The Growth of Populations Author(s): Raymond Pearl Reviewed work(s): Source: The Quarterly Review of Biology, Vol. 2, No.

4 (Dec., 1927), pp. 532-548 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/2808218 . Accessed: 26/12/2012 20:54
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THE GROWTH OF POPULATIONS

BY RAYMOND PEARL

ofpopu- and in particular, in thegrowth involved social forcesof various lation are two in number:the sorts,and perhapsothers, But it should measured ofnatality, bythe always be kept in mind,and this I parforce wish to emphasize, thattheseare the force ticularly birthrate,on the one hand; ancd a biologicalpoint measuredby the death rate, all secondary of mortality, factors from on the other hand. These primaryele- of view. They producewhatevereffect to the discussion they may have upon the final result, mentsare fundamental of the growthof populationsof any and namelythe size of the populationat any to given moment, whatever,fromAmoeba all organisms by acting, more or less man. In most of the lower organisms powerfully as the case may be, upon one plant or more of the threeprimary whether living in a state of nature, biological and migraor animal, these are the only first-ordervariables,natality,mortality, in a variables which have to be taken into tion. Thus an economicdepression account in discussingthe problem,and particularcountrymay affectadversely in experimentalstudies on population the birthrateof thatcountry, or eventhe of the experi- death rate if the degreeof the depression growth the arrangement great or its durationsuffimentsis usually such that only natality is sufficiently and mortalitycan possibly directlydc- cientlyprolonged. These effects will, in of individualsin the greateror smaller degree, reflectthemtermine the number in the size ofthepopulation. selvesfinally population at any given time. In most humanpopulations,especially This finaleffect upon the growthof the areas, population may, however,be extremely largegeographical thoseinhabiting directlythe slight,and difficult a thirdfactormay influence or even impossibleof statistical or measurerecognition size of the populationat any given mo- separate in less This of influences or ment, because compensating degree. greater ment, third factor is migrtion,and it is theoreti- at work at the same time. Logically, the operation as a primary of thesesecondary variable however, cally to be regarded in determining the growthof such human factorsmust always be recognized. But fromthe point of view of the theoryof populations. biological the primary three Besides population growth their influenceis and migra- always a second order one. They can of natality,mortality, factors the observed upon populationonly growth produceany effects tionwhich influence the of human populations there are various by operating upon primary biological factors which forces of natality, mortality,and misecondaryenvironmental the final gration. may play a part in determining
532-

Institute forBiological Research, TheJohns HopkinsUniversity

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variablessupply,the economicsituationin general biological primary

VARIABLES

result. These are such things as food

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GROWTH OF POPULATIONS
HOW POPULATIONS GROW

533

It is an observed which at thisstage f&ct, of the discussioninvolvesno theoretical implications whatever, or postulates special to it, that the growthof populations of the most diverse organisms andcharacteristic course. followsa regular In generaland everyday terms of common TABLE i manner sensethischaracteristic ofpopula- Growth ofa population of yeastcells(data from Carlson) tion growth may be described in the QUANTITY OF QUANTIYOF ASOLUTE ASLT YEAST PROYATQUANTITYO followingway. The population at first YEAST DUCED PER QUANTITYS IN PERHOUR HOU'RS grows slowly, but gains impetusas it POFUYEAS HOUR UP TO THAT grows, passinggraduallyinto a stage of GROWTH (==TIPER RATE HOUR (= PEROBCALCUTF CENTAGE RATE Of SERVED LATED rapid growth, which finallyreaches a GROWTH) OF GROWTH) maximumof rapidity. Afterthis stage (b) (c) (a) (e) (d) of most rapid growth the population cent per increases ever more and more slowly, 9.6 o 9. 8.7 until finally thereis no moreperceptible 9o.6 i6.8 I I8.3 I 87 58 growthat all. In short,the populations 2.8.2. 107 5.5 2.9.0o i8.2. 62..8 wax in their of variousforms of life first 46.7 47.2. 3 50.6 2.3.9 and thenwane. 76.0 speedof growing 7I.I 4 67.5 48.0 12.0.1 5 This characteristic phenomenon maybe 46.6 55.5 6 I74.6 i8i.9 more preciselydescribedin a varietyof 82..7 47.4 7 2-57.3 2.60.3 ways, depending upon which of several 36.3 93*4 8 350.7 348.2. mathematicalaspects of the matterwe 90.3 2.5.7 44I.0 433.9 9 10c i6.4 513.3l5l6.9y72-3 choose to stress. This may best be ilIo 559.7 5o62.9 9.0 lustrated by a concrete example. For this 46.4 I12. 594.8 6oo.8 6.3 we maywell take a relatively 34.6 simplecase 3. 62.9.4 62.5.8 13 biologically, such as is afforded by the i.8 14 I1.4 64o.8 64I.5 growth of a population of yeast cells. hoursdui which 4.8 0.7 The yeastplantis an organism low in the i6 655.9 656.7 o.6 37 17 659.6 66o.i scale of life. Its reproduction, underthe 03 2. . i8 66i.8 662..i is by the conditionsof the experiments, processof cell division,the cells present at any given momentforming new cells investigation of the subject made by by budding. An experimenton the Carison (3), is shown in table i. A discussionof these data will be growthof a population of yeast cells is further made as follows: A measured work (8). The figures amountof foundin an earlier and nutriment to the of columns wort, which furnishes (b) (c) are shown graphii. yeast plants, is seeded with a few cells. cally in figure column(a) gives the Then at equal intervals of timethereafter In table i the first the equivalentof a censuscount is made hours during which growth occurred; of the cells thenpresent. Because of the thesecondcolumn theobserved (b) records minute size of the individualcells, and of quantityof yeast, or in otherwords the other technicaldifficulties, somc sort of size of the population,at each successive
OF PRODUCE CENT OF TOTAL 2. I89.I 2.2

indirect method is commonly used to measurethe size of the population, in place of direct counts. But this is a mattermerelyof detail, which need not discussion. detainus in the present The outcomeof a typicalexperiment of this sort, the data being taken froman

QUAR. REV.

BIOL.,

VOL. II, NO.

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534

THE QUARTERLY REVIEW OF BIOLOGY Column(d) tells us that in the growth of this yeast population the absolute increment of new yeast cells per unit of timeincreases each hour as the population grows, up to a maximumincrement per hour,which is attainedin this case somewhere about the 7thor 8th hour. Thereafter as growth continuesthe absolute increment of new cells per unit of time decreases hour by hour until it almost

hour;the thirdcolumn(c), which maybe neglectedfor the present,gives the size of the populationat each successivehour equaas calculated froma mathematical (d) columa discussed tion to be later; givesthe simpletimerateof growth,that is the absolute amount of new yeast by the growthof the population produced in in each successivehour. The figures values of the thiscolumnare the observed

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3I. THE GRowTH3OF A POPULATIONq CE-LL OF Y-BAST

In the publication of this diagramin (8), through an error in proof-reading the abscissalunitswerestated to be days,whenthey shouldhave beenhours,as is correctly shownhere. (Data from Carlson.)

ratio dy/dt, y denoting the size of population and t denotingtime in hours. The last column(e) gives the relativerate of growth, in proportion to the already attained size of the population, in each timeinterval. In mathematical terminolin this column are the ogy the figures observed values of the double ratio _

dyforeach hour.

vanishes,and, for all practicalpurposes, the population has ceased to grow. If the figures of column(d) are plotted, as in figure2., they obviously form a curveshaped like a cocked hat or sugarloaf, low at the two ends and risingto a peak near the middle. As a naturalconsequenceof this trendof the first differences the relative growthrates given i column(e) are foundto be largestat the

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GROWTH OF POPULATIONS startof growthand to becomesmalleras the absolute size of the population increases. Although this relative rate of growthpresented in column(e) is the one most oftendiscussedby demographers in talkingabout populationgrowth,it tells us much less clearly what is the real natureof the characteristic curveof such growth than does the simple time rate, or first derivative of the population growthcurvegiven in column(d). recentbooks (6, 8). It has been shown populations of yeast, that experimental of bacteria, and of the fly Drosophila in their growthfollow this melanogaster, characteristic curvewith greatprecision. it has been demonstrated Furthermore statisticallythat populations of human beingshave grownaccordingto the same type of curve, so far as may be judged fromthe available census records,in at least the following countries:Sweden,

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2.. GROWTH INCRBMENTS OP YBAST POPULATION IN SUCCBssivz TImBINTERVALS

Data from column(d) of table i

France,Austria, The phenomenon shown in column UnitedStatesofAmerica, (d), namelythat the absolute increment Belgium, Denmark,England and Wales, with Hungary, Italy, Norway, Scotland, of growthper unit of timeincreases time to a maximumvalue, and then de- Servia, Japan, Java, Philippine Islands, growth BaltimoreCity, New York City, and the creasestill the end of measurable years world as a whole. is reached,has beenfoundin recent In illustrationof this statementfour to be the way in which populations of here. These actually cases arepresented graphically such a wide varietyof organisms regarded are the United States (fig. 3), Sweden grow, that it may now be fairly normal mode of (fig. 4), France (fig. 5), and the world as the characteristic, population growth, to at least a first as a whole (fig. 6). In these diagrams approximation. I shall not take the time the circles give the census counts (or, here to review all the evidencethat this in the case of the world as a whole, is so. This has alreadybeen done in two estimates)of the populationsexistingat

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536

THE QUARTERLY REVIEW OF BIOLOGY sucha conclusion in somedegree probable. And one cannotconduct with experiments human beings on this point, as can be done with lower organisms. But fortunatelyit has been possible to findone group of human beings, the indigenous native populationof Algeria,in which a

the given dates,while the smoothcurves are the fitted theoretical curvesof population growth. In the case of the demographicunits listed above the census records do not

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3. THE OBSERVED AND CALCULATED GROWTH OF THE POPULATION OF THE UNITED STATES

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OBSERVED AND CALCULATED GROWTH OF THE POPULATION OF SWEDEN

extend over a sufficiently long time to make the case conclusivethat population would follow in growth,if undisturbed, human groups the complete course exhibited by the yeast population just discussed. The available data only make

cycle of population growth has been practicallycompletedduring the period for which census records are available, these having been carefully made by the French. In this case the humanpopulationfollowedin its whole cycleofgrowth a curve of the same characteristic form that has been discussed for the yeast. This case has been fully describedand

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OF POPULATIONS GROWTH

537

ofPopu- mortality analyzedin my book TheBiology are so integrated and correlated here in theiraction as to lead to a finalresult lationGrowth (8), and is illustrated in figure 7. in size of populationwhich follows this particularcurve ratherthan some other THE LOGISTIC CURVE one. This demandsthat we shall push The equation to the curve which has the enquiry further. beenfound and observation by experiment MATHEMATICAL CONSIDERATIONS to be descriptive ofpopulationgrowth in a There are two lines along which this wide varietyof organisms was firstdisinvestigation mustproceed. The covered by the Belgian mathematician, further is a theoretical mathematical analysis Verhulst(i8, see also I9, SO) in I838. first His pioneer work was forgotten,and of the relation of birth rates and death consequentlyoverlooked by most sub- ratesto the logistic curve,undervarious sequentstudents of the populationproblem. In i9Z0 the present writerand his 1600 colleague, Lowell J. Reed, without any knowledgeof Verhulst'spriorwork, in"0WORLD dependently hit upon the same equation Verhulst called his curvethe "logistic." This usage we shall follow. The characteristic appearance, and some of the mathematicalpropertiesof the logistic 8. curve,are shown in figure The thirdcolumnof table i (column c) shows the degreeof accuracy with which this logistic curveis able to describethe observed growthof a populationof yeast cells. For thewhole seriesof I9 observations the root mean square deviation resultsand the betweenthe experimental curveis only 3.59. The equatheoretical curveis: tion of the fitted
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wherey = quantityof yeast, and x = time of growth in hours. The possessionof such a curveas this, which is found by actual experienceto the course of populadescribeaccurately tion growth in a wide variety of organisms,is a valuable firststep, but only a firststep, towards reaching an of the biology of the understanding process. What we want to know is how the biological forces of natality and

postulates. We have done a considerable amount of work, as yet unpublished, upon this subject. Certain aspects of it have also been discussed by Yule (z5) and Lotka (5). I shall not attempt to present the details of this analysis on this occasion, because of their technical character. Some of the results, however, may be stated in simple form and are pertinent to the present discussion. It is easilyshownthat ifbirthratesand death rates are assumedto remainconstant, at any values consistent with growthat all (i.e., birthrateslargerthan

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538

THE QUARTERLY REVIEW OF BIOLOGY

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GROWTH OF POPULATIONS death rates) the population will grow accordingto an exponentialcurve,on to infinity.The rate of this progression may be slow or rapid according to the assumptionsmade, but the form is exponential. Nothing like the slowing down of growthaftera period,which is

539

with this growth, pands proportionately so that each single elementalways has how large the plentyof room,no matter whole group becomes. But such an asfor any actual sumptionis unwarranted populationwhatever. All populationsof real organisms live in universes with

LOGISTIC CURVEAND ITS FIRRST DERIVATIVE CURVE ASYMPTOTE

O,

a b

TIME

FIG. 8. THE LOGISTIC CURVE AND ITS FIRST DERIVATIVE

seen in the logistic curve,appears under this postulate. But plainly,with actual populationsof living organisms, birth rates cannot In saying indefinitely. continueconstant that birthrates shall be constantregardless of the size of the populationalready attained, it is tacitly, but implicitly, assumed that the universein which the hypotheticalpopulation is growing ex-

limits. The absolute size of the definite universemay be small, as in the case of which holds the yeast cells, the test-tube or it maybe as largeas the earth,mostof which could conceivably be inhabited, on a pinch,byman. But in anyand every limit case there is ultimatelya definite to the size of the universein which any real populationlives. This consideration obviouslyalters the

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540

THE QUARTERLY REVIEW OF BIOLOGY

complexionof the case. For it at once and knownconditions. Not onlycan the follows that if the universesin which growthof such an experimental populalive are finite and inextensible, tion as a whole be observed, populations but also it is theremustcome a timein the growthof possible to set up experiments in which any population when the individuals each separate variable such as natality, composingit begin to get crowded, and mortality,density of population, etc., as the growth continues beyond this can be particularly studied, and its bepoint they will get more and more havior under controlled conditions decrowded. scribedand measured. It is to this mode hypothesis It is not an unreasonable a of attackupon the problem that muchof see there is the research priori,and as we shall presently energy of my laboratory has and observational been devoted duringthe past five abundantexperimental years. supportfor the view, that crowding,or The organismchiefly used has been the as it is technicallytermed,density of fruit-fly Drosophila the form melanogaster, as material population, when it reaches a certain so widelyemployed forgenetic upon both of studies. has an adverse effect degree, the two primary biological forcesunder- Before undertakingto present some lying population growth, natality and accountof theresults of theseexperiments mortality. This assumption has been I wish to point out that the significance made and the theoreticalconsequences of these, or any similar experiments on in the discussionof the worked out, underseveral simplepostu- lower organisms, lates as to the quantitative relationships problemof humanpopulationsshould be neitherover-rated nor under-rated.Obthevariables. between The net resultof this first, or mathe- viously in all such experiments the conmatical, attack upon the problem of ditions of life are not only simplerbut than any underwhich populationgrowthis to show that,with also moreuniform simple postulates as to the relations human populations live. Further the variables, organisms concerned are less complex between the two first-order and psychologically than birthrate and deathrate,and the second- physiologically reached beings. No conclusion ordervariabledensityof population,and arehuman without consideration of any other by the studyof experimental populations we are led rigorously of lower organismscan safelybe transvariableswhatever, to the conclusionthat underthesepostu- ferred by simple inferenceto human lates the growth of population must populations. It must in every case be follow that typeof curve(the conclusivelyshown by independent innecessarily found to vestiga-tion ofhumanpopulations logistic) which is empirically thatthe thegrowthof actual populations same conclusion holds for them, if it describe if the same resultis, does. Furthermore ofwidelydiverse organisms. in fact,foundto appear in both experiEXPERIMENTAL POPULATIONS mental and human populations, by no that the The second mode of attack upon the meansmay it safelybe inferred causes the of to this result are leading of population necessarily biology problem growthis the experimental. Its purpose the same in the two cases. They maybe is to observeat each stage the growthof or they may not. To find out which is truedemandsspecial ad hoc the populationof some actual organisms, alternative small enoughto be capable of laboratory investigation. On the otherhand it mustalso be kept under carefullycontrolled management,

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GROWTH OF POPULATIONS

54I

in mind that the forcesof natality and the effectof these factors on human arebasic biological attributes of populationgrowthhas in thepast, at any mortality suchas to makethegrowth of the oflivingthings, menas well as rate,been aggregates flies. The ultimate biological forces lead- population rather follow the logisticcurve, from it. During the past few ing to the observedgrowthof a human thandeviate has been made that population are these two, natality and years the argument the very populations which have in fact are in a as as mortality, just truly they to a according or yeastcells. Because grownwith greatprecision populationofflies of the much greaterease and precision logistic curve,cannotpossiblyhave done with which the behavior of these so, because this curve does not "take variables, under diverse controlledcon- account of" a lot of these second-order ditions,may be analyzedin experimental variables. This argument is rubbish, populations of lower organisms, the bornout of the conservative resistance to such studiesmaybe any new idea which the established order results obtainedfrom of great value in the directionof sug- of learninghas always shown, by that gesting pointsforstatisticalinvestigation wind-broken and spavined old stallion, in humanpopulations. Over and beyond faithin a priori logic as againstplain facts is the further one that this consideration of experience. As a matterof fact the populations of whatever organismsare, logistic curvedoes "take accountof" all in their very nature,aggregate wholes, these second- and third-order variables, and behave in growthand other ways as in thesensethatit describes theintegrated such (cf. Wheeler, The elements end effect 2I-2.4). upon population size of the their out behaviorarising of this con- aggregated of forces towardsincrease tending dition of aggregate wholeness are just as in numbers on the one hand, and decrease true in populationscomposedof individ- in numbers on theotherhand. The popuuals of one species as in those composed lation growthof the United States is an of individuals of another. excellentcase in point. If one plots the Perhaps the most impressive thing census counts of this population from which has come out of the statistical I790 to i92.0 it is impossible to detect in studyof humanpopulation growthis the the curveof growthany separateor disevidence that the steady onwardmarchof turbingeffect of immigration, although, this growthis not sensiblyalteredfrom as we have seen,migration is theoretically the logisticcourse,in the greatmajority a first-order variable in population of actual cases, by the host of economic growth. The actual observed growthof and social eventswhich are supposedof thepopulation oftheUnitedStates follows logical necessity to affect it. This state- the logistic curvewith remarkable preciment should not be taken to mean that sion. To suit various theoristsit preeconomic and social factors are unim- sumablyoughtnot to, but in factit does. portant in their relation to population BenjaminFranklinlong ago pointedout growth. So far am I fromholding any that for the growth of population agsuch opinion that it is my belief that gregates ofanyconsiderable size migration economic forces are probably the most was an unimportant factor,as compared important single elementin the biologi- with natural increase by reproduction. cally effective environment of civilized He was right. Whateverthe future may man today. All that is meantis that, in develop, the past historyof the matter the majorityof observedcases at least, shows plainly that human populations

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542.

THE QUARTERLY REVIEW OF BIOLOGY

have behavedin theirgrowthin the same way that experimentalpopulations of lower organisms do, with truly remarkable faithfulness.

literallydo nothingfurther exceptto let nature take its course and to count the fliesat intervals. What then happensis that the populationgrowsalong alogistic curvefora littlemorethanhalfof a comDROSOPHILA POPULATIONS plete cycle of such a curve. Then quite The details of the techniqueof making suddenly thepopulationbeginsto decrease on population with the fly in numbers. This decreasegoes on at an experiments have alreadybeen accelerating melanogaster rate, until thereare no surDrosophila described(8), and need not be repeated vivorsleft. to say that in bottles The resultis precisely here. It will suffice that which is to size definite amountsof food be expectedmathematically of measured on the postumaterial are placed, and then an initial lates that birthrates decreaseand death groupofa fewflies-say one male and one rates increase uniformly at a constant linear rate. This is shown in table 2. TABLE 2. In the actual experiments the changes assumed in the birth and death rates which lead anddecline population Thegrowth ofa hypothetical and tobesubject therewith twoindividuals tostart to this type of curve,with first growth birthand deathrates after to the indicated and then decline of population, are POPULAPERCENTPERCENTdirectlyassociated with the diminishing DEATHS BIRTHS TION AT AGE DEATH BIRTH IN IN PEIDAGE IN RATE IN food supply under the conditions of a PEIDBEGINNING PEIDRATE OF PERIOD PERIOD PERIOD limitedand closeduniverse. The secondtypeof experiment is one in 0.2. 2_0 2_.0 IO IOO I 3.8 2D o.8 3.4 290 ismadetoadd foodas the whichan attempt I.9 8o 30 5.2. 3 6.5 supply is used up. The technical diffi6.8 9.7 3.9 40 70 4 cultiesof doing this satisfactorily with a i2L.6 6o 6.3 7.6 5 50 Drosophila population are considerable, 6o 8.3 6 6.9 I3.9 50 caretheycan be overcome butby sufficient 70 8.7 40 I2-.5 7 5.0 8o 2_.6 8.7 8 7.0 30 in large degree. The resulting growthof 2.0 4.4 3.9 9 90 0.9 the population then follows a simple 0.1 IOO 10 IO I .3 I .3 logistic curve, as I have described in detail elsewhere (8), and as is shown female-is added. At regular intervals graphicallyin figure9. There are two the thenexistingpopulationin pointsin connection with experiments thereafter of each bottle is carefully counted and this typewhich seemto need emphasisin work. The firstis recorded,and the individualscomposing the light of further to the bottleto carryon their that,owing to the technical difficulties of it returned normallives until the next censuscount. adding new food to a bottle forreasons There are several ways in which an that are fully discussedin the reference are always subon the growthof a Drosophila cited,the foodconditions experiment of this type. The population can be carriedout. Some of optimalin experiments in a manner secondpoint is that the smoothness in principle with theseways differ in determining the which the populations followthe logistic that is of importance is probkind of result obtained. The simplest curvein this typeof experiment in theway ably a directconsequence of the factjust case is to set up an experiment that the conditions as to food described brieflyabove, and thereafter mentioned

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OF POPULATIONS GROWTH
are definitely sub-optimal. The nutritional level of the bottle is not high. It only a little more than barely permits continued growth of the population. Under these circumstances violent oscillationsof either birthratesor death rates do not occur. The observations of population size lie smoothlyon the logistic curve until the asymptote is nearly reached.
400

543

population. At the end of threedays a censuscountis taken and the population to a new bottle, A2, conis transferred new foodto thesameamount taining fresh as was presentinitially in A1. At the end of anotherthreedays a censuscount is again taken and the population again to anotherbottle, A3, which transferred food, and so on again containsnew fresh Meantime the lag bottles, indefinitely.

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censuscounts,and thesmoothcurveis thefitted areobserved The circles logistic. (FromPearl(8))

which I The thirdtypeof experiment, way for here reporton in a preliminary time,is so plannedas to have the the first food conditionsalways optimal both as to quality and quantity. This is accomplishedin the followingway. Supwith our pose a bottleA1 to be prepared

A1, A2, etc., which the adult population are examined has left by transference, in any such bottle any daily. Whenever eggswhich werelaid flieshatchout from by the femalesof the adult population in that bottle, while they were resident these newly hatched flies are counted food(9), whichhas been (as births)and addedto the adultpopulanew synthetic to lead to lower tion in whateverbottle it happensthen shown experimentally ratesthanany to be. So then we get a recordof the birth deathratesandhigher other natural or laboratoryfood tried, daily birthrate of the population. Also in which and to be seeded with an initial small each day the bottleis examined

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544

THE QUARTERLY REVIEW OF BIOLOGY this asymptotic level as long as the experimenterdesires. A striking result, however,is that both duringthe growth period and thereafter there are violent oscillations of the populations in size, about its mean position as given by the fitted curve. In fact thesewaves in the size of the population,produced by oscillations in the birthand death rates, are perhapsthe mostcharacteristic feature of of this particular populationexperiments type. It has not so farbeen possible to deviseanymethod of holdingthesepopulationsin a steadystateat thelevel of the when thereis at all timesan asymptote, abundance offresh food. The population simply waves up and down about an averagesize. I believethatthiscondition ofunstable equilibrium is, in partat least, with the optimalfood causallyconnected conditions. A detailed account of these will be published, it is hoped, experiments some timein the courseof the nextyear.
THE EFFECTS OF DENSITY OF POPULATION

the adult population is actually then resident,and the deaths recorded. So thenthe situationin theseexperiments is that (a) thereis a new and abundant crop of fresh food everythreedays,(b) all the youngare bornin a lying-in hospitaland transferred to and counted intothegeneral populationonlyafter theyhavebeenborn and driedoffthere,and (c) thereis complete registrationof both births and deaths, as well as census counts of the
TABLE 3 data ontheinfluence Experimental ofdensity ofpopulation in Drosophila onrateofreproduction
PAIRS PER BOTTLE AT START MEAN POPULATION 16 DAYS AVERAGE AEAE IMAGOES PRODUCED PER FEMALE PER DAY. OBSERVIED CALCULATED NUMBER OF PROGENCY PER FEMALE PER DAY

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4.I 3.1
5.2

It has been stated above that the growth of a populationalong the logistic 10 3.6 curve can be completely accountedforon 2 I.70 I2. 3.5 3.I mathematical theoretical grounds by makI5 2.6.o2. 2. 5 2..3 I.8 2..2. 34.02. 2.0 ingcertain simplepostulates the regarding I.0 I.I 2.5 47-75 relationsbetween three variables, birth 30 47-73 rate,deathrate,anddensity ofpopulation. 90.66 0.34 0.33 50 The basic postulatewas that increasing density of populationhas associatedwith populationas a whole. The population it adverse changesin birthratesand death is dependent upon any given singlefood rates. Is thereexperimental evidencein supplyonly over a period of threedays. supportof such a postulate? For several years an intensiveexperiDuringso shorta periodeven the largest has beengoingon in populations obtained do not begin to mentalinvestigation exhaustthe food. my laboratory regardingthe effectof Manyexperiments ofthistypehavebeen densityof populationupon natality and resultis thatthe mortality in Drosophila. The chief performed.The general results reviewed. populationfirst grows up to a maximum of these studiesmay be briefly or asymptotic Some years ago Pearl and Parker(ii) level, just as in the second typeof experiment described above. But showed that if a countwere made of the in thiscase the populationcan be keptat progeny (adult fliesor imagoes) produced
4.0
I.2I.I

I4.48 i6.62. 18.74

4.6

5.6

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GROWTHOF POPULATIONS
amount in bottlesofuniform size, uniform air space, but with of food, and uniform theresults densities ofpopulation, varying werethoseshown in table 3. These resultsare shown graphicallyin io. figure
22

545

rapidlyand then more and more slowly at higherdensities. The total numberof flies(23,922) is large enough to progeny in the results. give confidence The last column of table 3 gives the calculated ordinates of the equation, by the method fitted to the observations of least squares,
logy = I.54 - o.oo8 x - o.658 log x
TABLE 4 of densities at different oflifeofDrosophila Meanduration population
WEIGHTED AVERAGE MEAN DENSITY TO WHICH FLIES HAD BEEN SUBJECTED UP TO TIME OF DEATH MEAN DURATION OP LIFE I DAYS LF NDY

20

18

14

INITIAL DENSITY

4 6
0
IO I2.

I.77
3.30

17.3I
29.32

4o.6o

0o.58
Qo.6i

6.68
8. I5
9.72.

5.00

34-45 4o.65
34.2.0
34.31
37.07

d0.72. 36.2-.2

4 o. 6i
h0.55

I5
2.0

I.. 42. I6.69

37.92. 4o.66 37-47 t0o.49 39.43 to. 67 37-46 t0.-5I

40.04 -t0.53 3525 d0-.45 32.34 A-o.46 .36 30.?IOo
2.4-2.0 4-0.32.

2.5

35 45
55 65

2.0.68 2.8.85 37-.23

10

40

eoW 90

95 105
125 I50 .00

75 85

44.65 53.I6 59.66

66.95

MEANF JES P&

FIG. 10. SHOWING
CHANGES

BOM
OF

74.50 80.36 94.38

iii.88

o 17I7 A.36

I9.60 =Io.2.8 z-0.2.4 16. I7

II.93

DENSITY OPl WITHINCREASING OrDROSOPEILA TH:EMATED POPULATION

Ix RATE

REPrOtTcrioN

144-47

0 .2.o

and the smooth The circlesgive the observations curve is the graph of the logarithmic equationdiscussedin the text.

wherey denotesnumberof progenyflies perday overa sixteenperfemale produced day period, and x denotesmean density and of the mated population (measured as a profound There is evidently fliesper bottle). in therateofreproduction change regular that this equation It is at once apparent of underthe conditions of Drosophilag, facts with extraobserved the describes denwith increasing theseexperiments, even in a Rarely, ordinary precision. of progof population.The number sity does chemical experiment, or physical per day declincs perfemale enyproduced obbetween closer one agreement get at first as density extremely increases,

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546

THE QUARTERLY REVIEW OF BIOLOGY under different densities of population. The resultswill shortlybe published in detail. Here it is only necessaryto say that they show conclusively that the effect in populationsof primary ofdensity, is upon fecundity (numberof Drosophila, eggs laid). The number of eggs produced perfemale perdaydecreases with advancing density of population according to the same type of curve as that shown for progenyproducedin the last column of table 3. So then it may be concludedthat in-

servationand theorythan is here shown. of the Plainly the curveis the expression law relatingthese two variables,rate of and density. It is the inreproduction verse of Farr's law relating death rates and density in human populations resultson the effect These experimental of density of population on the birth many times in rate have been confirmed my laboratory(cf. io). so farmenIn all of these experiments of densityupon birth tioned the effect
45 40

(4, I, z).

o20

5 /0 20
FIG. II.

ao 40

60

60 70

O 90

Initial Density

100 /

/50 120 180 140

160 /70 /80 i90 200 210

DENSITIES OF POPULATION

MEAN DURATION

oF LIFE

oF WILD

TYPE DROSOPHILA AT DIEPRENT

rates was measuredby counts of adult progenyflies (imagoes) produced. This procedureleft in doubt the point as to its effect through produces whether density offecundity (egg production) an alteration or at some later stage in the process of reproduction.This doubt has now been cleared up by an elaborate series of excarried out in my laboratory periments duringthe past year by Dr. Arata Terao. carefulcountswere In these experiments made of the number of eggs produced

creasing density of population does in fact have associated with it in experimentalpopulationsof Drosophila just the sort of adversechanges in the birthrate which were postulatedin the theoretical discussionin the preceding sectionof this paper. Elsewhere(8) evidencehas been presented showingthat the same kind of effect of densityis observablein populations of other organisms. The resultsof a large amountof experimentalwork regarding the effect of den-

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OF POPULATIONS GROWTH
sity of population upon mortality have recently been publishedin detail. Only the broad outcomecan be presented here. This is done in table 4, which gives the meanduration oflifein days of Drosophila subjected to the indicated densities of population. The experimentsinvolved
I3,000

547

in figure ii. graphically What the figures in table 4 show is that aftera densityof 55 fliesper one-ounce bottleis passed,the meandurationof life steadily decreases as density of population increases. Anotherway of putting the same thingis to say that the death rates in these experiments increasedwith increasingdensityof population afterdensity 55 has been passed. The figures for mean durationof life are necessarily determined by the values of the death rates at ages. Other experiments have demonstrated that with extremely high densitiesof population the mean durations of life (or equally the death rates) approach an asymptote. In densities below 55, aftera relativelysmall initial rise,there is littlechangein meanduration oflife(or deathrates)with smallincreases in density.
SUMMARY

flies. The results are shown

To summarize: A large amount of mathematical, statistical, and experimental study has brought to light the the genfollowingbroad factsregarding eral biologyof populationgrowth: i. Populations ofthemost oforganisms diversekinds,rangingfrombacteria and yeast to man, are found statisticallyto follow in theirgrowtha particulartype thelogistic. ofcurve, 2.. Mathematical investigation shows that a curve of this type is necessarily undercertainsimplepostulates generated

betweenthe two as to the inter-relations first-order variables,birthratesand death rates, and the second-ordervariable, density of population. One particular set of such postulatesis that it shall be that birthrates are markassumed,first, by small increases edly affected adversely in density at relatively low densities, while after a certain density is passed further increases produce only slight decreases in birth rates down to an asymptotic limit; and, second,that death affected rates are insignificantly by increasingdensityat relativelylow densia certain density is passed ties,while after death rates markedlyincreasewith increasing density up to an asymptotic limit. investigation of popu3. Experimental under lations of Drosophilamelanogaster controlledconditionsshows that in fact the relationsbetween densityofpopulation and birthrates,on the one hand,and density and death rates, on the other hand,which actuallyexistin suchpopulations, are in accord with those theoretiin thepreceding paragraph. cally assumed In shortit is possibleto accountforall the main featuresof the growth of experimental populations of Drosophilaby a simple hypothesisas to the correlated behavior of threevariables. A great deal more work needs to be done on the problem,and the investigations are being continuedin my laborapaperis to be regarded tory. The present only as a progressreport. In condensed it was read at the WorldPopulation form in Geneva, August 3I, I92-7, Conference underthe title "The Biology of Population Growth." It has been revisedfor publicationin the QUARTERLY REVIEW OF BIOLOGY, and this publicationis to be reform of thepaper. gardedas thedefinitive

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548

THE QUARTERLYREVIEW OF BIOLOGY

LIST OF LITERATURE
J. Notes on the biology of a life table. Jour. Roy. Stat. Soc., Vol. 82., pp. Discussion,pp. 66-77. 34-65, I9I9.
. Density and death-rate: Farr's law. Ibid., Vol. 83, pp. 2-8o-i83, I92.0.

(i)

BROWNLEE,

representasince I790 and its mathematical

(I3)

tion. Ibid., Vol. 6, PP. 2.75-2.88, i92o0. . A further note on the mathematical

(X)

of populationgrowth. Ibid., Vol. 8, theory

PP- 365-368, i92.2.. . Predicted Growth of New York and and its Environs), I92.3. PP. 42..

T. (3) CARLSON,

undGrosse (I4) UberGeschwindigkeit in Wiirze. Biochem. der Hefevermehrung Ztschr.,Bd. 57, pp. 3I3-334, I9I3(4) FARR, W. Vital Statistics: A Memorial (I5) Volume of Selectionsfromthe Reports and Writingsof William Farr, M.D., D.C.L., C.B., F.R.S. Editedby Noel A. Humphreys. (i6)
(5) LOTrCA, A.
I92.5.

its Environs. New York (Plan of New York

. On the mathematical theory of population growth. Metron, Vol. 3, PP. . The probable constants error ofcertain of the population growth curve. Amer.
Jour. Hyg., Vol. 4, PP. 2.37-140, I92.4. . Skew growth curves. Proc. Acad. Sci., Vol. ii, pp. i6-12., I92.5.
VERHULSt,

6-I9, I92.3-

London, i885.

J. Elementsof Physical Biology. Baltimore (Williams and Wilkins Co.),

R. Studiesin Human Biology. Baltimore (Williams and Wilkins Co.), I92.4.
Pp. 653. The curve of population growth.
I0-I7,

Pp. xxiv + 563.

(17)

Nat.

Pp. xxx + 460.

(6)

PEARL,

(i8)

P. F. Noticesurla loi que la population suit dans son accroissement.Corr. math. et phys. publ. par A. Quetelet.T. X, d'accroissementde la population. Nouv. mem.de l'Acad. Roy. des Sci. et Belles-Lett.
PP- II3-I2I, I838. . Recherches mathematiques sur la loi

(7)

Proc. Amer. Phil. Soc., Vol. 63, pp.

I92.4.
.

(19)

(8)

The Biology of Population Growth. New York (Alfred A. Knopf, Inc.), I92.5.
Pp. xiv +
2.60. (2.0)

(g) PEARL, R., ALLEN, A. L., and W. B. D. PENNIMAN.

Culturemedia for Drosophila. III. on and its influence medium A new synthetic densitiesof population. at different fertility
Amer. Nat., Vol. 6o, pp. 357-366,
i92.6.

de Bruxelles, T. i8, pp. I-38, I845. . Deuxieme memoire sur la loi d'acde la population. Ibid., T. 2.o, croissement PP. I-32., I847. WHEELER, W. M. The ant colony as an organism. Jour. Morph., Vol. 2.2., pp. 307-32.5, I9II.
.

(2.1)

(IO)

PEARL,

R., MINER, J. R., and S. L. PARKER. studieson the durationof life. Experimental XI. Densityof populationand life duration in Drosophila. Ibid., Vol. 6i, pp. 2-89-3i8,
I92.7.

(2.2.)

Social Life Amongthe Insects. New

Les Societes d'Insectes. Leur Ori-

York (Harcourt, Brace and Co.), I92.3.

(2.3)

(II)

PEARL,

R., and S. L.

PARKER.

On the influence

gine-Leur Evolution. Paris (Doin), i92.6. PP. xii + 468.

(2.4) (2.5)

(I1)

in Drosophila. Proc. Nat. Acad. production Sci., Vol. 8, pp. 2.I2--i9, I92S. On the rate of PEARL, R., and L. J. REED. ofthepopulationoftheUnitedStates growth

of density of population upon rate of re-

. Emergent evolution and the social. Science, Vol. 64, PP. 433-440, i92.6.
YULE,

whichcontrolit. Jour.Roy. and the factors

Stat. Soc., Vol. 88, pp. I-58, I92.5.

G. UDNY.

The growth of population

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