattheentropyofanisolatedsystemtendstoincrease,itdoesnotinanyway
conflictorinvalidatethislaw,becausetheprinciplethatentropycanonlyincreaseorremain
constantappliesonlytoaclosedsystemwhichisadiabaticallyisolated,meaningnoheatcan
enterorleave.Wheneverasystemcanexchangeeitherheatormatterwithitsenvironment,an
entropydecreaseofthatsystemisentirelycompatiblewiththesecondlaw.[6]Theproblemof
organizationinlivingsystemsincreasingdespitethesecondlawisknownastheSchrdinger
paradox.[7]

In1964,JamesLovelockwasamongagroupofscientistswhowererequestedbyNASAto
makeatheoreticallifedetectionsystemtolookforlifeonMarsduringtheupcomingspace
mission.Whenthinkingaboutthisproblem,Lovelockwonderedhowcanwebesurethat
Martianlife,ifany,willrevealitselftotestsbasedonEarthslifestyle?[8]ToLovelock,thebasic
questionwasWhatislife,andhowshoulditberecognized?Whenspeakingaboutthisissue
withsomeofhiscolleaguesattheJetPropulsionLaboratory,hewasaskedwhathewoulddo
tolookforlifeonMars.Tothis,Lovelockreplied:

Idlookforanentropyreduction,sincethismustbeageneralcharacteristicoflife.[8]

Thus,accordingtoLovelock,tofindsignsoflife,onemustlookforareductionorareversalof
entropy.

https://en.wikipedia.org/wiki/Entropy_and_life 3/8
10/05/2017 EntropyandlifeWikipedia

Gibbsfreeenergyandbiologicalevolution
Inrecentyears,thethermodynamicinterpretationofevolutioninrelationtoentropyhasbegun
toutilizetheconceptoftheGibbsfreeenergy,ratherthanentropy.[9]Thisisbecausebiological
processesonearthtakeplaceatroughlyconstanttemperatureandpressure,asituationin
whichtheGibbsfreeenergyisanespeciallyusefulwaytoexpressthesecondlawof
thermodynamics.TheGibbsfreeenergyisgivenby:

TheminimizationoftheGibbsfreeenergyisaformoftheprincipleofminimumenergy,which
followsfromtheentropymaximizationprincipleforclosedsystems.Moreover,theGibbsfree
energyequation,inmodifiedform,canbeutilizedforopensystemswhenchemicalpotential
termsareincludedintheenergybalanceequation.Inapopular1982textbook,Principlesof
BiochemistrybynotedAmericanbiochemistAlbertLehninger,itisarguedthattheorder
producedwithincellsastheygrowanddivideismorethancompensatedforbythedisorder
theycreateintheirsurroundingsinthecourseofgrowthanddivision.Inshort,accordingto
Lehninger,"livingorganismspreservetheirinternalorderbytakingfromtheirsurroundingsfree
energy,intheformofnutrientsorsunlight,andreturningtotheirsurroundingsanequalamount
ofenergyasheatandentropy."[10]

Similarly,accordingtothechemistJohnAvery,fromhisrecent2003bookInformationTheory
andEvolution,wefindapresentationinwhichthephenomenonoflife,includingitsoriginand
evolution,aswellashumanculturalevolution,hasitsbasisinthebackgroundof
thermodynamics,statisticalmechanics,andinformationtheory.The(apparent)paradox
betweenthesecondlawofthermodynamicsandthehighdegreeoforderandcomplexity
producedbylivingsystems,accordingtoAvery,hasitsresolution"intheinformationcontentof
theGibbsfreeenergythatentersthebiospherefromoutsidesources."[11]Theprocessof
naturalselectionresponsibleforsuchlocalincreaseinordermaybemathematicallyderived
directlyfromtheexpressionofthesecondlawequationforconnectednonequilibriumopen
systems.[12]

Entropyandtheoriginoflife
Thesecondlawofthermodynamicsappliedontheoriginoflifeisafarmorecomplicatedissue
thanthefurtherdevelopmentoflife,sincethereisno"standardmodel"ofhowthefirst
biologicallifeformsemergedonlyanumberofcompetinghypotheses.Theproblemis
discussedwithintheareaofabiogenesis,implyinggradualpreDarwinianchemicalevolution.
In1924,AlexanderOparinsuggestedthatsufficientenergywasprovidedinaprimordialsoup.
TheBelgianscientistIlyaPrigoginewasawardedwithaNobelprizein1977forananalysisin
thisarea.Arelatedtopicistheprobabilitythatlifewouldemerge,whichhasbeendiscussedin
severalstudies,forexamplebyRussellDoolittle.[13]

EntropyandthesearchforlifeelsewhereintheUniverse
In2013AzuaBustosandVegaarguedthatdisregardingthetypeoflifeformthatcouldbe
envisionedbothonEarthandelsewhereintheUniverse,allshouldshareincommonthe
attributeofbeingentitiesthatdecreasetheirinternalentropyattheexpenseoffreeenergy
obtainedfromitssurroundings.Asentropyallowsthequantificationofthedegreeofdisorderin
https://en.wikipedia.org/wiki/Entropy_and_life 4/8
10/05/2017 EntropyandlifeWikipedia

asystem,anyenvisionedlifeformmusthaveahigherdegreeoforderthanitssupporting
environment.Theseauthorsshowedthatbyusingfractalmathematicsanalysisalone,they
couldreadilyquantifythedegreeofstructuralcomplexitydifference(andthusentropy)ofliving
processesasdistinctentitiesseparatefromtheirsimilarabioticsurroundings.Thisapproach
mayallowthefuturedetectionofunknownformsoflifebothintheSolarSystemandon
recentlydiscoveredexoplanetsbasedonnothingmorethanentropydifferentialsof
complementarydatasets(morphology,coloration,temperature,pH,isotopiccomposition,etc.).
Detectinglifeaswedon'tknowitbyfractalanalysis(http://journals.cambridge.org/action/displ
ayAbstract?fromPage=online&aid=9012687&fileId=S1473550413000177)

Otherterms
Fornearlyacenturyandahalf,beginningwithClausius'1863memoir"OntheConcentration
ofRaysofHeatandLight,andontheLimitsofitsAction",muchwritingandresearchhasbeen
devotedtotherelationshipbetweenthermodynamicentropyandtheevolutionoflife.The
argumentthatlifefeedsonnegativeentropyornegentropywasassertedbyphysicistErwin
Schrdingerina1944bookWhatisLife?.Heposed,"Howdoesthelivingorganismavoid
decay?"Theobviousansweris:"Byeating,drinking,breathingand(inthecaseofplants)
assimilating."RecentwritingshaveusedtheconceptofGibbsfreeenergytoelaborateonthis
issue.[14]Whileenergyfromnutrientsisnecessarytosustainanorganism'sorder,thereisalso
theSchrdingerprescience:"Anorganism'sastonishinggiftofconcentratingastreamoforder
onitselfandthusescapingthedecayintoatomicchaosofdrinkingorderlinessfroma
suitableenvironmentseemstobeconnectedwiththepresenceoftheaperiodicsolids..."We
nowknowthatthe'aperiodic'crystalisDNAandthattheirregulararrangementisaformof
information."TheDNAinthecellnucleuscontainsthemastercopyofthesoftware,in
duplicate.Thissoftwareseemstocontrolby"specifyinganalgorithm,orsetofinstructions,for
creatingandmaintainingtheentireorganismcontainingthecell."[15]DNAandother
macromoleculesdetermineanorganism'slifecycle:birth,growth,maturity,decline,anddeath.
Nutritionisnecessarybutnotsufficienttoaccountforgrowthinsizeasgeneticsisthe
governingfactor.Atsomepoint,organismsnormallydeclineanddieevenwhileremainingin
environmentsthatcontainsufficientnutrientstosustainlife.Thecontrollingfactormustbe
internalandnotnutrientsorsunlightactingascausalexogenousvariables.Organismsinherit
theabilitytocreateuniqueandcomplexbiologicalstructuresitisunlikelyforthosecapabilities
tobereinventedorbetaughteachgeneration.Therefore,DNAmustbeoperativeastheprime
causeinthischaracteristicaswell.ApplyingBoltzmann'sperspectiveofthesecondlaw,the
changeofstatefromamoreprobable,lessorderedandhighentropyarrangementtooneof
lessprobability,moreorder,andlowerentropyseeninbiologicalorderingcallsforafunction
likethatknownofDNA.DNA'sapparentinformationprocessingfunctionprovidesaresolution
oftheparadoxposedbylifeandtheentropyrequirementofthesecondlaw.[16]

In1982,AmericanbiochemistAlbertLehningerarguedthatthe"order"producedwithincellsas
theygrowanddivideismorethancompensatedforbythe"disorder"theycreateintheir
surroundingsinthecourseofgrowthanddivision."Livingorganismspreservetheirinternal
orderbytakingfromtheirsurroundingsfreeenergy,intheformofnutrientsorsunlight,and
returningtotheirsurroundingsanequalamountofenergyasheatandentropy."[17]

Evolutionrelatedconcepts:

Negentropyashorthandcolloquialphrasefornegativeentropy.[18]

https://en.wikipedia.org/wiki/Entropy_and_life 5/8
10/05/2017 EntropyandlifeWikipedia

Ectropyameasureofthetendencyofadynamicalsystemtodousefulworkandgrow
moreorganized.[19]
Extropyametaphoricaltermdefiningtheextentofalivingororganizationalsystem's
intelligence,functionalorder,vitality,energy,life,experience,andcapacityanddrivefor
improvementandgrowth.
Ecologicalentropyameasureofbiodiversityinthestudyofbiologicalecology.

Inastudytitled"Naturalselectionforleastaction"publishedintheProceedingsofTheRoyal
SocietyA.,VilleKailaandArtoAnnilaoftheUniversityofHelsinkidescribehowthesecondlaw
ofthermodynamicscanbewrittenasanequationofmotiontodescribeevolution,showinghow
naturalselectionandtheprincipleofleastactioncanbeconnectedbyexpressingnatural
selectionintermsofchemicalthermodynamics.Inthisview,evolutionexplorespossiblepaths
toleveldifferencesinenergydensitiesandsoincreaseentropymostrapidly.Thus,an
organismservesasanenergytransfermechanism,andbeneficialmutationsallowsuccessive
organismstotransfermoreenergywithintheirenvironment.[20]

Objections
Sinceentropyisdefinedforequilibriumsystems,[21]objectionstotheextensionofthesecond
lawandentropytobiologicalsystems,especiallyasitpertainstoitsusetosupportordiscredit
thetheoryofevolution,havebeenstated.[22]Livesystemsandindeedmuchofthesystems
andprocessesintheuniverseoperatefarfromequilibrium,whereasthesecondlawsuccinctly
statesthatisolatedsystemsevolvetowardthermodynamicequilibriumthestateofmaximum
entropy.

Ontheotherhand,(1)livesystemscannotpersistinisolationand(2)thesecondprincipleof
thermodynamicsdoesnotrequirethatfreeenergybetransformedintoentropyalongthe
shortestpath:liveorganismsabsorbenergyfromsunlightorfromenergyrichchemical
compoundsandfinallyreturnpartofsuchenergytotheenvironmentasentropy(heatandlow
freeenergycompoundssuchaswaterandCO2).

Seealso
Abiogenesis
Complexsystems
Dissipativesystem
Entropy(orderanddisorder)
Adaptivesystem

References
1.Adams,Henry.(1986).HistoryoftheUnited
StatesofAmericaDuringtheAdministration
ofThomasJefferson(pg.1299).Libraryof
America.
2.Adams,Henry.(1910).ALettertoAmerican
TeachersofHistory.GoogleBooks(https://b
ooks.google.com/books?id=gaLdOOzuiKAC
&pg=PA1&dq=A+Letter+to+American+Teach
ers+of+History#PPA10,M1),ScannedPDF(h
ttps://archive.org/details/alettertoamerica00a
damuoft).Washington.

https://en.wikipedia.org/wiki/Entropy_and_life 6/8
10/05/2017 EntropyandlifeWikipedia

3.Boltzmann,Ludwig(1974).Thesecondlaw
ofthermodynamics(Theoreticalphysicsand
philosophicalproblems).SpringerVerlag
NewYork,LLC.ISBN9789027702500.
4.McCulloch,RichardSears(1876).Treatise
onthemechanicaltheoryofheatandits
applicationstothesteamengine,etc.New
York:D.VanNostrand.
5.Schrdinger,Erwin(1944).WhatisLifethe
PhysicalAspectoftheLivingCell.
CambridgeUniversityPress.ISBN0521
427088.
6.Thecommonjustificationforthisargument,
forexample,accordingtorenowned
chemicalengineerKennethDenbigh,from
his1955bookThePrinciplesofChemical
Equilibrium,isthat"livingorganismsare
opentotheirenvironmentandcanbuildup
attheexpenseoffoodstuffswhichtheytake
inanddegrade."
7.Schneider,EricD.Sagan,Dorion(2005).
IntotheCool:EnergyFlowThermodynamics
andLife.Chicago,UnitedStates:The
UniversityofChicagoPress.p.15.
8.Lovelock,James(1979).GAIAANew
LookatLifeonEarth.OxfordUniversity
Press.ISBN0192862189.
9.Moroz,Adam(2012).TheCommon
ExtremalitiesinBiologyandPhysics.
Elsevier.ISBN9780123851871.
10.Lehninger,Albert(1993).Principlesof
Biochemistry,2ndEd.WorthPublishers.
ISBN0879017112.
11.Avery,John(2003).InformationTheoryand
Evolution.WorldScientific.ISBN981238
3999.
12.Kaila,V.R.Annila,A.(8November2008).
"Naturalselectionforleastaction".
ProceedingsoftheRoyalSocietyA.464
(2099):30553070.
Bibcode:2008RSPSA.464.3055K.
doi:10.1098/rspa.2008.0178.
13.RussellDoolittle,"TheProbabilityandOrigin
ofLife"inScientistsConfrontCreationism
(1984)Ed.LaurieR.Godfrey,p.85
14.Higgs,P.G.,&Pudritz,R.E.(2009)."A
thermodynamicbasisforprebioticamino
acidsynthesisandthenatureofthefirst
geneticcode"Acceptedforpublicationin
Astrobiology(http://adsabs.harvard.edu/cgi
bin/bib_query?arXiv:0904.0402)
15.Nelson,P.(2004).BiologicalPhysics,
Energy,Information,Life.W.H.Freemanand
Company.ISBN0716743728
16.Peterson,Jacob,Understandingthe
ThermodynamicsofBiologicalOrder,The
AmericanBiologyTeacher,74,Number1,
January2012,pp.2224
17.Lehninger,Albert(1993).Principlesof
Biochemistry,2ndEd.WorthPublishers.
ISBN0879017112.
18.Schrdinger,Erwin(1944).WhatisLifethe
PhysicalAspectoftheLivingCell.
CambridgeUniversityPress.ISBN0521
427088.
19.Haddad,WassimM.Chellaboina,
VijaySekharNersesov,SergeyG.(2005).
ThermodynamicsADynamicalSystems
Approach.PrincetonUniversityPress.
ISBN0691123276.
20.LisaZyga(11August2008)."Evolutionas
DescribedbytheSecondLawof
Thermodynamics".Physorg.com.Retrieved
20080814.
21.Callen,HerbertB(1985).Thermodynamics
andanIntroductiontoStatistical
Thermodynamics.JohnWileyandSons.
22.BenNaim,Arieh(2012).Entropyandthe
SecondLaw.WorldScientificPublishing.

Furtherreading
Schneider,E.andSagan,D.(2005).IntotheCool:EnergyFlow,Thermodynamics,and
Life(http://www.press.uchicago.edu/ucp/books/book/chicago/I/bo3533936.html).
UniversityofChicagoPress,Chicago.ISBN9780226739366
LaCerra,P.(2003)."TheFirstLawofPsychologyistheSecondLawofThermodynamics:
TheEnergeticEvolutionaryModeloftheMindandtheGenerationofHuman
PsychologicalPhenomena".HumanNatureReview,Volume3:440447.Fulltext(http://h
umannature.com/nibbs/03/lacerra.pdf).
Moroz,A.(2011).TheCommonExtremalitiesinBiologyandPhysics(http://www.scienced
irect.com/science/book/9780123851871).ElsevierInsights,NY.ISBN9780123851871

https://en.wikipedia.org/wiki/Entropy_and_life 7/8
10/05/2017 EntropyandlifeWikipedia

Externallinks
ThermodynamicEvolutionoftheUniverse(https://web.archive.org/web/20070612172500/
http://pi.physik.unibonn.de/~cristinz/thesis/t/node7.html)pi.physik.unibonn.de/~cristinz

Retrievedfrom"https://en.wikipedia.org/w/index.php?title=Entropy_and_life&oldid=773937098"

Categories: Thermodynamicentropy Biologicalevolution Biophysics

Thispagewaslasteditedon5April2017,at09:18.
TextisavailableundertheCreativeCommonsAttributionShareAlikeLicenseadditional
termsmayapply.Byusingthissite,youagreetotheTermsofUseandPrivacyPolicy.
WikipediaisaregisteredtrademarkoftheWikimediaFoundation,Inc.,anonprofit
organization.

https://en.wikipedia.org/wiki/Entropy_and_life 8/8