AsGilbertWhite,Darwin,andothersobservedlongago,allspeciesappeartohavetheinnatecapacitytoincreasetheirnumbersfromgenerationtogeneration。Thetaskforecologistsistountangletheenvironmentalandbiologicalfactorsthatholdthisintrinsiccapacityforpopulationgrowthincheckoverthelongrun。Thegreatvarietyofdynamicbehaviorsexhibitedbydifferentpopulationmakesthistaskmoredifficult:somepopulationsremainroughlyconstantfromyeartoyear;othersexhibitregularcyclesofabundanceandscarcity;stillothersvarywildly,withoutbreaksandcrashesthatareinsomecasesplainlycorrelatedwiththeweather,andinothercasesnot。
Toimposesomeorderonthiskaleidoscopeofpatterns,oneschoolofthoughtproposesdividingpopulationsintotwogroups。Theseecologistspositthattherelativelysteadypopulationshave“density-dependent“growthparameters;thatis,ratesofbirth,death,andmigrationwhichdependstronglyonpopulationdensity。Thehighlyvaryingpopulationshave“density-independent“growthparameters,withvitalratesbuffetedbyenvironmentalevents;theseratesfluctuateinawaythatiswhollyindependentofpopulationdensity。
Thisdichotomyhasitsuses,butitcancauseproblemsiftakentooliterally。Foronething,nopopulationcanbedrivenentirelybydensity-independentfactorsallthetime。Nomatterhowseverelyorunpredictablybirth,death,andmigrationratesmaybefluctuatingaroundtheirlong-termaverages,iftherewerenodensity-dependenteffects,thepopulationwould,inthelongrun,eitherincreaseordecreasewithoutbound。Putanotherway,itmaybethatonaverage99percentofalldeathsinapopulationarisefromdensity-independentcauses,andonlyonepercentfromfactorsvaryingwithdensity。Thefactorsmakinguptheonepercentmayseemunimportant,andtheircausemaybecorrespondinglyhardtodetermine。Yet,whetherrecognizedornot,theywillusuallydeterminethelong-termaveragepopulationdensity。
Inordertounderstandthenatureoftheecologist‘sinvestigation,wemaythinkofthedensity-dependenteffectsongrowthparametersasthe“signal“ecologistsaretryingtoisolateandinterpret,onethattendstomakethepopulationincreasefromrelativelylowvaluesordecreasefromrelativelyhighones,whilethedensity-independenteffectsacttoproduce“noise“inthepopulationdynamics。Forpopulationsthatremainrelativelyconstant,orthatoscillatearoundrepeatedcycles,thesignalcanbefairlyeasilycharacterizedanditseffectsdescribed,eventhoughthecausativebiologicalmechanismmayremainunknown。Forirregularlyfluctuatingpopulations,wearelikelytohavetoofewobservationstohaveanyhopeofextractingthesignalfromtheoverwhelmingnoise。Butitnowseemsclearthatallpopulationsareregulatedbyamixtureofdensity-dependentanddensity-independenteffectsinvaryingproportions。