ForestScience,Vol 39, No 3, pp 594-616

Analysisof Growth Equations

Bores ZEIDE

ABSTRACT. Growthof plantsresultsfromtwo opposing factors:the intrinsictendencytowardun-

limitedincrease(bioticpotential)andrestraintsimposedby environmental resistanceand
ag/ng.The expansiontendencyprevailsin the beginningof a tree's life, while growth
declinebecomesprominenttowardthe end.The existinggrowthequations canbe trans-
formed(by differentiation,decomposition intothe divisioncomponents, andtakinglog-
arithms)sothatthe components thatcorrespond to thesetwo factorsare exposed.This
transformationrevealstwo basicformsintrinsicin mostof the analyzedequations.Their
commonfeatureis that growthexpansionis proportionalto currenttree size. Growth
declineof individual
trees appearsto be morevariableandcanbe renderedwith equal
accuracyby a variety of expressions.This may reflect that a greater numberof factors
hindergrowth:scarcityof resources,competition,
reproduction,diseases,herbivory,
disturbances,
etc. Consequently,the growthpath is inherentlyimpreciseand canbe
viewedas a wide valleyrather thana singleline. This analysislaidgroundworkfor the
classification
of knownequations andmadepossiblethe discoveryof a promisingnew
equationform. FOR.SCL 39(3):594-616.
ADDITIONAL KEYWORDS.Basicequationforms,classification of growthequations,com-
ponentsof growth,decomposition of equations,exponentialincrease.

ROWTH
EQUATIONS
DESCRIBE
THE
CHANGE
INSIZE
OFANORGANISM
ora
population
withage.Biological
growth,
theoutcome
ofnumerous
and
enormously
complexprocesses,appearsremarkablysimple,particularly
for trees. As we combinemore and more similartrees, the increasein their size
followsanever smoothersigmoid curve.In thebeginning the curveis concave up,
whilein later life it becomesconvex.Althoughgrowthrespondsto environmental
trendsandfluctuations, this long-termpatternremainssurprisingly stable.
Many equationshavebeenproposedto describeplantgrowth. Kiviste (1988)
described75 of themin a comprehensive two-volumemonograph. Althoughonly
a few haveprovenuseful,probablyno biologistbelievesthatoneequationwould
suitallgrowthprocesses. Thisseemsto be a beliefpeculiarto biology.A physicist
wouldnot usemorethanoneequationto describe,for instance,the fallof a body
in a vacuum."If physicshasits laws,biologyhasitsvariety"(Dover1988,p. 623).
The varietyof existinggrowthequations bringsup a numberof questions. Are
there any relationships amongthe equations? Is it possibleto reducethem to a
smallnumberof basicforms?How canthe adequacy or inadequacy of anequation
be interpretedso as to contributeto our understanding of nature?
Besidessatisfyingour intellectualcuriosity,answeringthese questionsmay
solvemanypracticalproblems.For example,an optimalrotationage extracted
from a more accuratepictureof foreststandgrowthcoulddifferby 10 or more
yearsfromthe agecomputedusingtraditionalmethods(suchasyieldtables).This
meansthat just by doingmore accuratecalculations, we can utilize forest re-

594/FORES'•S•
sourcesmore efficiently.Anotherimportantareaof application is the detectionof
changesin the environment,be it negative,suchas stress,or positive,suchas
fertilizationor thinning.These changescanbe revealedby comparing the actual
growthin an alteredenvironment withthe growthpredictedby reliableequations
from the data of growthprior to the change.
Althoughtheseproblemsare not new, few wouldclaimthat they are solved.
Therefore,giventheirtheoretical andpractical
implications,
I believethatfurther
analysisof the existingequationscouldcontributeto a better understanding of
tree growth.

GROWTH EQUATIONS AND

COMPUTER MODELING

Not everyonesharesthe beliefthatgrowthequations meritfurtherconsideration.

Someview the curve-fittingapproachas an ossifiedremnantof the precomputer
past and doubtwhetherit is worthwhileto studytheseold-fashioned formsof
modeling.Now we havemuchmorecomprehensive andflexiblemethodsof com-
puter modelingthat utilizesuchmethodsas dynamicprogramming, difference-
basedequations, andneuralnetworks.There are alsoconceptual objections to
usinggrowthequations. Hustonet al. (1988)believethatgrowthequations andall
modelsthat dealwith a population as a whole("aggregatedlarge-scalemodels")
arebasedonunrealistic ecologicalassumptions, glossoverecological mechanisms
andindividualvariability,andignorereality (in particular,feedbackinteractions).
These authorsview it as unacceptable to "combinemanyindividuals andassume
that they canbe describedby a singlevariable,suchas population size. This
procedureviolatesthe biological principlethat eachindividual is different,with
behaviorand physiologythat result from a uniquecombination of geneticand
environmentalinfluences"(Hustoneta. 1988, p. 682). Accordingto these au-
thors, the futureof ecologybelongsto the individual-based modelingapproach,
whichwill soonproducea mechanistic understanding of ecological systems.
This beliefis not new. It hasbeen knownin philosophy for centuriesunderthe
nameof nominalism.It is basedon the exaggeration of differencesamongindi-
vidualsof the samespeciesor population andis equivalent to sayingthattheyhave
nothingin common, apartfromthe species name.Nominalism is logicallyincon-
sistent:were there nothingcommonto all individuals, they couldnot be consid-
ered as membersof the samespeciesandcombined underonename.Following
the sameline of reasoninga nominalistshoulddo awaywith the conceptsof not
onlypopulation andspeciesbut of organism aswell. An organism canbe viewed
as a population of individualcellsthatare differentin behavior,physiology, loca-
tion, and manyother characteristics. Cellsin their turn includemanydifferent
components.
Oneof the reasonsofferedby Hustonet al. (1988)in supportof individual-based
modelsis that, alongwith detailedinformation
on individual
interactions,they are
capableof producing a generalandintegralpictureof a system.Following this
suggestion,we coulddetermine,for example,temperatureof an organismby
measuring the speedandmassof eachconstituent particle(onemolecontains
6.02 ß102aparticles).It is likely,however,thatevenintheremotefuturemany
wouldpreferto usea thermometer whichdescribes
theintegralbehaviorof many
individual
particleswith a singlevariable,temperature.

AUGUST1993/595
 Experienceshowsthat alongwith differences,individuals sharemanycommon
featuresandthat it makessenseto useanaverageto characterizea population as
a whole. Both unicellularandmulticellular organismsoftenexhibitaggregatebe-
havior,whichis considered oneof the majorcharacteristics of complexadaptive
systems(Holland1992). Problemswith the individual approach are well-knownin
forest ecologyin whichindividual-based modelshave been appliedsince 1950.
Substantial researchhasrevealedthat, whilerequiringmuchmore effort, these
modelsprovidepracticallyno improvementin the accuracyof growthpredictions
as comparedwith an aggregateapproach (Alemdag1978, LarocqueandMarshall
1988). Interactionsamongindividualtrees are often so complicated that they
precludereliablepredictions.
Even if it were possible,not all our problemscouldbe solvedby tracingthe
effectsof numerousagents,detectingassorteddisturbances, andfleetingday-to-
day perturbationsin growth. There is a need to describelastingfeatures of
growth and to expresslong-termtrends, suchas aging.In studiesof overall
trends, a certain degree of rigidity is an asset rather than a liability. For this
reasongrowthequations havenot losttheir significance evenwith the adventof
computermodeling,althoughthey are no longerviewedas biological versionsof
Newton'slaws. Actually,growthequations are usedmore thanever beforebe-
cause,in addition to theirindependent role,theequations serveasbuilding blocks
for computermodels. Of course, these two approachesdo not exclude,but
complementeachother. Each is tunedto its own frequency:detailedcomputer
modelsare designedto reflectshorterwavelengths of dailychanges,whilegrowth
equations considerthe entirelifespanas onewave.
Preciselybecauseof theirwidescope,growthequations, alongwitha descrip-
tionof the change,afforda glimpseat the constancy resultingfromthe invariance
of the geneticmechanism responsible for growth.The form of an equationis a
meansto achievestabilityof parameters.The moreaccuratean equationis, the
lessvariableare its parameters.A growthequation bringstogethertwo currents,
age and size, to make explicitthe hiddeninvariancethat governstheir relation-
ship.

THE PRIMARY COMPONENTS OF GROWTH

Growth resultsfrom the interactionof two opposingforces. The positivecom-

ponent,most vividlymanifestedin expansionof an organism,representsthe
innatetendencytowardexponential multiplication.
This component is assodated
with bioticpotential,photosynthetic
activity,absorption
of nutrients,constructive
metabolism,anabolism,etc. The opposingcomponentrepresentsthe restraints
imposedby external(competition, limitedresources,respiration,andstress)and
internal(self-regulatorymechanisms, andaging)factors.Thosefactorsthat ad-
verselyaffectgrowthhavebeenreferredto asenvironmental resistance,destruc-
tive metabolism,catabolism,respiration,and so on.
Appropriately,hws or postulatesof growthare oftenformulatedin pairsthat
reflect boththe multiplicative
andlimitingcomponents. Hutchinson's (1978) two
postulatesof populationgrowthare:
1. Every livingorganism
hasarisenfromat leastoneparentin likekind(thepostulateof
parenthood);

596/FORESTSCmNCE
2. In a finitespacethereis an upperlimitto the numberof finitebeingsthatcanoccupy
or utilizethe spaceunderconsideration (the postulate
of an upperlimit).
In 1941 Medawarformulatedfive lawsof biological
growth.Whenhe returned
to the sameproblem4 decadeslater,he condensed thesefiveintotwo basiclaws
(Medawar and Medawar 1983). They are similarto Hutchinson'spostulates:
1. Fundamentally, growthis multiplicative.
That whichresultsfrom biological
growthis
itself,typically,capableof growing;
2. The relativegrowthrate is alwaysdecreasing(Minot'slaw).

The conflictbetweeninfinityimplicitin multiplicativereproduction andthe limit

imposedby finitespaceis the chiefsourceof allchangein livingbeings,including
growth.Thisconflictis the drivingforceof evolution andis crucialto understand-
ing virtuallyall biologicaland socialphenomena.Growth equationsprovidea
succinctexpression of this conflictandits resolution.

ANALYZED EQUATIONS

This studyanalyzesexistinggrowthequations, includingpopularequationssuch

as the Chapman-Richards, the Gompertz,and the logisticmodels,as well as
severallessknown,yet promising models(Table1). The structureof any other
equation canbe analyzed in a similarmanner.Polynominal-based equations were
not consideredbecausethey are devoidof any biological interpretation.
Let's briefly review the equationsstartingwith the oldest, the HossfeldIV
equationproposedfor the description of tree growthas earlyas 1822 (Peschel
1938). Despiteits age, this equationperformsremarkablywell. Accordingto
Kiviste(1988), it is the thirdmostaccurateof 31 three-parameterequations when
the three mainstandvariables(totaltree height,stem diameter,andvolume)are
considered together.Kivistefoundit to be the bestequationfor volumegrowth.
The HossfeldIV equation is almostasaccurate asthe Chapman-Richard equation,
whichdominates growthstudiesin thiscountry.The mostaccurateequations with
three (LevakovicI andIII equations) andmore (YoshidaI equation)parameters
are modifications of the Hossfeldequation.
The Gompertz(1825) equationwas designedto describeage distribution in
humanpopulations. A centurylater it was appliedas a growthmodel(Winsor
1932).The equation presentsrelativegrowthrate (theratioof incrementof size
to sizeitself,y'/y) as an elementaryexponential functionof age. Therefore,this
equation wascalled(Lairdet al. 1965),the equation of exponentialdecay.Another
characteristics featureof the Gompertzequationis that the positionof the inflec-
tionpointis controlled by onlyoneparameter,final(asymptotic) size,a. Thispoint
occurswhencurrentsizeis equalto a/e, that is, at aboutone-third(1/e = 0.3679)
of the finalsize.Nokoe(1978,p. 41) appliedthe Gompertzequationto threetree
speciesand concluded that this equation"demonstrated sufficientflexibilityto
warrantits use."The Gompertzequation wasfoundby CaustonandVenus(1981)
andmanyotherresearchers(Lairdet al. 1965,ZweifelandLasker1976, Zullinger
et al. 1984) to be more appropriatein biological work thanany other. Besides
these empiricalresults,it was deducedtheoreticallyby Medawar (1940) that
growthshouldfollowthe Gompertzmodel.

AUGUST1993/597
598• FO•Scm•cE
 The logisticequation(Verhulst1838)is probablythe mostfamousequationin
ecology.An outstanding exposition of its historyis givenby Hutchinson (1978).
The forcesthat counteractthe exponential increaseare assumedto be propor-
tionalto the squareof size.No reasonfor thischoiceof the exponentis provided
otherthanthat 2 is the next integerafter 1. It is doubtfulthat the principleof
parsimony,invokedanddiscussed in depthonthisoccasion by Hutchinson (1978),
is suffidentto precludethe application of numbersother than integers.The
inflectionpointof the logisticequationcorresponds to one-halfof finalsize. The
relativegrowthrate of the equationdeclineslinearlywith size. Severalother
assumptions inherentin the logisticequationappearto be questionable (Krebs
1985, p. 220). Despiteits untenableassumptions andlack of accuracy--itwas
found(Zeide 1989) to be the least accurateamongsigmoidequations for the
descriptionof diametergrowthof trees--the equationis still usedin research,
mostlyby zoologists (Ricklefs1979, Murtaugh1988).
The monomolecular equation,the simplestamongthe analyzedequations,is
not inflectedandthereforepresentsa rather unrealisticpictureof growth.This
equationis knownas the law of diminishing returnsin agricultureandeconomics
and as the law of massactionin chemistry.Ricker (1979) attributesthe first
biologicalapplicationof a specialform of this equationto Patter (1920), while
Richards(1969)refersto it asthe Mitscherlich formula,afterthe Germanagron-
omistwhousedit at the beginning of thiscentury.I havefoundanevenearlieruse
of thisequationin a studyof tree growthby Weber(1891).
The appealof the Bertalanffy(1957) equationlies in the intendedrigor of its
theoreticalfoundation.Bertalanffy(1957, p. 223) claimsto have succeededin
developing "a generaltheoryof growthwhichestablishes rationalquantitative
lawsof growthandindicatesthe physiological mechanisms uponwhichgrowthis
based."This theory considersanimalgrowthto be the result of the combined
actionof two opposing processes,anabolism andcatabolism. Bertalanffy(1957)
derivedhisequationfromthe assumptions, whichhe attributedto Patter (1920),
thatthe rate of anabolism is proportionalto the surfaceareaof an organism(or to
is massraisedto the powerof g/a),whilecatabolism is proportionalto the organ-
ism'smass.These assumptions definewhat he calls"the first metabolictype."
Bertalanffyalsodescribes two othertypesof metabolism andtheircorresponding
growthtypes. Ricker (1979, p. 707) questioned theseassumptions considering
themto be "fanciful speculations."Nevertheless, to honorthe presumably original
authorof thesespeculations, Rickerrefers to the Bertalanffyequationas the
Patter Growth Curve No. 2.
The originof thisconcept,however,goesdeeperthanBertalanffyandRicker
believe.It mightbeoneoffewviableideasfromtheprodigious legacyofSpencer.
He was concernedwith the question,"Why has individualgrowtha limit?"and
proposed severalanswersthatresemblestructuralandmechanical considerations
putforthby Galileoin his"Dialogues concerningtwo newsciences." In particular,
Spencer(1898, p. 151) wrote, "In similarbodies,the areasvary as the squares
of thedimensions, andthemasses varyasthecubes;it followsthattheabsorbing
surfacehas becomefour times as great, whilethe weightto be movedby the
matterabsorbed hasbecomeeighttimesasgreat."As doesBertalanffy, Spencer
restrictsthisreasoning to animals,believingthattree growthis unlimited.
The trademarkof the Chapman-Richards equationis its flexibility.Althoughthe
equationwas reportedby Mitscherlich(1919), it becameknownto American

AUGUST1993/599
researchersfrom the articlefittinglytitled "A flexiblegrowthcurve for empirical
use"by Richards(1959). This equationis valuedfor its accuracyandis usedmore
than any other functionin studiesof tree and stand growth. The Chapman-
Richardsequationwas derivedfrom the Bertalanffyequation"when limitations
imposedby its theoreticalbackground are discarded"(Richards1959, p. 291).
The differencebetweenthe equations is thatthe parameterc, restrictedto a value
of three in Bertalanffy'scase, can assumeany value in the Chapman-Richards
equation.This modification dispensed with the biological
interpretation proposed
by Bertalanffy.In his review of thispaper,RolfeA. Leary remarkedthat com-
paredwith Bertalanffy'sequation,the one by Chapman-Richards is "a giantleap
backwards from explanationto description."
It is not clearwhetherflexibilityis a desirablequalityof growthmodels.When
the numberof parametersis equalto the numberof datapoints,anyequationwill
passthrougheachpoint,thusexhibiting the ultimateflexibility.I doubtthatwe are
lookingfor thissortof flexibility.Our understanding of growthwouldbenefitlittle
froman equationthatpassivelyfollowsall datapoints.We needan equationthat
setsits own, andhopefullycorrect,paththroughall datapoints,an equationthat
exposesa growthtrend in the maze of dataandseparatesthe essentialfrom the
accidental.The law of physicsthat statesthat the distancecoveredby a falling
bodyin a vacuumis proportional to the squareof the time of fallrigidlyadheres
to the exponentof 2. This "rigidity"is the essenceof the law. An equationwith
a variableexponentwouldcertainlyfit empiricaldatabetter thanthislaw. Yet, this
flexibleequationwouldbe a meaningless formularather than a cornerstoneof
science.
Flexibilitydependsonthe numberof parametersin anequation,andthereis no
reasonto expectthat the Chapman-Richards equationwouldbe moreflexiblethan
any other equationwith three parameters.Still, this equationis set apart from
othersby its computational properties.Ratkowsky(1983,pp. 83-84) showedthat
thisequationis "the onlymodelthat hasan unacceptable intrinsicnonlinearityas
the solutionlocusdepartssignificantly
froma hyperplane."Thispropertyleadsto
so much instabilityin parameter estimatesthat it makes them useless.In his
personalcommunication (of March23, 1992), RichardWoolions writesthatpa-
rmeter estimates ofthe Chapman-Richards equationareworthless, especiallyfor
largedatasets,becauseconvergence is achievedby "meansof a fait accompli."
The LevakovicI and III (1935) equationsare modifications of the Hossfeld
equation.They were publishedover halfa centuryago in Serbianandare little
knownin othercountries.The LevakovicIII equationmaylookstrangebecause
it is unclearwhyageshouldbe squared.Althoughsquaring doesnotcontributeto
the accuracyof the logisticequation,it workswell for the Levakovicequation:
Kiviste(1988)foundit to be the mostaccurateamongall three-parameter equa-
tions that he investigated.The LevakovicI equationis one of the best four-
parameterequations.
Korfs equationwasproposedin 1939(Kiviste1988)in Czechoslovakia andhas
beenrediscovered severaltimes,inparticularby Lundqvist(1957).He, aswellas
subsequentresearchers(Stage 1963, Brewer et al. 1985), appliedit to model
heightgrowthof foreststandswith moderatesuccess.Zarnovican (1979), who
alsousedKorfs equationin a studyof heightgrowth,was better versedin the
literatureandcitedthreepapersby Korf, including hisoriginalpaperof 1939. This
equationis especiallysuitablefor the descriptionof diametergrowthof a fixed

600/F9•SCmNCE
numberof trees. Zeide(1972, 1975)andZeideet al. (1972)foundthatthe relative
growthrate of diameteris a power,ratherthananexponential functionof ageand
by integration arrivedat the Korf equation.Usingaveragegrowthof thousands of
stem analysesof differentspeciesfrom differentlocations,it was shownthat the
Korf equationis substantially moreaccuratethanothergrowthequations(Zeide
1989). Its standarderror of estimatewas 2.1, 2.3, 3.4, and4.8 timeslessthanthe
errors of the Chapman-Richards, Weibull,Gompertz,andlogisticequations,re-
spectively. A specialformof thisequation withc = i wasindependently proposed
by Terazakiin 1915 (citedin Peschel1938),Johnson (1935), andSchumacher
(1939).
Originallyintendedto describea probab•itydistribution, the Weibullequation
has provento be a goodempiricalmodelof tree growth. Yang et al. (1978)
reportedthat this equationis more accuratethan the Gompertzor Bertalanffy
equations. No comparison withthe Chapman-Richards equation wasgivenin their
work. WhenDolph(1991) compared theseequations in the processof construc-
tionof site indexcurvesfor red fir, he foundthatthe Weibullequationwasmore
accuratethan the Chapman-Richards equation.Unlike all other functions,the
Weibullequationpresentsthe increaseof growthas a powerfunctionof age. It
occupies the fourthplacein Kiviste's(1988)rankingof three-parameter equations
and is particularlygoodfor modelingdiametergrowthof stands.Zeide (1989)
foundthatWeibull'sequationislessaccuratethanthe Koff andChapman-Richards
equations.
The YoshidaI equation,proposedin 1928 (Peschel1938), is anothermodifi-
cationof theHossfeld equation. Kiviste(1988)foundthatthisequation is themost
accurateamong21 four-parameter equations. In additionto thoseof the Hossfeld
equation,the YoshidaI equationcontains anadditiveterm (parameterc in Table
1) that representsinitialtree size.Becausethissizeis negligible,sois the term.
The Sloboda(1971)equationdiffersfromthe Gompertzequationby an addi-
tionalparameterd. The presenceof thisparameterprobably is responsiblefor the
greateraccuracy of the Sloboda equation.According to Kiviste(1988),it is the
secondbest four-parameter equation.

DECOMPOSITION OF EQUATIONS

In mostcases,growthequations are usedin anintegralformwhichdescribes the

accumulated sizeof anorganism. The formof the equations affectsthe perception
of differences or similarities
amongthem.Usingthe integralform, Kiviste(1988)
dividedgrowthequationsinto sevenclasses.He placedthe logisticand Korf
equations in the sameclassof exponential functions.The Bertalanffy, Weibull,and
Chapman-Richards equations were classified
as Mitscherlich functions.The Korf
andLevakovicequations were locatedin two differentclasses.
It is easierto understandthe processof growthandthe structureof growth
equations whenwe consider themin the differential
formwiththe currentincre-
ment,y', as the dependent variableandtree age, t, as the independent variable
(Table1). In thisformgrowthequations canbe decomposed intotwo components
that representgrowthexpansionand decline.The expansioncomponentde-
scribesthe multiplication tendencyandis responsible for an increasein the in-

AUGUST 1993/ 601

crementwith age or keepingthe incrementconstant.The componentof growth
declinecausesthe incrementto decrease.Theseoppositeeffectson growthallow
one to detect the componentsand decomposean equation.
The componentthat expressesgrowthdeclineis notjust an optional"modifier"
of exponentialincrease,asis sometimes suggested. Thiscomponent is oneof the
two irreplaceable parts of any modelof biological growth.Equationscontaining
onlyonecomponent,suchas Malthus'(1798)law of population increase(geomet-
ric progression),cannotbe considered completegrowthequations. Thislaw (pro-
posedwith a balancing conjectureaboutthe arithmeticprogression of the means
of subsistence) servedas a progenitorandcatalystof growthequationsbecause
many sdentists, includingits author, felt its incompleteness. For this reason
Verhulst(1838)augmented the law (in its differential
form)witha subtractorthat
offsetsthe multiplicative component.The sameopposition is achievedin the
Gompertzequationby division.
Thesetwo example(VerhulstandGompertzequations) are typicalof allgrowth
equations.Components with positiveparametersare connected by subtraction or
divisionbut not by additionor multiplication.

SUBTRACTION

All the investigatedequationscan be presentedin the differentialform as a

differenceof two components.For example,usingthe integralformof the mono-
molecular
equation
(Table1), we canpresent
thetermace-otasa - y (a, b, c,
andd throughout thispaperare constantparameters).Substituting
this resultin
the differentialform of the sameequation,onewouldobtain:
y' = ab - by (1)
The expansioncomponentin this equationis a positiveconstant(ab), while the
declinecomponent (by)is proportional
to the size,y. In thispapersizerefersto
nondiminishing
tree or standvariables,suchas height,diameter,or volume.
As anotherexampleof decomposition, theintegralformof the Weibullequation
can be written as:

exp(-bD = i - y/a (2)

Substitution in the differential form results in

y' = abct
c- • - bcyt
•- • (3)
Becausethe constantsare positive,the first term causesincreaseof the incre-
ment (y'), while the secondterm contributesto its decrease.
As a form of connectionof components,subtractionhas received the most
attentionin previousaccountsof equationstructure.Our intimatefamiliaritywith
thissimpleoperation(afterall, we balanceour checkbooks by subtraction,notby
divisionor exponentiation)probablyplayssomerole in this preference.Berta-
lanffy (1957, p. 223) consideredgrowth as the result of "a counteractionof
synthesisanddestruction,of the anabolism andcatabolism of the buildingmate-
rialsof the body."He viewedthiscounteraction exclusivelyin termsof subtrac-
tion.In a seriesofpublications,
Savageau (1979,1980)andhisstudents presented
the mostthoroughandconsistentdevelopmentof this approach.He produceda
generalized growthequationthat "is not simplyanotherempiricallyderivedfor-

602/FORESTSCII•-NCE
mula but is baseduponthe nature of the elementalmechanisms in synergistic
systems"(Savageau 1979,p. 5416).This equationis constructed asthe difference
of two terms. In orderto presentthe Gompertzequationas a particularcaseof
hisgeneralized equation,Savageau admitsthe existenceof two (andmore)dom-
inantprocessesof growthwithinthe samesystem.The technicaland linguistic
difficulties(dominant,after all, meansthe most influential,prevailing,and, there-
fore, unique)with this approachcan be avoidedby decomposing the Gompertz
equationinto divisioncomponents.

DIVISION

Notwithstanding
the familiarity
andsimplicity
of subtraction,
it is not the only
possibilityfor decompositionof growthequations.Division,whichis the subtrac-
tion of logarithms,is an equallyvalidoperation.Many, if not the majorityof all
biologicalphenomena are multiplicative
in natureratherthanadditive.The com-
ponentsof the Gompertzequationcanbe presentedequallywell as dividend(y)
anddivisor(e½5.
For theGompertz
equation
andmanyothers,components
of the
divisionmethodof decomposition are simplerthanthoseof subtraction(Table 1).
For example,eachof the divisioncomponents of the Korf equation(obtainedby
substitutingthe right sideof the integralform into the differentialform) contain
only one of the equation'svariables,while the subtractioncomponents contain
both.
Decomposition of someequationsdependsnot onlyon their form, but alsoon
the valuesof parameters.In the $1oboda
equation,for example,whenthe param-
eterd > 0, the termta- • increases
the increment
in thecourseof timeand,
therefore,is a part of the expansion
component. When0 < d < i the sameterm
playsthe oppositerole andbecomesa part of the declinecomponent.
Decomposition by divisionallowsone to further simplifyequationsby taking
logarithms.In many casesthe equationsbecomelinear. The three steps de-
scribedabove(differentiation, decompositionintodivisioncomponents,andtaking
logarithms)transformequationsinto a form that allowsone to do the following:
1. Simplifythe equationsandlinearizemostof them.
2. Homogenizetheir variance.
3. Applywell-developed methodsfor the investigation
of linearequations.
4. Facilitatethe designof new equations.
5. Exposebasicformsor familiesof equations.

The last point is pursuedin somedetailbelow.

BASIC FORMS OF EQUATIONS

The describedtransformation
revealstwo basicformsbehindmostof the analyzed
equations.To presenttheseformsmore vividly,the equationsare rewritten to
simplifythe notationof the constantparameters.All interceptsare designated as
k. Constantsof size,y (or In(y))are denotedbyp, whilethoseof age, t andIn(t),
are expressedby q (Table2). For example,for the Korf equationthe parameter
k in Table 2 is equalto In(bc)in termsof the parametersof Table 1. Similarly,for
the sameequation p = 1 andq = -(c + 1).

AUGUST1993/603
 TABLE 2.

Growthequationsin uniformnotation.k > 0, p > 0, andq < 0 = parameters

of equations.
Equationname Logarithmof differentialform

HossfeldIV ln(y') = k + 2In(y) + qln(t)

Gornpertz ln(y') = k + In(y) + qt
Logistic ln(y') = k + 2In(y) + qt
Monornolecular ln(y') = k + qt
Bertalanffy ln(y') = k + (g/3)ln(y) + qt
Chapman-Richards ln(y') = k + pin(y) + qt
LevakovicI ln(y') = k + pin(y) + qln(t)
LevakovicIII ln(y') = k + pin(y) - 3In(t)
Korf ln(y') = k + In(y) + qln(t)
Weibull ln(y')= k + pin(t) + qt•+•
YoshidaI ln(y') = k + 21n(y- c) + qln(t)
Slobode
if d > 1 ln(y')= k + [In(y)+ (d - 1)In(t)]+ qt•
if0<d< 1 ln(y')= k + In(y)+ [(d- 1)In(t)]+ q•

It becomestransparentthat all the equations,exceptWeibull's,are particular

casesof the two followingforms:
ln(y')= k + pin(y)+ qln(t) or y' = k•yPtq (4)
ln(y')= k + pln(y)+ qt or y' = klyPe qt (5)
wherep>0, q<0, andk• = ek.
In bothformsthe expansion component is proportional
to In(y)or, in the antilog
form, is a powerfunctionof size. The formsdifferin the way the declinecom-
ponentis presented.In Equation(4) it is proportionalto the logarithmof age, t.
This form will be referred to as the LT-decline or LTD form. The decline com-
ponentof Equation(5) is directlyproportionalto age,t. Accordingly,
Equation(5)
canbe calledthe TD (T-decline)form.In the integralform,the declinecomponent
is either a power functionor an exponentialfunctionof age.
The LTD form includesthe HossfeldIV, LevakovicI andIII, Korf, andYoshida
I equations.Eachof theseequationscanbe derivedfrom the generalequation
form [Equation(4)] whenits parametersassumea particularvalue.Thus, the
LevakovicIII equationis distinguished
by q = 3 andthe Korf equation byp = 1.
The peculiarityof the YoshidaI equation(Table2) is that its dependent variable
is the differencebetweenthe currentsizeandthe initialsize(c = y(0)). The same
numberof equations (Gompertz,logistic,monomolecular, Bertalarfffy,andChap-
man-Richards) belongto the TD form. The Slobodeequationcanbe viewedas a
hybridbetween the two forms.
Dependingon the valuesof p andq, severaldistinctintegralequationscanbe
obtainedfromthe sameequationform. The Korf equationresultsfromintegration
of the LTD formwhenp = 1 andq • - 1. The LevakovicI followsfromthe same
equationwhenp > 1 andq < - 1. The selectionof a particularintegralequation
forcesits parametersintoa certainrange.The parameters c (= - 1/(1 - p)) and
d (= -q - 1) in the LevakovicI equation,for instance,mustbe greaterthan
zero.

The transformed
equations
revealquitedifferentandsimplerrelationships
than

604/FORESTSCIENCE
those of Kiviste's classification.
Thus, despitethe differenceof their integral
forms,HossfeldIV andKorf equations are varietiesof the samebasicform(Table
2). On the otherhand,differentiation
showsthat the outwardsimilaritybetween
the Chapman-Richard andWeibullequationsis misleading.

COMPARISON OF THE BASIC EQUATION FORMS

The precedinganalysisbringsforth a question:whichof the two basicformsis
more accurate?The answerprovidedin thispapershouldbe consideredas pre-
'hminary
becauseit was obtainedfrom one data set.

MATERIALS

Probablythe best-knowndataon tree growthare the measurements of Norway

spruce(Picea abies[L.] Karst) publishedby Guttenbergin 1915. This set is
considereda touchstoneof tree growthandhas been usedrepeatedlyby many
researchers(for example,Assmann1970, Sloboda1971, Zeideet al. 1972, Zeide
1989). The data containmeasurementsof 107 average-sizetrees from five site
classesselectedfrom healthy,fully stockedstandsgrowingin the Alps. Gutten-
berg (1915) provided actual data for each tree as well as corrected (hand-
smoothed)averagesfrom 10 to 150 yr by site classes.The numberof analyzed
trees andtheir averagesize at age 50 are shownin Table 3. Despitethe differ-
encesin locationandspecies,the growthpatternof thesetrees is s'm•ilarto that
of severalspeciesin the westernUnited States(Zeide 1989).
Becausestemdiameterwasmeasuredat the heightof 1.3 m aboveground, this
variablewasregressedonthe agesincethe tree reachedthisheight.Thisagecan
be easilycalculated
for eachtree andsite classfromheightgrowthdata(Table3).
Tree heightand volumewere regressedon the age at stem base.

DATA SCREENING

Severaltrees showedan erraticpatternof growth.Unlikethe majorityof trees,

their incrementjumpedup anddownwithoutapparentpattern. Sometimesout-
liersmay providevaluableinformationandevenleadto new discoveries. Unfor-

TABLE 3.

Averagesize of Norway spruce(Piceaabies[L.] Karst) trees at 50 yr, number

of trees, andage at whichtrees reachedthe heightof 1.3 m by site class.
Site class

Variable 1 2 3 4 5

Height, m 20.0 16.4 12.2 9.4 6.1

Diameter, cm 24.4 20.0 15.1 12.6 9.3
Volume,m3/1000 433 245 104 56 23
Number of trees 21 37 20 21 8
Age at 1.3 m 9.5 10.0 12.0 12.4 20.0

AUGUST 1993/605
tunarely,this doesnot happenoften. Usually,they obscurethe patternwithout
revealinganythingmeaningful. In orderto excludeanybiasfor or againstanalyzed
equations,the outlierswere detectedusinga completelydifferentequation(a
polynomial). Currentannualincrement,i (= y'), wasusedto computestatistics
of the followingequationfor eachtree:
i= a + bt + cF (6)
where t is age anda, b, andc are constants.
The choiceof the equationdidnot
affectthe detectionof outliers;the sametrees showa poor fit usingany other
three-parameter
equation.
Seventreeswiththe smallest
adjusted
R2 for each
variablewere consideredoutliersandwere deletedfrom the data set. Any mea-
sure of fit clearlyshowedthat thesetrees were not typicalas is seenfrom the
distribution
of R2 for Equation
(6) describing
diametergrowth(Table4). The
deletedtreeshadR2 lessthan0.32forheight,0.60fordiameter,
and0.64for
volume.The remainingdata set containedexactly 100 trees.
ENVIRONMENTALCHANGESAND TREE GROWTH

Amongotherfactors,tree growthis affectedby long-termenvironmental change.

This factor,however,is not reflectedby the growthequationsconsidered above
thatpresenttree growth(increment)asa functionof tree sizeandage.To deride
whether growth equationsshouldcontaina term responsiblefor environmental
changeit is necessaryto investigateits effecton tree growth.
The analyzedtrees were cut at agesrangingfrom60 to 150 yr. This makesit
possibleto dividethem,withineachsiteclass,intotwo groupscontaining younger
and older trees. The differencebetweenmean ages betweenthe groupswas
40-60 yr. For eachgroupmeanheight,diameter,andvolumewere calculated at
the age of 50 yr.
The results(Table 5) showsthat the size of 50-yr-oldtrees in both groupsis
practicallyidentical.The differencesare neither significant
nor consistent.In
someclassesthe trees cutat an olderagewere slightlybiggerat 50 yr, whilein
othersthey were smallerthanthe trees cutat a youngerage.Theseresultsallow
one to concludethat duringthe 40-60 yr prior to Guttenberg'sanalysis,the
environmentdid not changeenoughto affectgrowthof the investigatedtrees.

TABLE 4.

Distribution
ofR2 forEquation
(6) describing
diameter
growthoftrees.
Adjusted
R2 Frequency Cumulated
frequency
0.05 2 2
0.15 1 3
0.25 0 3
O.35 0 3
0.45 3 6
0.55 1 7
0.65 3 10
0.75 16 26
0.85 23 49
0.95 58 107

606/FoP, mTSC•'•CE
 TABLE 5.

Height,diameter,andvolumeof average50-yr-oldtrees in youngerandolder

groupsby site class.SD = standarddeviation.
Height(m) Diameter(cm) Volume(dins)
Site No. of Group
class trees age Mean SD Mean SD Mean SD

i 8 80.0 19.9 1.6 23.0 2.5 402.5 92.2

i 12 139.2 20.1 1.6 25.1 4.2 453.3 169.0
2 26 92.3 17.2 1.5 19.9 2.8 258.7 85.1
2 9 148.9 15.5 2.0 18.6 3.6 206.4 81.3
3 9 108.9 12.9 1.1 14.7 1.2 102.3 21.2
3 10 150.0 11.1 2.2 14.7 3.5 96.4 51.4
4 9 113.3 10.0 0.9 13.5 2.6 72.4 30.4
4 11 150.0 8.9 1.1 12.1 2.1 52.4 20.5
5 3 116.7 6.1 1.8 8.2 0.6 18.3 5.0
5 5 150.0 5.9 2.3 8.5 2.6 22.0 11.9

ACCURACYOF THE BASICEQUATIONFORMS

The two basicequationforms, LTD andTD, [Equations(4) and(5)] were com-

paredusingGuttenberg'sdatafor three variables(totalheight,stem diameterat
breastheight,andstemvolume).The equationformswere fittedto the following
three typesof data:(1) eachtree separately,(2) all tressof the samesite class
(referredto in Table 6 as pooleddata), and(3) smoothedaveragegrowthseries
providedby Guttenbergfor each site class(referred to in Table 6 as average
data). Comparisons were madeusingthe standarderror of estimate(Table 6).
For individualtrees the meanof the errors, foundseparatelyfor eachof the 100
trees,wascalculated.
Otherstatistics
(suchasR2 or Mallows'
Cpstatistic)
pro-
duced similar results.
The results (Table 6) show that:
1. The accuracyof the equationform dependson the datatype.
2. For individualtress, both equationforms are equallyaccuratefor any tree variable
(diameter,height, volume).
3. When trees are pooledby site class,both equationforms providean identicalfit for
heightandvolume.LTD is more accuratefor diameterin all site classes.
4. LTD is more accuratethanTD for averagegrowthseriesdatain all classesandfor all
tree variables.In the integralform, this differenceis likely to be more substantial.
Thus, the standarderror of estimateof Chapman-Richards equation(an integralform
of TD) is twice as large as that of Korfs equation(an integralform of LTD) for the
same data (Zeide 1989).
5. The shapeof growthcurvesdependson the data type, as is evidentfrom the sub-
stantialdifferencesin parametersof the same equation.For example,when LTD
[Equation(4)] wasappliedto volumethe parameterq wasequalto - 3.05, - 1.55, and
- 2.29 for individual,
pooled,andaveragegroupings of the samedata, respectively.
These differencesindicatethat even in homogeneous tree groups(trees of average
sizestratified
by siteclass)thegrowthofindividual
treescannotberepresented
by the
averagegrowthof the group.
6. If we are analyzing
the processoftree growth,it is saferto investigate
individual
tress
andavoidarbitrarytree groupings,averaging,andother kindsof datamanipulation.

AUGUST1993/ 607
 TABLE 6.

Standarderrors of estimatefor the basicequationforms, LTD andTD

[Equations(4) and (5)] by datatype andvariable(height,diameter,
and volume).

Height Diameter Volume

Tree
•oup LTD TD LTD TD LTD TD

Mean results for individual trees

Single 0.15 0.16 0.15 0.15 0.15 0.15
Pooleddataby site dasses
SC = 1 0.21 0.21 0.26 0.32 0.24 0.23
SC = 2 0.24 0.24 0.27 0.33 0.26 0.25
SC = 3 0.24 0.20 0.22 0.27 0.25 0.21
SC = 4 0.24 0.26 0.25 0.33 0.25 0.25
SC = 5 0.32 0.28 0.27 0.36 0.29 0.29

SC Mean 0.25 0.24 0.25 0.32 0.26 0.25

Averagedataby site classes

SC = 1 0.06 0.11 0.06 0.08 0.10 0.13
SC = 2 0.07 0.11 0.03 0.05 0.08 0.10
SC = 3 0.05 0.07 0.03 0.04 0.02 0.03
SC = 4 0.05 0.06 0.03 0.05 0.06 0.06
SC = 5 0.03 0.04 0.03 0.03 0.01 0.02

SC Mean 0.05 0.08 0.04 0.05 0.05 0.07

COMPARISON OF EQUATIONS WITH ALL

COMBINATIONS OF TREE SIZE, AGE, AND
THEIR LOGARITHMS

The LTD form clearlydiffersfrom TD. In the TD form the carrier of growth
declineis age, whilein the LTD formthe samecomponent is representedby the
logarithmof age.Yet, it wasfoundthatthe accuracy of theseformsis equalwhen
theyare appliedto individual trees. If thesetwoformscannotbe distinguished by
accuracy,does the choiceof an equationmatter?Wouldother combinations of
variablesbe equallysuccessfulin growthprediction?
To answerthesequestions,the accuracyof growthpredictionwastestedfor 15
differentialequationsin whichindependentvariablesincludedall possiblelinear
combinations of tree size, age, andtheir logarithms.Limiteddegreesof freedom
precludedcalculation of equationswith 4 variablesfor the 2 out of 100 trees with
the shortestlife span(60 YD. Calculations were performedfor datapooledfrom
all 98 remainingtrees andby site class(Table 7) as well as for individualtrees
(Table 8).
Standarderrors of estimatefor analyzedequationswere calculatedfor each
tree. Their meanvaluesare givenin Table8. Most of thesemeanshada standard
error of 0.01. Onlyfourerrorsfor volumeincrementwere greaterthan0.01. For
equations with two or moreindependentvariablesthe coefficients
of determina-
tion(R2)exceeded
0.80,0.90,and0.95forheight,diameter,
andvolume
incre-
ments, respectively.

608/FORESTSCmNCE
 TABLE 7.

Standarderrorsof estimatefor equations predictingincrementfrom all

combinationsof tree size (x), age (t), andtheir logarithms(In(x), In(t)). Size (x)
designates height,diameter,andvolume.The errorswere calculated using
pooleddatafor all 98 trees. Comparative accuracyof equations fitted to all
trees coincidedwith that fitted to the datapooledby site class.
Independent
variables Height Diameter Volume

In(x) 0.52 0.48 0.46

x 0.50 0.50 1.00
In(t) 0.41 0.34 0.94
t 0.37 0.36 1.06
In(x), x 0.48 0.48 0.44
In(x), In(t) 0.26 0.26 0.27
In(x), t 0.27 0.35 0.26
x, In(t) 0.38 0.29 0.89
x, t 0.32 0.33 0.97
In(t), t 0.35 0.34 0.88
In(x), x, In(t) 0.23 0.26 0.25
In(x), x, t 0.26 0.32 0.26
In(x), In(t), t 0.24 0.26 0.25
x, In(t), t 0.31 0.28 0.73
In(x), x, In(t), t 0.22 0.26 0.24

These calculations showedthat the basicequationforms(LTD andTD) were

amongthe mostaccurate.In addition,for individual trees the equationcontaining
tree size and its logarithmas independentvariables
ln(y')= k + pin(y)+ qyory' = k0PeqY (7)
wasequallysuccessful (Table8). Becausethe declinecomponent is proportional
to the sizey, this form will be referred to as the Y-decline,or YD form.
Other results of these calculations indicate that:

1. The accuracyof equationsincreasesdrasticallywhen the numberof independent

variableschangesfromoneto two. Subsequent additionof variableshadlittle effect.
The besttwo-variable
equationswere almostas accurateasequations withthree and
four variables.
2. In mostcasescomparative
accuracy
of equations
fittedto alltreescoincided
withthat
fitted to the datapooledby site class.
3. The effectof datatypeonthe accuracy
of a givenequation,
noticed
for thetwobasic
equationforms,wasmorepronounced whenthe set of 15 equationswasconsidered
(Table 7 and8). This effectis espedallyclearfor the YD form: it was the best form
for heightanddiameter(Table8) growthof individual trees andthe worstformfor
pooleddatafor the samevariables(Table7).
4. The distinguishing
featureof the threebesttwo-variable equationforms(LTD, TD,
andYD) is thatgrowthexpansion is proportional
to the logarithm
of size.

AUTOCORRELATION
OF RESIDUALS

Whencomparing accuracyof equations,

it isnecessary
to consider
autocorrelation
amongresidualsthat oftenappearsin time seriessuchas tree growthdata.
Autocorrelation
doesnot changeparameterestimatorsin least squareregres-

AUGUST 1993/ 609

 TABLE 8.

Standarderrorsof estimatefor equations predictingincrementfrom all

combinations
of tree size (x), age (t), andtheir logarithms(ln(x), ln(t)). Size (x)
designates
height,diameter,andvolume.The errorsare averagesof individual
regressionsof 98 trees.
Independent
variables Height Diameter Volume

In(x) 0.32 0.22 0.30

x 0.25 0.17 0.82
In(t) 0.28 0.18 0.41
t 0.23 0.18 0.65
In(x), x 0.15 0.14 0.15
In(x), In(t) 0.15 0.15 0.15
In(x), t 0.16 0.15 0.15
x, In(t) 0.18 0.15 0.18
x, t 0.21 0.14 0.26
In(t), t 0.16 0.14 0.17
In(x), x, In(t) 0.14 0.14 0.13
In(x), x, t 0.13 0.13 0.13
In(x), In(t), t 0.14 0.13 0.13
x, In(t), t 0.14 0.14 0.15
In(x), x, In(t), t 0.12 0.12 0.12

sions,but inflatestheir variance(as comparedwith completelyuncorrelateddata)

and introducesbiasinto the standarderror of estimate.This happensbecause
autocorrelation reducesthe effectivenumberof degreesof freedomfor estimating
the parameters,whichis equivalentto reducingthe samplesize. Therefore, the
estimatesof standard errorsandthe coefficientof determination
(R2) shouldbe
supplemented with an analysisof autocorrelation.
The investigationof autocorrelation
amongresidualsof the analyzedequations
showedthat autocorrelationis by far the largest at the first step. This fact
permittedthe restrictionof thisinvestigation
to first-orderautocorrelation.
It was
calculated (usingthe Durbin-Watson, DW, optionin the regressionprocedureof
SAS)forpooleddata,whichcontained 100trees,andfor eachtree separately.For
individualtrees the results(Table 9) representedmeanabsolutevalues.

TABLE 9.

First-orderautocorrelation
of equations
predictingtree increment.
Autocorrelations of individual trees are mean absolute values.

Pooled data Individual trees

Equation Equation
Tree Number of
variable observations LTD TD YD LTD TD YD

Height 1099 0.57 0.58 0.86 0.29 0.31 0.28

Diameter 1024 0.65 0.68 0.80 0.28 0.26 0.26
Volume 1088 0.56 0.56 0.80 0.24 0.26 0.25

610/FORESTSCIENCE
 These calculations demonstrated that:

1. Autocorrelation
waspractically
identical
for all equations
whentheywereappliedto
individual trees.
2. Whenappliedto pooleddata,autocorrelationofthe YD formwassubstantially greater
thanthat of the LTD andTD forms.Autocorrelation of the lattertwo wasstatistically
indistinguishable.
3. Autocorrelation for individual
trees wasequallylikelyto be positiveor negative,and
the mean of actual (not absolute)valueswas never differentfrom zero.
4. Autocorrelationvariedby datatype andequationin a way that was s'nuilarto that of
their standarderrors(Tables7 and8). Thisfactmakesthe analysisof autocorrelation
redundantfor Guttenberg'sdata.
5. This analysisof autocorrelationsdid not changethe previousconclusion basedon
standarderrors;for individual
trees allthreetwo-variable
equations,
in whichgrowth
expansion wasrepresentedby the logarithmof size, were equallyaccurate.

SINGLE EQUATION FORMS

The precedingresultbringsforththe question of whetherit is possible
to achieve
the ultimate reductionin the number of equationsand obtaina singleform. An
easyway to do this is to combineequations(4), (5), and(7) at the expenseof
introducingadditionalparameters:
y' = klyP•tq•
+ k2yP2e
qzt+ k3yP3e
qay (8)
The LTD formcorresponds to k2 = k3 = 0, andthe TD formariseswhenkx =
k3 = 0. Whenallthreeparameters are differentfromzero,Equation(8) becomes
a singlegeneralform that includesthe discussed forms(LTD, TD, andYD) as
specialcases.
A similargeneralequationwith five parameterswas suggestedby Dr. Insarov
(personalcommunication,
May 1, 1992):
y' = k•yPtqe
k2•3 (9)
It summarizes all the equationsgivenin Table 1.
An undeservedly ignoredpaperby Grosenbaugh (1965)described an equation
thatgeneralizesmanynonlinear functionsincludingseveralgrowthequations such
as Bertalanffy,Gompertz,andJohnson-Schumacher equations.In this equation,
Y = H + A(eav•-vu- NUfviq-x (10)
A, H, M, andN are parameters.U is an elementaryfunctionof an independent
variableandcontainstwo additional parameters.According to the author(personal
communication, March 23, 1992), no one ever madeuse of his equation.
An interestingattemptto arriveat a singleequationformwasmadeby Schnute
(1981). BredenkampandGregoire(1988)were the first to introducethisequation
to forestry.Whilemanyscientistsstudiedgrowthrate, y', andrelativegrowth
rate, z = y'/y, Schnutewent furtherandinvestigatedthe rate of a rate, that is,
acceleration of growth.Schnute(1981, p. 1129)believedthat the relativegrowth
rate of a relative growth rate is a linear functionof the relative growth rate
becauseit is "the simplestpossibleassumption":
ldZ
W-z dt- (a+ bZ) (11)

AUGUST 1993/611
where a and b are parameters(unrelatedto thoseof Table 1). Schnute(1981)
showedthat manyexistingequationsare specialcasesof his Equation(9). For
example,the GompertzequationfollowsfromEquation(9) whena > 0 andb =
0. When parametersa > 0 andb = - 1, the resultis the logisticequation.
The first thingthat comesto the mindof an empiricalresearcheris to test this
linearassumption (Schnutedidnotprovideone).Perhapsthisassumption is good
for describingthe growthof fish (the area of Schnute'sresearch)but not that of
trees.WhenI plottedrelativeacceleration, w, overrelativegrowthrate calculated
usingGuttenberg's(1915) averageheightgrowthfor five site classes,it became
evidentthat the relationshipis not linear(Figure1). A powerfunction
w = azb (12)
appearedto be more appropriate becausewhenplottedon the log-logscale,the
relationship
becamestraight,especially if three outliersin the lower left comer
are disregarded(Figure2). This relationship is characterized by a coefficient
of
determinationof about0.95. Both interceptsand slopeschangedlitfie with site
class.
Althougha singlegeneralsolutionis attractive,it is not clearwhetherit is worth
the costof additional
variables[Equations(8) and(9)]. It is probablybetter to rely
on two or three equationforms.

DISCUSSION

This investigationrevealedthat manyof the existinggrowthequationsbelongto

oneof two basicforms.The type of datauseddrasticallyaffectedthe comparative
accuracyandshapeof equations.Growthcurvesfor groupsof trees belongingto
the samesite classandaveragecurvesdifferedsubstantially from the growthof
individualtrees from the samegroups.
This investigation of knownequationsnot only providedthe basisfor their
classification
but alsomadepossiblethe discoveryof a new promising equation

0.06 +
I 3
12
4
2 4
•0.04 + 2 13 3
I 1
I I I 3 5 5
I 123 35
I 1245
0.02 + 122 35
41444
I 11
I
I
•u o.oo +
I
-+ ......... + ......... + ......... + ......... + ......... + ......... + .....
0.00 0.02 0.04 0.06 0.08 0.10 o.x2

Rela•.ive •ro•ch rate

FIGURE1. The relationshipbetweenrelativegrowthacceleration
andrelativegrowthrate for Gut-
tenberg's(1915)averageheightgrowthdata.The plottednumbersindicatesiteclassof investigated
trees.

612/FOP,m'rscm•cE
 o I
ß•1 I 1
o -3 + i 5 43 5
1 13 3
1 3. 3 3 5 5
3 3. 2 3 5 5
3. 23.2 52 535 35 5
ßel 0 I 3434 4 4 4 4

m I
•
0
•-•
-5 +I 5
• I 3
m I 2
• -6 +
k ---4 •. • ............. + ............. +--
-6 -5 -4 -3 -2

Logarithm of relative growth rate

FIGURE2. The relationship
betweenrelativegrowthacceleration
andrelativegrowthratefor Gut-
tenberg's
(1915)averageheightgrowthdataonthelog-log
scale.Theplotted
numbers hndicate
site
classof hnvestigated
trees.

form(the Y-decline
equation
form).Bothcomponents
of thisequation
are func-
tionsof tree size. It doesnot containage, the mostdifficultvariableto measure
(especiallyin olderhollowhardwoods).
Stronglinearrelationships
existbetweenthe threeparameters
of the consid-
ered two-variableequations.Thus, for LTD
q = -0.29 - 1.30p(R2 = 0.95,SEE = 0.22) (13)
and

k = -1.74 - 2.35p - 3.95q = -0.60 + 2.77p

(R2 = 0.92, SEE = 0.67) (14)
Theserelationships, calculated
fromparameters describing
heightgrowthof in-
dividualtrees, are independentfromsite class(andage)andcansimplifyincre-
ment estimations.
All three best two-variableequations (LTD, TD, andYD) haveonecommon
feature:growthexpansion is proportional to the currentsizeof a tree. This
indicatesthatgeometrical progression is a fundamentalcharacteristics
of growth,
aswaspostulated twocenturies agobyMathus(1798).Fortrees,unlikebacteria,
thisfeatureis notself-evident.In dicotyledon tree species
the numberof dividing
cells(cambium) per unitof stemsurfaceremainsconstant. Therefore,the expo-
nentialtendencyin diametergrowthprobablyresultsfromdecreased intervals
betweensuccessive divisionsandnotfromproliferation of dividingcellsasis the
casein a population
of bacteria.
Unlikethe expansion component, growthdeclinein individual
trees canbe
renderedwith equalaccuracy by a varietyof expressions.
This resultmaybe
explainedby the greatnumberof factorsthat hindergrowth:scarcityof re-
sources,competition,reproduction,
aging,diseases,herbivory,disturbances,
etc. Thismakesthe growthpathinherently
imprecise.It should
be regarded as
a broadvalleyratherthana singleline.
Thisresultdoesnotmeanthatwe loosetheclarityof ourpictureof growth.We
dispense
withamisleading
precision
readintogrowthequations.
Instead,
wegain
knowledgeabout
theactual
variability
ofthegrowth
process,
basiccomponents of
thisprocess,
andappropriate
analytic
expressions.
Thisconceptual
shiftfroma

AUGUST1993/613
misleadinglypreciseline to a fuzzybut realisticstripbringsclarityto our under-
standingof tree growth.

LITERATURE CITED
ALEMDAG, I.S. 1978. Evaluationof somecompetitionindexesfor the predictionof diameterincrement
in plantedwhite spruce.Can. For. Serv. Inf. Rep. FMR-X-108.39 p.
ASSM•, E. 1970. The prindplesof forestyieldstudy.PergamonPress,New York. 506 p.
BE•T•U•U•FF¾,
L. VON.1957.Quantitative
lawsin metabolism
andgrowth.Quart.Rev. Biol. 32:217-
231.

B•DENKa•MP,
B.V., andT.G. G•c, oI•. 1988.A forestryapplication
of Schnute's
generalized
growth
function. For. Sci. 34:790--797.

B•EwE•,J.A., P.Y. Btn•s, andQ.V. C^o. 1985.Short-termprojection

accuracyof five asymptotic
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AUTHOR AND ACKNOWLEDGMENTS

Boris Zeide is Professorof Forestry, Departmentof Forest Resources,Universityof Arkansasat
Monticello,Arkansas AgriculturalExperimentStation,Monticello,AR 71656.Thisreportis approved
for publicationby the Director, Universityof ArkansasAgriculturalExperimentStation.Valuable
commentsby JohnL. Greene,TimothyG. Gregoire,Lew R. Grosenbaugh, GregoryInsarov,Rolfe
A. Leafy, DanielJ. Leduc,RobertA. Monsemd,LynneC. Thompson,SuzanneWiley, RichardC.
WoolIons,andVoyteckT. Zakrzewskiare gratefullyacknowledged.

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