A Lei Da Evolução Como Um Princípio Máximo - Alfred Lotka

THE LAW OF EVOLUTION AS A MAXIMAL PRINCIPLE
Author(s): ALFRED J. LOTKA

Source: Human Biology, Vol. 17, No. 3 (SEPTEMBER, 1945), pp. 167-194
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HUMAN BIDLDfiy
a record of research
SEPTEMBER , 13)4 5
VOL. 17 No. 3
THE LAW OF EVOLUTION AS A

MAXIMAL PRINCIPLE
BY ALFRED J. LOTKA
| NDOUBTEDLY fundamentalto the conceptof evolution

is the idea thatit is in some significant
sense a directedpro-
cess. As soon,however,as we seek to definein preciseterms
■"=-
____ its presumablycharacteristic directionwe meetwithdifficul-
ties. To say,as somehave said or implied,thatthe directionof evolution
is thedirectionof progress,is merelyto substituteforan undefinedterm
another at best ill defined,1and contaminatedwith anthropomorphic
flavor. Such phrases as "passage fromlower to higherforms"which
are sometimesused to describethe directionof evolution,are open to
the same objection. If, on the other hand, it is stated that evolution
proceedsfromsimplerto more complex forms,fromless specializedto
more specializedforms,the directionof evolutionis but poorlydefined,
for the rule is one withdisturbingexceptions,and what we seek is not
an empiricalrule,but a law of naturethatbrooksno exceptions.
One descriptionof the directionof evolutionwhich has a certain
superficialappearance of greater scientificprecisionis that evolution
1 B. PetronievicsScience distinguishes
( Progress,1919, p. 407) expressly
progressive,
regressive,and mixedevolution. I citethis,notin evidence for,or
refutation
of,progress as a characteristic
of evolution,
butmerely to showthatthe
term"progress"is itselftoo indefinite
to be usedto characterize
thedirection of
evolution.

168 HUMAN BIOLOGY
proceeds from less probable to more probable states. That this ap-
pearanceof precisionis fallaciouscan be seen froma simpleexample.
In a new pack of cardsthefirstthirteen cardsare theAce to Ten, Knave,
of
Queen, King, Spades, arranged in that order. If we shuffle
the cards
thoroughly, we shall presentlyfind the suit of spades distributedap-
proximatelyevenlythroughout the pack. We can expressthisby saying
thatin the originalarrangement the spades had a densityof one, or 100
percent, in the firstquarterof the pack, and a densityzero, in the rest
of the pack, whereasat the end of the shuffling theyapproachedmuch
more nearlya uniformdensityof 13/52 or 25 percentthroughoutthe
whole pack. In an actual trial the followingdistribution was observed
after thoroughshuffling:
Spades
Number Density
(percent)
Firstquarter
of pack 3 23.1
Secondquarter 3 23.1
Thirdquarter 2 15.4
Fourthquarter 5 38.4
In a certainsense it can here be said thatthe pack of cards, in the

shuffling process,has passed froma less probableto a more probable
state. But this is not an objectivepropertyof eitherthe pack or the
process of shuffling.It is a subjectivepropertyof the shuffler, or the
observer,who takes a special interestin the suit of spades picked out
for observation,and who uses two fundamentally differentmodes of
description for the initialand the finalstate of the pack: In describing
the initial state he tells us specificallywhat the first13 cards are; he
describesthe finalstate only in termsof a density. There is only one
specification that correctlydescribesthe arrangementof the new pack;
there are a great many differentarrangementsthat would have the
particulardensityobserved in the shuffledpack. If the final state is
definedwith the same detail as the initial state,namelyby indicating
each card and its positionin the pack, then the finalstate is just as
probableor improbableas the initialstate. In the shuffling of the cards
the passage is not froma less probableto a more probablestate,but
froma preciselyknownto a less well knownstate. The distinctionis
not objectivebut psychological.The futureis here knownonly statis-

THE LAW OF EVOLUTION 169
ticallybecause we deliberatelyshufflein such manneras to have only

statisticalknowledgeof the course of events.
Evidently,then, the statementthat evolutionproceeds from less
probableto more probablestates is, not so much a false statement,as
it is a statementdevoid of meaning,unless it is furthersupplemented
by a specification as to how and in what respectsthe probabilitiesof
thesestatesare measured.2
The example of the card shuffling is also suited to illustratean-
otherpoint. It has been remarkedby various writers3(includingthe
present),4that evolution is an irreversibleprocess: Now the term
irreversibleprocesshas acquireda veryspecialmeaningin thevocabulary
of thephysicist,and noteveryprocessthatmightbe describedin ordinary
speech as irreversiblefalls into the categoryof irreversibleprocesses
as the physicistunderstandsthe term. The sufflingof the pack of
cards is an irreversibleprocessin the sense thatneverby mereshuffling
could the initial state of the pack be restored. The cards would be
worn to rags before the spades ever appeared in order at the top of
the pack.
But note that, except for the peculiar psychologicaleffectwhich
the marks on the cards have upon certainhuman minds,the 52 cards
of thepack are physicallyas good as identical. Except again as through
theirpeculiaraction on humanminds,the particularorderof the cards
is practicallywithouteffectupon the course of physicalevents. Such
shuffling as this is physicallyin the same categoryas the interdiffusion
of two quantitiesof the same gas when intercommunication is establish-
ed betweenthe two vessels containingthem at the same temperature
and pressure. This is not the typeof processwhichthe physicistterms
irreversible;it is thermodynamically indifferent.An irreversiblepro-
2 The of theparticularkindof passagefromlessprobable to more
significance
probablestateswhichplaysa role in statistical mechanics arisesfromits con-
nectionwithcertainenergyrelations, namelyfromthe factthatprocessesso
characterizedcanbe madeto yielda balanceof work.The shuffling of a packof
cardsis devoidof thischaracteristic.
• In particular,L. Dolio. See B. Petronievics, ir-
Sur la loi de l'évolution
réversible,ScienceProgress, 1919,p. 406. See also pp.418-419forbibliography.
« A. J. Lotka,Elementsof PhysicalBiology , 1925,pp.23 et seq. Butthecon-
ceptof irreversibility
is thereintroduced mainlyto distinguish an evolutionary
processfroma purelyperiodic processandto characterize theforward direction
intime.

170 HUMAN BIOLOGY
cess, as the physicistuses the words, is associated with decrease in

thermodynamic potential,withcapacityfor yieldinga balance of work,
as when one gas diffusesinto anothergas. By a purelyphysicaldevice
(semipermeablemembraneor porous partition) such diffusioncan be
made to operate,for example,a pointermovingover a graduateddial,
to warn of the presenceof an explosivegas mixture.
Now it may be objected that the shuffling of the pack of cards,
thoughit cannotaffectthe course of eventsthroughthe action of any
simple physicaldevice,could indeed be made to do so by the aid of a
sufficientlyintricate"purely physical"contrivance,such as an electric
eye (as distinguishedfrom psychologicalinterference).This is true,
but the fact is itselfinstructive.For thereis an importantdifference
betweenthe operationof the danger signal pointerunderthe influence
of gaseous diffusion, forexample,and theautomaticdoor openeractuat-
ed by the aid of an electriceye: The energyfor the operationof the
formeris furnishedby the diffusing gas itself,whereasthe electriceye
acts merelyas the "trigger"or "key" to release energysuppliedby a
separatesource. In thisit trulycopies thehumaneye,and indeedall our
sense organs, the role of which,in their part in human behavior,is
typicallyto act'as "triggers,"5operated,as a rule,by the applicationof
a verysmall dose6of externalenergy,the "stimulus,"to set in actiona
whollydisproportionate amountof energydrawn fromthe totalsources
available in the livingbody,or, by secondarytriggeraction,fromstill
more abundant sources outside the body.These are cases of "relay"
action,of whichprobablythe most advanced example,in contrivances
of humanconstruction, is the amplifierof a radio set.
The fact is, in the last analysis irreversibilityis a relativething,
and is due to restrictionsplaced naturally(by physicalor economic
5 The significance
of trigger actionseemsto be havebeenfirstpointed outby
N. L. S. Carnot, andmorespecifically byBarréde SaintVenant, whospeaksof
it as travailde décrochement , as in the pullingof a déclic (trigger). See
Boussinesq, Coursde physique mathématique, TomeIII, p. XXXII (in section
"Conciliationdu véritabledéterminisme avec l'existence
de la vie et de la liberté
morale").Paris,Gauthier- Villars,1922,butreferring to originalpublications in
1872,1878.
e to S. Hecht,underthe mostfavorable
According conditions, the smallest
amount of lightwhichthehuman eyecandetectis 58-148quanta,representing an
energyof 2-6X 10-10 ergs. Actuallyonlyaboutone-tenth of thisis absorbed
bytheretina ; therestis lostbycornealreflection,absorptionbyocularmediaand
passingon beyond theretina.(Abstract in Nature, July1, 1944,p. 13)

constraints)or arbitrarilyupon operationspermitted.Just as the tri-

sectionof an angle is or is not an impossiblefeat,accordingas we pro-
hibitor allow the use of instruments otherthan straightedge and com-
pass, so a process may be described as irreversibleor not, according
as we are restricted(by physicalor economicconstraints)to operations
in bulk,or are allowed to operate separatelyupon individualelements.
While these factsbear an importantrelationto the centralproblem
of organic evolution,the vagueness of the concept of irreversibility
in its broader interpretation demands very special caution in its ap-
plication to the general problemof evolution. It is quite true that a
of
species irreversibility verylike thatcontemplatedin thermodynamics,
as applied to processes on a molecularscale, plays an importantrole
also in large scale processes,in whichour facultyof "sorting,"or "un-
shuffling," i. e. reversingthe spontaneous"mixing"processesof nature,
gives us a materialadvantage. But merelyto state that "a biological
species neverretracesits steps,"or that"when a race has lived its term
it comes no more again," does not characterizean "irreversible"pro-
cess in any usefulsense,forthisdoes not distinguishthe chainof events
froma mere "changefulsequence." If the world's eventstakenin his-
toricalorderA, B, C, . . . are a changefulsequence,the same would be
true of the invertedsequence . . . C, B, A. This kind of "principleof
irreversibility" has no prognosticvalue. It tells us at best that if the
sequence А, В , С happens,thenthe sequence С, В, A will not happen,
and vice versa; but it does not tell us which of the two actuallywill
happen. Aftertheeventit calls theone thathas happened"irreversible/'7
Another characterizationof the direction of evolution that has
been given is that it is the directionof increasingorganization.Those
who have in the past propoundedthis view have not told us how to
measureorganization,as would be necessaryin ordereitherto establish
or to refutesuch characterization.8But assumingthat an appropriate
measurewere forthcoming, this could at best lead to an empiricalob-
1 "Dollo'sLaw of Irreversibility,
evenin its modern phrasing:Evolution is
reversiblein thatstructures oncegainedmaybe lost,butirreversible
or functions
inthatstructures oncelostcanneverbe regained,"is notborneoutbyfacts.A. J.
E. CaveandR. WheelerHaines,Nature , Nov.4, 1944,p. 579.
9 С. C. Lienauhas presented an interesting
suggestion forthemathematical
treatmentof thisproblem. Butthisappliesto specialcases. It doesnotenableus,
forexample, and to comparewitheach other,as to
to appraisequantitatively,
degreeoforganization, theelephant,
thefrog,thehousefly,theoyster or amoeba.

172 HUMAN BIOLOGY
servationthatevolutionin a large numberof cases has proceededalong

the line of increasingorganization.While such studieshave a definite
interest,theybelongin the domainof descriptivescience. What we seek
is a deductiveschemein whichthe law of evolutionis seen to flowas a
necessaryconsequencefromfundamental laws,as a resultof thephysical
properties of the organisms and the systemof whichtheyformpart.
The standardsof the exact sciences look beyondmere description.
They aim at establishingcoherentdisciplineswithinwhich,by the ap-
plicationof relativelyfew fundamentalprinciples,the course of events
can be rigorouslydeduced for innumerablespecificsituations. So, for
example,an applicationof the firstand secondlaws of thermodynamics
(the laws of conservationand of dissipationof energy)to a systemcom-
posed of any liquid and its saturatedvapor enables us to deduce the
latentheat of vaporizationof the liquid if threecharacteristic proper-
ties of the substanceare known,namelyits specificvolumein the liquid
and vapor form,and the temperature of its vapor pressure.
coefficient
The phenomenonof increasingorganization,which in a somewhat
loose qualitativesense may be admittedto be a commoncharacteristic
of organicevolution,is far too complicatedand too inexactlydefined9
a conceptionto be classed withsuch basic principlesas the two laws of
thermodynamics, for example. It is ratherthe sort of thingthat we
mighthope to derive as a consequenceof the propertiesof livingor-
ganismsand some othermorebasic principleor principles.
To accept a "law" of increasingorganizationas a basic principle
would be to shutthe door to furtherimportantanalysisby prematurely
pronouncingas solved a problemthatactuallyis hardlyeven approach-
ed : to establishthe fundamentallaw governingthe course of evolution
in a systemcomposed of aggregationsof energytransformersof a
9 To of increasing without
speakvaguely organization, anymethod
indicating of
measuring,is merely another wayof statingtheobviousfactthatcertain of the
mostsuccessfulcompetitors owetheirsuccessto adaptation themto deal
enabling
witha greaterandgreater anddiversity
variety of circumstances.Thisis achieved
bytheacquisition of certainstructures
performing - thatis, by
certainfunctions
Butthisis onlyonedirection
organization. inwhichevolution mayproceed.While
certainspecieshavethusmovedin thedirection of greaterand greatercapacity
forindividualsurvival undervariedconditions,othershaveassuredracesurvival
bygreatfertilityof thespeciesin thefaceof a lowerdegreeof adaptation of the
individual.
Thesetwotypessurvivetogether, theone,in manycases,depending
on theotherforfood.A shining exampleof thisbilateraladaptationis manand
certaindomesticanimalskeptforslaughter.

peculiar type,competingfor the substancesand energiesrequisitefor

theircontinuedexistenceand operation. In this search we are looking,
not fora new empiricalformulation, but fornecessaryrelationsdeduced
fromknownuniversalprinciples.
From a purely formal10statementin mathematicalsymbolsof the
factthatin generalthe rate of growthof each organicspecies is "some
function"of the amountsof coexistingspecies,it is possible,on rather
broad assumptions,to set up a function(or, indeed,a whole class of
functions)of the amounts (masses) of the several species,whichhas
the propertyof approachinga maximumor a minimumas the several
coexistingspecies proceed toward a stable adjustment. Such purely
formalexpressionshave been indicatedby Lotka,11and in greaterde-
tail and perfectionby Volterra.12The latterauthornames the function
thusestablishedby himthe "demographicpotential"of the system.This
use of the term "potential" cannot be commended. Potential is a
10Sucha haveanydeepersigni-
purelyformalstatement doesnotnecessarily
ficance.Cf. G. D. Birkhoff, "One can manageto obtaina variational principle
appropriate to almostanyphysical or mathematical theory," Scientific Monthly,
1944,vol. 58, p. 54-
11A. J. Lotka,Evolution and Irreversibility, ScienceProgress,1920,vol. 14,
pp.406,412.
12V. de biologiemathématique, Acta Biotheoretica,1937»
Volterra, Principes
vol.3, partI. See especiallypp.18,20,30,wherealso thetermsénergie démogra-
phiqueand principe de moindre actionen biologieare usedin a sensenotcon-
forming withtheirstrictusagein physics.Otherauthors whohaveemployed the
term"potential" in similarlyloosefashion are R. N. Chapman, AnimalEcology ,
1926,p. 145,and L. Hersch,Publications des scienceséconomiques et socialesde
l'Universitéde Genève , 1944,p. 64.
Another loosetransference of a verbalism fromthedomainof physics to that
of biologicalphenomena, madewithout adequatecritical examination of theunder-
lyingassumptions andtheconditions foritsvalidity, is thetentativeapplication of
thePrinciple of Le Chatelier by variousauthors(e. g., O. D. Chwolson, F. Le
Dantec,W. D. Bancroft).How incautious suchapplications are canbe seenfrom
thefactthatevenin thedomainof physical systems, whereconditions are much
simpler and moreclearlydefined, erroneous applicationshavebeencommon, as
pointed outbyP. Ehrenfest ( Zeitschr. /. phys.Chemie ., 1911,p. 735). An analysis
of theconditions forthevalidapplication of theprinciple cannotbe entered into
within thescopeof thisarticle.For suchthereadermustbe referred to myPrin-
ciplesof PhysicalBiology(1925), ChapterXXII, Displacement of Equilibrium,
especiallypp.281et seq.,andthereferences therecited,in particular, A. J. Lotka,
The GeneralConditions fortheValidityof thePrinciple of Le Chatelier, Proc.
Amer.Acad.ArtsandSciences f 1922,vol.27,p. 21; The Principle of Le Chatelier
in BiologyandMedicine, Am.Jl.of Hygiene , 1923,vol.3, p. 355.

174 HUMAN BIOLOGY
technicaltermin physics,of the dimensionsof an energy,and Volterra's

analysisnowhereintroducesconsiderationsof energyor of the peculiar
mechanicalpropertiesof the organism. If we were dealingin problems
whollyunconnectedwithenergetics,the borrowingof a termfromthat
fieldin a purelyfigurativesense mightbe condoned. But, quite on the
contrary,the energyrelationsof the systemof organic nature are a
deeply fundamentalfeature,in which potentialsof some kind, in the
physicist'smeaning,may be expectedto play a role. To christena func-
tionas a potential,whichdoes nothave thedimensionsand qualitiesof a
potentialin the physicist'smeaning,is to create the impressionthat a
problemhas been solvedwhich,actually,stillawaits solution. This does
notmake fortheadvancementof science.
A bettersignpostto pointus in the rightdirectionis given in Bert-
rand Russell's An Outline of Philosophy:13
Everylivingthingis a sortof imperialist, seekingto transform

as muchas possibleof itsenvironment intoitselfand its seed....
We mayregardthewholeof evolution as flowingfromthis'chemical
imperialism* of livingmatter.Of thismanis thelastexample(so
far). He transforms thesurfaceof theglobebyirrigation, mining,
quarrying, making canalsand railways,breedingcertainanimalsand
destroyingothers ; andwhenwe askourselves, fromthestandpoint of
an outsideobserver, whatis theendachieved byall theseactivities,
we findthatit can be summed up intoone verysimpleformula:to
transform as muchas possible oftheearth'ssurface intohuman bodies
.... Andinpursuing thesimplepurposeofmaximizing theamount of
human life,wehaveat anyratetheconsolation of feelingat onewith
thewholemovement of livingthings
fromtheirearliest originonthis
planet.
Now a good signpostpoints the way, but it also tells us that we
have not yet arrived. If each species seeks to maximizeits own mass,
what is the over-allresult? It is, of course,conceivablethat ultimately
one single species (perhaps man) mightdominatethe scene to the ex-
clusion of all others. But shortof this eventuality,
or at any rate until
it is attained,the problemis one of distribution.What determinesthe
distribution of the totalmatterof the systemamong the several species
and individualorganisms,and thesuccessivechangesin thisdistribution?
Is theresome significantphysicalquantitywhich these competingor-
ganismsmaximizeby theircollectiveactivities?
13Bertrand
Russell,An Outlineof Philosophy
, London,1927,p. 27.

Problemsof distribution are not new to the physicalsciences. The

whole topic of Change of State deals with preciselysuch problems:
The distribution of a mass of water betweenthe liquid and the vapor
phase, or betweenthe solid and the liquid phase, under specifiedcon-
ditionsof volume,temperature, etc.; the distributionof hydrogenand
oxygenbetweenthe threecomponentsof a systemcomposedof these
substancesin elementaryform,H2 and 02 on the one hand, and in
combinationas water,H20, on the other,again under specifiedcondi-
tions,etc. As applied to thesecases the law of evolutionis well known
and clearlydefined. It statesthat for an isolated system,for example,
the entropyof the systemincreasesto a maximumas the systemap-
proaches equilibrium;or, more generally,that certain clearly defined
functionsof the parametersspecifyingthe state of the system - its
thermodynamic - approach a minimum.
potentials
A special example whichhas oftenbeen cited,because of a certain
superficialanalogy to the growthof a colonyof bacteriain a suitable
nutrientmedium,is that of the crystallizationfrom a supersaturated
solution,upon the introductionof a crystal"germ" of the dissolved
substance. So long as the supersaturatedsolutionis left undisturbed
it mayremainas such forlong,perhapsindefinite periods,thoughseem-
ingly spontaneouscrystallizationmay also take place. This restricted
type of stabilityhas sometimesbeen spoken of as the metastablestate.
Its thermodynamic potentialis not at the minimumpossiblefor the sys-
tem,and hence,as soon as an available "path" is presented(by the in-
troductionof the crystalgerm), the transformation to the crystalform
is initiated. It continuesuntil that particular amount of the solid has
formedwhich is requiredto bringabout stable equilibrium,that is, to
makethethermodynamic potentiala minimum.
If this were merely outward appearance of analogy, the oft re-
peated example would hardlybe worthquoting again here. But it is
literallytrue that the nutrientsolutionis "metastable"in the absence
of a germof an organismcapableof growthin it. Whetherin primordial
naturesomewherelocallysuch a state presenteditself,and whetherthe
metastablestate could pass over into some kind of elementary"living"
matterwithoutthepresenceof a preexistinggerm,just as crystallization
of a supersaturatedsolutioncan take place even in the absence of a

176 HUMAN BIOLOGY
crystalgerm,thatis todaystillone of theunsolvedproblemsof science.14

But if such was the originof life, we know that it was followedby a
long chain of evolutionarydevelopment.And just as the continued
growthof the crystal,fromthe firstgerm to the completelyformed
solid phase, is governedby the law thatthis processproceedswithcon-
tinuousdiminutionof thermodynamic potentialof the systemsolution
plus crystal, so we may expect that the law of organicevolutiontakes
the formthatit is accompaniedin the long run withdiminutionof some
function,analogous to thermodynamic potential,of the parametersde-
finingthephysicalstateof the systemas a whole.
While the problemof organic evolution,as a problemin the dis-
tributionof matteramong the componentsof a material system,is
formallyin the same generalcategoryas theseproblemsof distribution
in physico-chemical systems,it differs,of course, in certainimportant
respects from the specific problems considered in physico-chemical
dynamicsand statics. What are actuallythe significantpoints of dif-
ference?
First,thatthe ultimateindividualcomponentsrecognizedin physico-
chemicaltransformation - moleculesand atoms- escape directobserva-
tion by any of the ordinarymethods. It is the bulk propertiesand ef-
fects,such as volume,pressure,temperature, etc., of the massive com-
ponents,which are directlyaccessible to observation,and in termsof
whichthermodynamics receivesits data and states its conclusions. The
investigationof the detailedactionsof individualmoleculesand atoms
requiresaltogetherdifferent, highlyrefinedtechniques.
The establishment of the statisticalrelationsbetweenthe individual
actions and the bulk effectsis an additionaldisciplineof great inter-
est- statisticalmechanics- but is not primarilynecessaryfor the know-
ledge of the bulk effects,since theyare directlyobservedwith greater
ease thanthe individualactions.
Diametricallyoppositeis the situationwhenwe studyorganicevolu-
tion. Here it is the individuals,the separate organisms,that are ac-
cessible to direct observation,while the collectiveor bulk effectsof
aggregatesof organisms,as a whole, (in termsof whichwe must ex-
14This observation,
casuallyintroduced
by Ostwaldin one of his lecturesin
1902,wasthe"trigger"
thatsetoffthetrainof thought inmysubsequent
developed
andsummarized,
publications, in part,inmyElementsof PhysicalBiology(1925)
and ThéorieAnalytique
des Associations
Biologiques
(1934,1939).

pect the law of organicevolutionto be primarilyexpressed) will neces-

sitatethe developmentof a special branchof statisticaldynamics.
Secondly, physico-chemicaltransformations, in all their variety,
are neverthelesslimitedby certainnarrow restrictionswhich hold for
cil time. These transformations run, as it were, along a single track.
So for example in the case already cited, in a systemcomposed of
specifiedamountsof hydrogenand oxygen,therelativeamountsof H2, 02
and H20 may undergoa progressivechange (toward an equilibrium
if externalconditionsare constant), but they will never give rise to
hydrochloric acid, for example. This may be expressedby sayingthat
ordinaryphysico-chemical transformations (exclusive of transmutations
of elements) are bound by certainequations of constraint(the "reac-
tionsequations") whichare fixedfor all time.
The transformations betweenand withinorganic species are of a
different order,becausetheyare in themaintransformations of structure,
which is capable of infinitevariation. It is true that here also there
are equations of constraint.An eagle is not hatched from a duck's
egg. But a white duck may have parti-coloredducklings. Again, it
is not possibleat the presentepoch of the world's historyfor grass or
hay to be convertedintomammoth.But some tensor hundredsof thou-
sands of years ago this transformation, or somethingvery much like
it, was not onlypossible,it was a daily occurrence.Thus the equations
of constraintthatdelimitthe possibletransformations of organicevolu-
tion are not onlymore liberalthan those of physico-chemical evolution,
they are distinguishedby the fact that they are functions of the time,
whereas the physico-chemical equations of constraint (e. g. the "reac-
tionequations"of chemistry)are rigidlyfixedforall time.
One consequenceof this distinctivecharacterof the equations of
constraintin organic evolutionis that the process is one that never
ends,since structureis capable of infinitevariation. In contrastto this,
a reaction such as that already cited between hydrogenand oxygen
proceeds(undergivencondition)to an equilibrium, oftenreachedwithin
a shortspace of time.16
The fact that the equations of constraintin organic evolutionare
functionsof thetimeis closelyrelatedto a thirdcharacteristic.Where-
as the componentsof the systems considered in physico-chemical
15Thatis, forall theendstatemayneverbe

practical
purposes.Theoretically
butonlyapproached
fullyreached, to within
a negligible
distance.

178 HUMAN BIOLOGY
dynamics- - elements,compounds,"phases"- are strictlyhomogeneous,

the componentbiologicalspecies in the systemof nature are only re-
lativelyhomogeneous.In comparisonwith the discontinuousgaps that
separatedifferent species,the variationswithineach species,describable
in terms of frequencydistributionsof characteristicsamong its in-
dividuals,are eitherpracticallycontinuous,or, if discontinuous,pro-
ceed by steps much less extremethan those that separate the several
species.
In accordance with this, analytic formulationdistinguishestwo
aspectsof organicevolution: changesin thedistribution of matteramong
the differentspecies- inter-speciesevolution; and changes in the dis-
tributionof the individualswithineach species,as definedby suitable
frequencyfunctionsof their characteristics-intra-speciesevolution.
It is the changingcharacterof organic species that impartsto the
equationsof constrainttheirdependenceon time. This featurehas no
counterpartin physico-chemical evolution,since there the components
are homogeneousand invariableand presentno opportunityfor intra-
species evolution. On the contrary,in the systemof organic nature,
intra-speciesevolution has so prominentlyengaged the attentionof
biologists,that attemptsto indicate the directionof evolution have
commonlybeen couchedwith referenceto individualspecies. Actually,
as alreadyhinted,any adequate treatment of the problemmustenvisage
the evolvingsystemas a whole- the aggregateof coexistingspeciesand
their inorganic environment.16 Anyone accustomed to the discipline
of physico-chemical dynamics knows that it would be quite out of the
question to formulate the law of evolution of the systemby reference
to one of the componentsalone. The law of organicevolution,likewise,
must undoubtedlybe formulatedin terms of the systemas a whole
evolvingunder the floodof lightreceivedfromthe sun.
And this bringsus to the fourthtypicaldifference that distinguish-
es organicevolutionfrommostof the physico-chemical transformations
usually considered (adiabatic or isothermalreactions), namely that
the latterare envisagedas takingplace under conditionsleading to an
equilibrium. Absolutely fundamentalto organic evolution is that it
takes its course in a systemexposed to a continuousstreamof energy
fromthe sun, so that the approachis not towardsequilibrium,but to-
ie For a
particularly surveyof the planetary
illuminating relations
involved,
see W. Vernadsky,La Géochimie,
Paris (Alean), 1924.

wards a stationarystatemaintainedwithcontinuousexpenditureof free

energy.17
And directlyrelatedto this is the fact to which referencehas al-
ready been made in passing, namelythat triggeraction plays an im-
portantrole in organicevolution.For triggeractionpresupposesa fund
of free energyreadyto be released,and, as was pointedout years ago
by Boltzmann,the life struggleis primarilya competitionfor available
energy.What we have beforeus in organicnatureis a systemcomposed
of aggregatesof energytransformers adapted by theircompositionand
structureto guide available energyinto such channelsas lead to their
maintenanceand growth. A special branch of physics needs to be
developed,the statisticaldynamicsof systemsof energytransformers .
In the developmentof this it will not be necessaryor even desirableto
deal primarilywith specificliving organisms,but rather with trans-
formertypespossessingcertainpropertiescharacteristic of the physical
modus operandiof livingorganisms. The kind of problemthen to be
studied will be the relationbetweenthe distributionof matterin the
systemon the one hand, and on the otherthe particularpropertiesand
variationin propertiesof the several types of transformers of which
the systemis composed. In so far as the systemof organicnature is
subject to physicallaw, certainat least of the relationsdeduced for
the systemsstudiedin thisbranchof physicsshouldbe foundapplicable
to the systemof organicnatureand its evolution,just as thatsystemis,
forexample,subjectto thelaw of conservationof energywhichis usual-
ly thoughtof as applicableprimarilyto inorganicsystems.18
Among the transformerswhich this branch of physics will have
to consider,one importantclass obtainsits supplyof available energy
from sources distributeddiscontinuously, in separate masses, through
the systemin whichtheyoperate. This necessitatesstructuresor "ap-
paratus" whereby"collisions" or "encounters"between those trans-
formersand their energy sources are renderedprobable. The more
perfectthe functioning of this apparatus,the greaterwill be the prob-
17In thisrespectit is similarto photochemical and indeed,
reactions, it might
be regardedas a photochemicalreaction on a heroicscale,accompaniedbyimport-
ant structuralchanges.However,in organicevolution, owingto intra-species
changes,thestateis onlyrelatively stationary;it resemblesratherwhathas been
(inaccurately)termed a "moving equilibrium."
18Thoughtheprinciple was originallyrecognized byJ. R. Meyeron thebasis
of physiological
observations.

i8o HUMAN BIOLOGY
abilityof theseencounters,and the less the energyexpendedin propor-

tionto the energycapturedin the encounter,so as to yielda net balance
of gain.
In the apparatus by which the probabilityof favorableencounters
is ensured or enhanced four typicalelementsare to be distinguished,
as follows:
1. Receptors.The action of the transformermust be a function
of thepositionof the sources. But thatactionis primarilya functionof
the state of the transformeritself. Hence there must be apparatus
to providethatthe state of the environment is in some manner"repre-
sented"or "depicted"in the transformer.This may be by opticalimage
formationby a lens,as in thehumaneye or in an automaticdoor opener
operatedby a photoelectric cell; but moregenerally,the processof "de-
picting"19 may take a varietyof forms.What is characteristic is thatthe
amountof energyinvolvedin the processmay be verysmall.
2. Free energy. Associated with the transformer is a fund of free
20 to be released that
energy ready by triggeraction, is, by an expendi-
tureof an amountof energybearingin generalno quantitativerelation
to the energyreleased,which can thus be far in excess of the energy
applied at the "trigger."
3. Effectors,that is, structuresby which the energy released by
triggeraction fromthe available fund is applied to react upon the en-
vironmentso as to bringabout an encounterwith an energysource to
be captured,or so as to avoid an encounterin which the transformer
itselfwill be capturedor injured,i. е., made more or less inoperative.
19That of a one-to-one betweencertainfeatures

is, establishing correspondence
of theenvironment andcertain appropriatefeatures
of thetransformer. It is no
mereaccident of languagethatwe speakof thisgeneralized processof "image
formation" in ourselvesas information. By it we are informed of conditions in
theenvironment.
In thecase of mantheprimary representation
(depiction)of theenvironment
in theindividual is further supplemented or elaboratedby-a facultyor a setof
facultieswhichwe maydesignate as theelaboratore.
Theirmostrefined product
is thescientificworldpicture.
* It can be notedhere
onlyin passingthatthepossession of sucha fundof
freeenergy is whatgivestheopportunity forpurposive actionandtheexhibition
of thephenomenon of will, thatis, actionteleologically
aimedat future effects.
For a discussion of thisthereadermaybe referred to Chapters XXVlII et seq.
ofmyElements ofPhysical Biology, and,fora morerecent review,to A. J.Lotka,
Evolution andThermodynamics, ScienceandSociety,1944,vol.8, p. 161.

For since these transformers possess a fund of free energythey can

themselvesbecomeobjects for captureby competingtransformers.The
avoidanceof captureis made possibleby the existenceof refugesscat-
teredthroughthe topographyover which theyoperate.
4. Adjustors. Lastly, in order that the receptorsand effectorsmay
functionsuccessfullyin ensuringthe probabilityof encounterswith
energysources and in wardingoffunfavorableencounters,the trans-
formermust compriseapparatus by which the action of the effectors
upon the environment is suitablyadjusted to the "information"supplied
by the receptors.
Since it is characteristicof triggeraction that the energyreleased
may farexceed theenergyspentin workingthetrigger,whilethe energy
capturedmay still furtherexceed the energyexpended fromthe avail-
able fund of the transformer, it followsthat a transformer or an ag-
gregateof transformers equipped withapparatusof the typedescribed
is capable of continuedfunctioning,obtainingsufficient energy from
captured sources to the
defray energy cost of capturingfurther sources.
In fact,theremay be a balance left over for accumulation,whichmay
take the form of growthof the transformeror of the aggregateof
transformers.
In this discussionit has been assumed that there exist "discrete"
sources available for capture,scatteredthroughthe fieldof operation
of the transformers of the typeso far considered.
That such sources exist and are continuallyreplenishedin nature
is due to the existenceof anothertype of energytransformers, which
themselvesdraw their supply of energyand substance from sources
continuously distributedin space, namelysunlighton the one hand,and
on the other,gaseous or dissolvedsubstanceswhichdriftto themspon-
taneouslyby diffusionand seepage. These transformers can according-
ly remain essentiallypassive and do not require the "ingenious" ap-
paratus for capturingenergyby triggeraction. At the same timethey
lose the benefitof the power activelyto avoid unfavorableencounters.
They do, of course,possess variouspassivemeansof protection, some of
which,incidentally,are available also to animals.
Such are the fundamentaldata with which the statisticaldynamics
of aggregatesof energytransformerswill have to deal. Indications
of the directionin which the developmentof this branch of physics

I82 HUMAN BIOLOGY
may be expected to move have been given by the writerelsewhere.21

For our presentpurpose only certainbroad observationswill here be
made.
While the transformers in natureare "living" animals and plants,
it will be desirableto develop the disciplineof statisticaldynamicsof
aggregatesof transformer irrespectiveof thisfact. The resultsachieved
must then be applicable to the world of organismsin so far as they
are energytransformers having the physicalpropertiestaken in view
in thatdiscipline,just as the law of conservationof energy,is applicable
to them.
Dealing with purely physical data affordscertain importantad-
vantages. We have completecontrolof the particularpropertieswhich
we may assign to the transformers under consideration,withoutrais-
ing the question whether such a transformer actuallyexists or ever has
existed. As Karl Pearson has remarked: "In dealing with natural
phenomena.... the mathematician has to simplifythe conditionsuntil
theyreach the attenuatedcharacterwhichlies withinthe power of his
analysis."
This process can be carriedmuch fartherboth experimentally and
in
conceptually purelyphysical than in biologicalsystems. The relative
simplicityand directionof physicallaws is in no small measure due
to the factthatexperimentally we can controlconditionsso as to isolate
the factorsessentialto our enquiry,fromadventitiouscircumstances in
which for the momentwe are not interested.22
The factthatwe ourselvesare of the dimensionsof the units (trans-
formers) contemplatedin this disciplinegives us an inside view of
21A. J.Lotka,Elements ofPhysical Biology, pp.36,37,andespeciallyChapters
XXIV to XXXIV. Also,Familiesof Curvesof Pursuitand TheirIsochrones,
American MathematicalMonthly to theMathe-
, 1928,vol.35,p. 421; Contribution
maticalTheory ofCapture, ProceedingsoftheNational Academy ofSciences,1932,
vol.18,p. 172.See alsoV. Lalan,Compt. Rend.,1931,vol.192,p. 468.
32The history of Sciencewell illustratesthe importance of this simplifica-
tionfortheprogress, andespecially
theearlyprogress, ofa science.Thusastronomy
was ableto makea veryearlystartbecausenatureherself has simplifiedsomeof
thebasicproblems by reducingthedata practically to mathematical points,the
of stars(and forthenakedeyealso theplanets)in thesky.At theother
position
extreme wehavethecaseofchemistry, oneofthelastsciences to developeffective-
ly,becausenoconsiderable couldbe madeuntilrecognition
progress andseparation
of thefundamental - elements
unitsof chemistry and purecompounds - had been
achieved.

the processes under discussion,and thus enables us to select the ap-

propriateensembleof propertiesto assignto theconventionalized models
of the actual transformers presented to us in nature's organisms. At
the same timewe shall retainfull freedom,in this process of conven-
tionalization,to place the developmentof the subject on a postulational
basis, if we conductthe enquirysolely in termsof physicallydefined
transformers, since we thus avoid carryingon the discussionin terms
of "hypotheticalanimals and plants," to which some biologistsmight
take exception.
An admittedlyprimitiveillustrationof the mode of approach to
the problemis the case of a "pursuing" transformerof velocity
seekingencounterwith a "pursued" transformerof velocityv2 while
thelatterfleesin a straightline towarda refugeof safety. The pursuer,
constantlydirectedtowardthe pursued,then followsa so-called"curve
of pursuit." Whethercaptureoccursor not thendependson the relative
positionof the pursuerand the pursuedat the timewhen firststage of
the pursuit,the "stalk," begins (that is, when the pursuerfirstdirects
his course towardthe pursued) ; and dependsfurtheron the timewhen
thesecondstagebegins,namelywhenthe pursuedfirstreactsby "flight"
to the "attack" of the pursuer. The problemthus conventionalizedre-
solves itselfinto a problemof geometry.Capture certainlyoccurs,or
does not occur,accordingas a certaincircledoes or does not fall with-
in a certainellipse of dimensionsdependingon the efficiency of the
receptor-effector apparatus of the two types of transformers. If the
circleand the ellipse intersect,captureoccurs for some initialpositions
and does not occur for others. In this conventionalizedcase we can,
for example,discuss the influence,upon the probabilityof capture,of
variationsin thevelocities and v2or in the degreeof perfectionof the
receptors,etc. Carryingthe model somewhatfurther,the influenceof
thedensityof distribution of refugesin the territorymay be discussed.23
It may be objectedby some that such primitive - not to say naive-
conventionalizations of the natural phenomenaof pursuitand capture
observedamong livingorganisms,are too far removedfromany reali-
ties to be of any value.
To this the answer is, first,that the case describedis used merely
as a simpleexampleof thekindof procedureto follow,withappropriate
28For detailsthereaderis referred

to mytwopaperscitedabove,on curves
andonthetheory
of pursuit of capture.

I84 HUMAN BIOLOGY
refinements and extensions.And,second,thatit is well to be verycau-

tious in pronouncinguseless a conventionalization merelybecause it is
a seemingover-simplification. Such highlyattenuatedconventionaliza-
tionshave done excellentservicein certainfields.
The Carnot heat engine, with its perfectlyconductingand per-
fectlyinsulatingparts,is a case in point,and so is the use of a "perfect
gas"24in illustratingthederivationof thesecondlaw of thermodynamics
by a Carnot cycle. As is well known,the resultsso derivedare of far-
reachingconsequence and of the widestapplicability.
While the future developmentof such a disciplineof statistical
dynamicsof aggregatesof energytransformers should eventuallyprove
of great interestand importancein relationto the problemsof organic
evolution,certainbroad observationswith which its resultpresumably
will conformmay be statedeven now.
It has been remarked25 that the influenceof living organismsis to
retardthe dissipationof energy. So far as the plantworld is concerned
this seems essentiallytrue. Green plants act as accumulatorsstoring
sunlightwhich would otherwiseshortlybe convertedinto heat at ter-
restrialtemperatures.
But the influenceof animals is essentiallyin the oppositedirection,
sincetheiractivitiesare conductedwithdissipationof energy,catabolism
exceedinganabolism.
At firstsightthereis somethingperplexingabout thesetwo opposing
tendencies. But this perplexityexists only so long as we seek to ex-
press the driftof organicnaturein termsof a retardationor a promo-
34As to this,it is admitted thatbothlogicallyand fordidacticreasonsit is

preferableto avoidtheuse of an unrealworking substancein thisderivation of
theefficiencyof a reversible heatengine.It is nottheidealwayof treating the
though
subject, it has a certain heuristic value. I havenotedthat
andillustrative
somewhohadso beeninitiated intothefoundationsof thermodynamics gainedthe
impressionthattheuseof a "perfect" gas wasa necessary pointof theargument -
whereasjusttheopposite shouldhavebeenimpressed uponthem - thattheargu-
mentanditsconclusions are independent of theworkingsubstance,andthatthere-
fore,in the interest of simplicity of themathematical manipulations it is per-
missible
to conduct theargument on thebasisof a fictitious
working substance -
theconclusion is independent of the factthatsucha particular substance does
notexist.
38See J. Johnston, The Mechanism of Life,1921,pp.220,221,especially the
footnoteon page221.

tion of the dissipativeprocess. As soon as we franklyaccept the fact

thatbothare going on side by side the perplexityvanishes.
A simile may be helpful in illustratingthe situation. Consider a
reservoirconstructed to collectrainwater.Now let therebe two agencies
at work, one tendingto enlarge the catchmentarea, the othertending
to enlargethe outletor outletsfromthe reservoir. In one sense these
two influencesseem antagonistic,one tends to increase the volume of
water in the reservoir,the othertendsto decrease it. But in one thing
theycooperate: Togethertheyincrease the total flow throughthe re-
servoir.
From this illustrationby analogy,let us returnto the actual case
before us. The argument,as I have presentedit elsewhere,26 runs as
follows: As alreadynoted,"it has been pointedout by Boltzmannthat
the fundamentalobject of contentionin the life struggle,in the evolu-
tion of the organicworld,is available energy." In accord withthis ob-
servationis the principlethat in the strugglefor existencethe advan-
tage must go to those organismswhose energy-capturing devices are
most efficientin directingavailable energyinto channelsfavorablefor
the preservationof the species.
"Thefirsteffectofnaturalselectionthusoperating uponcompeting
specieswillbe to giverelativepreponderance to thosemostefficient
in guidingavailableenergyin themanner indicated.Primarily the
Pathof theenergyfluxthrough thesystem willbe affected.
"Butthespeciespossessing superiorenergy-capturing and direct-
ing devicesmayaccomplish something morethanmerelyto divert
to itsownadvantage energyforwhichothersare competing withit.
If sourcesare presented,capableof supplying availableenergyin
excessof thatactuallybeingtappedby theentiresystem of living
organisms,thenan opportunity is furnishedforsuitably constituted
organisms to enlargethe totalenergyflux2T through the system.
Whenever such organisms arise , naturalselectionwill operateto
preserveandincreasethem.Theresult, inthiscase,is nota merediver-
sionof theenergy fluxthrough thesystem of organicnaturealonga
newpath,butan increase of thetotalfluxthrough thatsystem.
"Again,so longas sourcesexist,capableof supplying matter of
suitable
a character forthecomposition of livingorganisms, in excess
of thatactuallyembodiedin thesystem of organicnature, so longis
* A. J.Lotka,Contribution
to theEnergetics of Evolution, of the
Proceedings
National
Academy 1922,vol.8, p. 147.
ofSciences,
27The termenergyfluxis hereusedto denotetheavailableenergy absorbed
byanddissipated
withinthesystem perunitoftime.

186 HUMAN BIOLOGY
opportunity furnishedforsuitably constituted organisms to enlarge

thetotalmassof thesystem of organicnature.Whenever suchor-
ganisms arise,naturalselectionwilloperateto preserve and increase
them, provided alwaysthereis presented a residueof untapped avail-
ableenergy.The resultwillbe to increase thetotalmassof thesys-
tem,and,withthistotalmass,also thetotalenergy fluxthrough the
system, since,otherthingsequal,thisenergyfluxis proportional to
themassof thesystem.
"Wherea limit,eitherconstant or slowlychanging,28 is imposed
uponthetotalmassavailablefortheoperation of lifeprocesses, the
availableenergyperunitof timeplacedat thedisposalof theor-
ganisms forapplicationto theirlifetasksandcontests, maybe capa-
bleofincrease byincreasingtherateofturnover oftheorganic matter
through thelifecycle.Suppose,as a simple, though ratherextreme
illustration,thatmanfoundmeansof doubling therateof growth of
crops,andof growing twocropsa yearinsteadof one. Then,with-
outchanging theaveragecropactually standing onthefields, theland
wouldbe capableof supporting doublethepresent population. If this
population wereattained, theenergyfluxthrough thesystem com-
posedof thehumanpopulation and theorganisms uponwhichit is
dependent forfood,wouldalso be doubled.Thisresultwouldbe at-
tained, notbydoubling themassof thesystem (forthematter locked
up in crops,etc.,at a givenmoment wouldbe, on theaverage,un-
changed)butbyincreasing thevelocity of circulation of massthrough
thelifecyclein thesystem.Oncemoreit is evident thatwhenever a
ж ariseswhichis so constituted as to increase the
group of organisms
rateof circulation of matter through thesystem in themanner ex-
emplified, naturalselectionwilloperateto preserve andincrease such
a group, provided alwaysthatthereis presented a residue of untapped
availableenergy, and,wherecircumstances requireit,also a residue
of masssuitable forthecomposition of livingmatter.
"To recapitulate: In everyinstance considered, naturalselection
willso operateas to increasethetotalmassof theorganicsystem,
to increase therateof circulation of matter through thesystem, and
to increase thetotalenergy fluxthrough thesystem, so longas there
is presented an unutilizedresidueof matter and availableenergy.
"Thismaybe expressed by sayingthatnaturaiselection tendsto
maketheenergy fluxthrough thesystem a maximum, so faras com-
patiblewiththeconstraints to whichthesystem is subject.
88As, forexample, if thetotalmassof thesystem is capableof accretion, but
onlyat a limited velocity,inwhichcasethephenomenon of a "moving equilibrium"
maypresent itself.CompareLotka,A. J.,Proc. Nat. Acad. Sci., 1921,vol. 7,
p. 168.
*
Owingto the factthatin existing organisms theanabiotic and catabiotic
functions are verylargelysegregated in different types(plantsand animals),
evolution willhereoperateuponsystems or groupsof at leasttwospecies,one
speciesofautotrophic anabions,andoneofheterotrophic catabions.

"It is notlawfulto inferimmediately thatevolution tendsthus

to makethisenergyfluxa maximum. For in evolution twokinds
of influences are at work: selecting influences,and generating in-
fluences. The former select,thelatterfurnish thematerial forselec-
tion.
"If thematerialfurnished forselection is strictly limited, as in
thecase of a simplechemical reaction,whichgivesriseto a finite
number of products, therangeof operation of theselective influences
is equallylimited.
"In thecase of organicevolution thesituation is verydifferent.
We haveno reasonto supposethatthereis anyfinite limitto the
number of possibletypesof organisms. In thepresent stateof our
knowledge, or ratherour ignorance, regarding the generating in-
fluences that furnishmaterialfor naturalselection, for organic
evolution, an element of uncertainty entershere.It appears, however,
at leasta prioriprobable that,amongthecertainly verylarge(if not
infinite)variety of typespresented forselection,sooneror laterthose
willoccurwhichgivetheopportunity forselection to operatein the
direction indicated,80namelyso as to increasethetotalmassof the
system, therateof circulation of massthrough thesystem, and the
totalenergyfluxthrough thesystem.If thiscondition is satisfied,
thelaw of selection becomes also thelaw of evolution :
"Evolution, in thesecircumstances, proceedsin suchdirection as
to makethetotalenergy fluxthrough thesystem a maximum com-
patiblewiththeconstraints"*1
over relativelyshortperiodsof time,when
As for theseconstraints,
we are dealing mainly with inter-speciesevolution,they are defined
essentiallyby transformationcoefficientsindicatingin what ratio the
mass added in the growthof one species stands to the masses captured
fromotherspecies or fromthe environment.For, growthin mass of
organismstakes place by the assimilationof food, thoughgrowthin
numberstakes place by reproduction(births), whichthus acts as pace-
makerforthe growthin mass.
Over extendedperiodsof time,as alreadynoted,the constraintsare
functionsof thetime,and the empiricaldata whichdefinethemare sup-
30One is inclinedhereto give at least qualifiedassentto the sayingof

Herodotus:"If oneis sufficientlylavishwithtime,everything possiblehappens."
(CitedbyC. E. Guye,Physicochemical Evolution,Methuen & Co.,London,1925,
p. 30. Thisis essentially in Englishof a seriesof papersoriginal-
a republication
lyappearing in variousFrenchjournals,
1917 to 1920.)
31A further analytical
developmentbaseduponthisprinciple has beensug-
gestedby N. Rashevsky, Outlineof a New Mathematical Approach to General
Biology, II, Bulletinof MathematicalBiophysics,1943,vol. 5, p. 49.

188 HUMAN BIOLOGY
plied mainly- but with one highlysignificantexception - by the find-

ings of genetics.
The one outstandingexceptionis the human species. Here evolu-
tion,especiallyin more recenttimes,has followedan entirelynew path.
In place of slow adaptationof anatomicalstructureand physiological
function in successive generations by selective survival, increased
adaptationhas been achievedby the incomparablymore rapid develop-
ment of "artificial"aids to our native receptor-effector apparatus,in
a process thatmightbe termedexosomaticevolution.32
Until recentlythe net effectof these new implementsof evolution
has been essentiallyin the same directionas the main current. Man,
one of thelatest,and in his own judgmentthehighestproductof evolu-
tion, has hithertosignallyconformedwith the principleof increasing
energyflux.83By ingeniouscontrivanceshe has immenselyrefinedand
multipliedthe operationof his receptor-effector apparatus. The excess
of energycaptured,over the energybarelysufficient for mere mainten-
ance,has in his case grownto a whollyunparalleledmagnitude.Normal-
ly this leaves him with a large balance available for "play" activities
and luxuries. And some of his play activitieshave turnedout to be a
mostprofitablereinvestment. For amongthemmustbe classed scientific
research indulged in primarilyout of curiosity,but resultingamong
otherthingsin thatcompleterecastingof methodsof productionwhich
is knownas the industrialand agriculturalrevolution.Aside fromits
directbenefitsthis"has made it possibleto spendrelativelylargeamounts
on sanitaryimprovements, on medical education and research,and,
above all, on betterliving among the masses of the people. This in-
33I haveelsewhere
( Proceedings of theAmerican Philosophical
Society,1939,
vol. 80, p. 625) graphically representedin quantitativemeasurethe rocket-like
ascent,in modern times,of humanknowledge and technicalskill. See also,the
editorial"FortyYears of Plenty"in theBulletinof theAmerican Association
fortheAdvancement of Science,1944,vol.3, no. 10,p. 73.
33Of thisthe of solarenergystoredup frompastages in coal
exploitation
is in thepresent era theoutstanding, thoughperhapstransitory, example."The
problem of economy in husbanding resourceswillnotriseto its fullimportance
untiltheavailableresources aremorecompletely tapped thantheyaretoday.Every
indication is thatmanwilllearnto utilizesomeof thesunlight thatnowgoesto
waste.The generaleffect willbe to increasetherateof energy fluxthrough the
system of organicnature "
(A. J.Lotka,Elements of PhysicalBiology,1925,
PP 357-358).Compare also F. H. Pike,The DrivingForcein Evolution, Ecology,
1929,p. 167.Thisalso contains notes.
bibliographical

crease in the productivity of humanlabor may,therefore,veryproperly

be called the basis of the declinein the deathrate whichhas takenplace
in almostall parts of the WesternWorld duringthe last century."34
This is one side of thepicture.But thereare also less pleasingaspects.
The same ingeniouscontrivancesthat extend our view, that speed our
travel, and multiplyour strengthin beneficentpursuits,are equally
potentto destroy.Then, in a worldat war as today,contendingarmies
exchangedeath against death,36while among the inhabitantsof the oc-
cupied countries"a disproportionate expenditureof energyis required
merely to maintain life and satisfythe most elementarydemands of
food and cleanliness,with the almost inevitableresult that in many
cases habitsof personalhygieneare relaxedand all interestsbeyondthe
merestruggleforexistenceare stifled.,,3e
It is not onlyin war thatthe trendof humanevolutionhas brought
special problems. With the vastly multipliedefficiency of modernin-
as
dustry, compared with primitivelabor, the necessitiesof life can be
producedin much less than the full workingtimeof the gainfullyem-
ployedpopulation.37Hence, to secure full employment much of the in-
dustrialactivitymust be expended on the productionof luxuries and
near-luxuries.This in itselfwould be no disadvantage. Quite on the
contrary,if continuedfull or approximatelyfull productiveactivity
were assured,this would mean the enjoymentof these luxuriesby the
consumers,among whom are the producersthemselves. But there is
84W. S.
Thompson, Plentyof People,JaquesCattellPress,1944,p. 64.
" The latest in thisfieldis theunlocking of atomicenergyfor
development
purposesof war. It remains to be seenwhether thisnewchannelforthefluxof
energy through theevolving system of whichhumansocietyformspart,will in
thecourseof timebe putto constructive use. In thatcase thiswillbe thesuper-
lativeexampleof theprinciple of maximum energyfluxas characterizing the
directionof evolution.
3eInternationalLabourOffice, The Healthof Children in OccupiedEurope,
Montreal, November 1943,p. 28.
" W. B. Kaempffert, Advancein Relationto Popula-
Compare Technological
tionTrends,Proceedings of theAmerican PhilosophicalSociety,1939,vol.80,p.
563: "In thedecadefrom1920to 1930.... thenation's outputincreased 46 per-
cent,butthelaborforceonly16percent." Compare also theeditorial
"FortyYears
of Plenty,"Bulletinof theAmerican AssoiationfortheAdvancement of Science,
1944,vol.3, no. 10,p. 73. In thefourdecades1900to 1940thepopulation of the
UnitedStatesincreased 70 percent, whilethenumber of farmworkers decreased
morethan10 percent.

içk> HUMAN BIOLOGY
this serious drawback: While the necessitiesof life, fromtheirvery

nature, insistentlydemand continued production,no such powerful
inherentdrive exists for the luxurytrades. And so, with our present
economicsystem,when fromtime to time periods occur duringwhich
the productionof luxuriesand near luxuries is depressed,this results
in a plague of unemployment, since idlenessat such timesis quite un-
evenlydivided,some retainingfull time employment, while othersare
wholly without work. Then we have the incongruousspectaclethat in
the midstof plentythereis widespreadwant.38
While the occurrenceof the more severe economicfluctuations are,
to say the least,annoyingand do cause muchhardship,it cannotbe said
thattheyofferany seriousthreatto the maintenanceof the species,and
it mayreasonablybe hopedthatin timebetteradjustmentsof production,
distribution and consumption maybe achieved,withreductionor elimina-
tionof at least the more extremefluctuations.
But the effectof the greatly increased productivityof labor in
shiftingthecenterof gravityof thetotalactivitytowardluxuryproduc-
tion has othermore serious results. It is obviouslyin the interestof
producersto stimulatedemand for luxuries. This demand,unlikethat
for necessities,is very elastic. The demand for personal necessities
needs no particularstimulation - that is what we mean by necessities-
nor could it be very greatlyenhancedby any attemptat stimulation.
People's appetitefor food is limited. Their appetite for automobiles,
radios,furcoats,jewelry,actuallyseemsto followthe ruleof theFrench
proverbl'appétitvienten mangeant.At any rate it is highlysusceptible
to suggestion,whichour advertisingfraternity has developed,if not to
a fineart,at least to an importunatetechnique.
Now overstimulation of the demand for luxuriesis not withoutits
dangers. Necessities for the species are not always felt as necessities
for the individual. Few personsmay be temptedto go hungryin order
to buy jewels. But many actuallydo go withoutchildren,or without
an adequate numberof them,in orderto maintaincertainstandardsof
38The measureadoptedby Germany, of absorbing

excessproduc-
desperate
forand wagingaggressive
tivityby preparing war,has proveda catastrophic
failureevenfroma criminally
nationalistic
pointof view.

living.39Nature tricksthe individualinto doing thingsadvantageousto

thespecies,by linkingthedoingof themwithpleasurefortheindividual.
The trickworkswell, untilthe individualdiscoversways of separating
the links, and reaping the pleasure withoutacceptingthe cost. Here
again man's ingenuityhas workedagainstthe advantageof the species,
or at least of certainbranchesof the species,as the statisticsshow. The
net reproduction rate,thatis the ratio of total birthsin successivegen-
erations,has fallenwell below unity,the requiredminimum,for many
of thecountriesof Westerncivilization,as shownin Table 1.
TABLE i
Net reproduction *
rate according to age-specific and fertility
mortality
at specified
dates
COUNTRY DATE RATE
Austria 1935 -64

Belgium 1941 .67
Czechoslovakia 1935 .79
Denmark 1941 .96
Englandand Wales 1933 .73
1938 .81
Finland 1938 .96
France 1939 .90
Germany 1933 .70
1940 .98
Holland 1941 1.16
Norway 1939 .86
Sweden 1940 .79
Switzerland 1941 .90
UnitedStates(whitepersons) 1931-5 -98
1939-41 1.01
* Ratioof totalbirthsin twosuccessive
generations.
The meaningof these figuresis that certaincontingentsof man-
kind are headed for extinction,if presentattitudescontinue. This is
not sayingthatin the case of the humanspecies the principleof maxi-
38For a discussionof thesocialinfluences

makingforrestrictionin thesize
of families
seeJ.J. Spengler,FranceFacesDepopulation, DukeUniversityPress,
pp. 157et seq. Also thesameauthor's
1938,especially article"Paretoon Popula-
tion"in theQuarterly Journal of Economics
, August,1944.

192 HUMAN BIOLOGY
mumenergyfluxis on the pointof failing. For, in the firstplace, that

principlecontainsa reservation:a maximumcompatiblewith the con-
straints,
. The principletells us what is selected; what is presentedfor
selectionis determinedby otherprinciples - just as in physico-chemical
evolutionthe principlesof thermodynamics tells us which of possible
transformations actuallytake place. What transformations are possible
is determinedby the factsof stoichiometry.
Secondly,the principleof maximumenergyflux presumablycon-
tinuesto be operativeamong the remainingcomponentsof the system
of nature,if one of themdrops out. Any species,or contingentof a
species,thatsignallyfailsto keep in thecurrentof the law of maximum
energyfluxis headed for eliminationfromthe evolvingsystem. Thus
the law is preserved.
That such eliminationof species occurs from time to time is a
familiarfact engravedin the earth's fossil record,thoughthe precise
cause of these disappearancesmay remain unknown. One conjecture
is that, in certaincases at least, the extinctionwas due to a certain
fatal driftin the cumulativeinheritanceof certaincharacteristics from
generationto generationby whathas been termedorthogenesis.Wheth-
er physiologicallyspeaking orthogenesisis fact or fictionis a matter
of dispute,but the exosomaticevolutionof the human species is in-
disputablysubject to orthogenesis.Knowledge breeds knowledge,and
withpresent-daymethodsof recording,thismeans unceasingaccumula-
tion of knowledgeand of the technicalskillsbased upon it.40 But in a
way hardlydreamedof by Tennyson,it is truethat "knowledgecomes,
but wisdom lingers,"if by wisdom we understandthat adjustmentof
action to ends whichis for the good of the species. It is preciselythis
thathas gone awry in the schemesof men: The receptorsand eťřectors
have been perfectedto a nicety;but the developmentof the adjusters
has lagged so far behind,that the resultantof our effortshas actually
been reversed. From preservationof life we have turnedto the de-
40This of knowledge has beenmuchexaltedbyA. Korzibski( Man-

property
hoodof Humanity, Dutton,1921) who seemedto be undertheimpression that
he had discovered
a newprinciple. The idea was of coursefar fromnew. It
hadforexample beenclearlysetforth byW. Ostwaldin Energetische Grundlagen
der Kulturwissenschaften
, 1909,pp. 121, 122. Korzibski'scharacterization
of
manon thisaccount, as "timebinder,"can hardlybe praisedeitheras a happy
choiceof termsor as altogether
settingmanapartfromotherspecies.

structionof life; and fromexpansionof the human race we have, in

some of themostadvancedcommunities, turnedto its curtailment.
If certaincontingentsof the humanspecies are to be preservedand
perpetuated,a revisionof prevailingvaluationswill be necessary. As
R. S. Lillie has remarked,"The avoidance of useless conflict,and the
subordinationof individualintereststo the integratedwhole,whichin-
cludes the individual,would seem to be a rationalaim for conscious
beings."41In particular,as procreation,fromthe essentiallyinstinctive
drive,is passing into the domain of deliberatelycontrolledactions,it
becomesnecessarythatthe adjustmentof reproductionbe equally con-
trolledin accordancewiththeneeds of the species. This may be difficult
to accomplish.For, as Whelptonhas pointedout, "The average size of
planned familieshas been substantiallysmallerthan that of unplanned
familiesin the past and will be smallerin the future,thoughperhaps
in lesser degree. Any tendencyfor the average size of plannedfamilies
to increasein the futurewill be more than offsetby the increasein the
relativefrequencyof planned familieswhichwill result fromthe con-
tinuingspread throughthe populationof informationabout effective
methodsof contraception."42
We findourselvesat a most remarkableepoch in history.As never
before,man lookingback contemplatesa vista of miraculousprogress;
and as neverbeforehe findshimselfcastingsearchingglances into the
future.Our gratification in beingat once spectatorsand participators
in
a captivatingdramais considerablydampedby the factthattheprospect
does not please.. In the solutionof scientific
and technicalproblemswe
have achieved brilliantresults. Social problems,as conceivedbroadly
fromthepointof view of the species,we have bungled.
To us, at this criticalperiod,the two-foldthreatto the very exist-

ence of large sectionsof the human race dominatesour immediatein-
terestin world history.In the longerview all this is but an episode in
the great drama of the evolutionof the earthand its inhabitants.The
41R. S. Lillie,The
PsychicFactorsin LivingOrganisms, Philosophy of
Science
, 1943,vol. 10,p. 270. Similarreflections
also are developed,
especially
in theclosingchapters,
in myElements of PhysicalBiology(1925).
42P. K. War and theBirthRate. Paper presented beforethe
Whelpton,
American PublicHealthAssociation,Oct.2, 1944.

194 HUMAN BIOLOGY
larger scientificoutlookaspires to an objectivesurveyof eventsquite

aside fromany peculiarrelationwhichtheymay have to humanaffairs.
It is withthis broaderconceptionthatwe have viewedthe evolution
of life on our planetas the changingdistribution of matteramong the
various organic species and their inorganic environment.Whatever
line the developmentof thisbranchof sciencemay ultimatelytake,this
can be confidently asserted,thatno treatment can be effective
whichfails
to give a prominentplace to the competitionfor substanceand energy
amongthe diverseorganicspecies,in relationto the characteristic physi-
cal and dynamicalpropertieswhich the individualorganismpossesses
for wagingthe competitivestruggle.We may furtherset it down that
the formulation of the law of evolutionmustunquestionably be made in
termsof the evolvingsystemas a whole; that it cannot be adequately
expressedby referenceto onlyone component,such as a singlespecies.
It is thesystemas a whole,underthefloodof lightenergyreceivedfrom
the sun, that evolves. From our experiencein the disciplinesof the
physicsof changeof stateand physicalchemistry we knowthatit would
be quite out of the question to formulatethe law of evolutionof a
physico-chemical systemby referenceto one of the componentsalone.
No more can we expect to express the basic directionalprincipleof
organicevolutionin termsof the "organization"or any otherproperty
or assemblageof propertiesof individualspecies consideredseparately.
It is in thecollectiveactivitiesand effectsof theorganismsthatwe must
look for an indicationof the directionof evolution. These collective
effectstendto maximize,on the one hand,the energyintakeof organic
naturefromthe sun, and on the other,the outgo of freeenergyby dis-
sipative processes in living and in decayingdead organisms. The net
effectis to maximizein this sense the energyflux throughthe system
of organic nature.
Such, it seems, is the shape of the basic principledefiningthe
directionof organicevolution.


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THE LAW OF EVOLUTION AS A MAXIMAL PRINCIPLE

Author(s): ALFRED J. LOTKA

This content downloaded from 130.63.180.147 on Tue, 22 Oct 2013 17:26:30 PM

THE LAW OF EVOLUTION AS A

| NDOUBTEDLY fundamentalto the conceptof evolution

This content downloaded from 130.63.180.147 on Tue, 22 Oct 2013 17:26:30 PM

In a certainsense it can here be said thatthe pack of cards, in the

This content downloaded from 130.63.180.147 on Tue, 22 Oct 2013 17:26:30 PM

ticallybecause we deliberatelyshufflein such manneras to have only

This content downloaded from 130.63.180.147 on Tue, 22 Oct 2013 17:26:30 PM

cess, as the physicistuses the words, is associated with decrease in

This content downloaded from 130.63.180.147 on Tue, 22 Oct 2013 17:26:30 PM

constraints)or arbitrarilyupon operationspermitted.Just as the tri-

This content downloaded from 130.63.180.147 on Tue, 22 Oct 2013 17:26:30 PM

servationthatevolutionin a large numberof cases has proceededalong

This content downloaded from 130.63.180.147 on Tue, 22 Oct 2013 17:26:30 PM

peculiar type,competingfor the substancesand energiesrequisitefor

This content downloaded from 130.63.180.147 on Tue, 22 Oct 2013 17:26:30 PM

technicaltermin physics,of the dimensionsof an energy,and Volterra's

Everylivingthingis a sortof imperialist, seekingto transform

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Problemsof distribution are not new to the physicalsciences. The

This content downloaded from 130.63.180.147 on Tue, 22 Oct 2013 17:26:30 PM

crystalgerm,thatis todaystillone of theunsolvedproblemsof science.14

This content downloaded from 130.63.180.147 on Tue, 22 Oct 2013 17:26:30 PM

pect the law of organicevolutionto be primarilyexpressed) will neces-

15Thatis, forall theendstatemayneverbe

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dynamics- - elements,compounds,"phases"- are strictlyhomogeneous,

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wards a stationarystatemaintainedwithcontinuousexpenditureof free

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abilityof theseencounters,and the less the energyexpendedin propor-

19That of a one-to-one betweencertainfeatures

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For since these transformers possess a fund of free energythey can

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may be expected to move have been given by the writerelsewhere.21

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the processes under discussion,and thus enables us to select the ap-

28For detailsthereaderis referred

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refinements and extensions.And,second,thatit is well to be verycau-

34As to this,it is admitted thatbothlogicallyand fordidacticreasonsit is

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tion of the dissipativeprocess. As soon as we franklyaccept the fact

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opportunity furnishedforsuitably constituted organisms to enlarge

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"It is notlawfulto inferimmediately thatevolution tendsthus

30One is inclinedhereto give at least qualifiedassentto the sayingof

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plied mainly- but with one highlysignificantexception - by the find-

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crease in the productivity of humanlabor may,therefore,veryproperly

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this serious drawback: While the necessitiesof life, fromtheirvery