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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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mortality.mortality. or eventhe of the experi. more or less man. Logically. in thegrowth involved social forcesof various lation are two in number:the sorts.the economicsituationin general biological primary VARIABLES result. But fromthe point of view of the theoryof populations. reflectthemtermine the number in the size ofthepopulation. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . biological the primary three Besides population growth their influenceis and migra. In most of the lower organisms powerfully as the case may be. may play a part in determining 532- T Institute forBiological Research. upon one plant or more of the threeprimary whether living in a state of nature. mentsare fundamental of the growthof populationsof any and namelythe size of the populationat any to given moment. thattheseare the force ticularly birthrate. They producewhatevereffect to the discussion they may have upon the final result.and perhapsothers.cientlyprolonged. population may.and particularcountrymay affectadversely in experimentalstudies on population the birthrateof thatcountry. In most humanpopulations. These are such things as food This content downloaded on Wed. factors from on the other hand. whatever. in a variables which have to be taken into tion. These effects will.of view.always a second order one.natality.and difficult a thirdfactormay influence or even impossibleof statistical or measurerecognition size of the populationat any given mo. They can of natality. ancd a biologicalpoint measuredby the death rate.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.separate in less This of influences or ment. these are the only first-ordervariables. because compensating degree.THE GROWTH OF POPULATIONS BY RAYMOND PEARL ofpopu.and in particular.fromAmoeba all organisms by acting. all secondary of mortality. bythe always be kept in mind.mortality. third factor is migrtion. Thus an economicdepression account in discussingthe problem. But it should measured ofnatality.on the one hand. selvesfinally population at any given time. 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.and misecondaryenvironmental the final gration. however. TheJohns HopkinsUniversity IHE variablessupply. These primaryele.and it is theoreti. biological and migraor animal. greater ment. the operation as a primary of thesesecondary variable however.at work at the same time. in of individualsin the greateror smaller degree.be extremely largegeographical thoseinhabiting directlythe slight. cally to be regarded in determining the growthof such human factorsmust always be recognized.and this I parforce wish to emphasize.especially This finaleffect upon the growthof the areas.

8 6. 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. i8 66i.8 6oo. BIOL.5 2.2 indirect method is commonly used to measurethe size of the population.7 o.1 5 This characteristic phenomenon maybe 46. mathematicalaspects of the matterwe 90.7 until finally thereis no moreperceptible 9o.9 more preciselydescribedin a varietyof 82. 3 50. depending upon which of several 36. 9.6 simplecase 3.8 wax in their of variousforms of life first 46. A discussionof these data will be growthof a population of yeast cells is further made as follows: A measured work (8). or postulates special to it.5 6 I74.6 i8i.7 47. yeast plants. is seeded with a few cells.8.4 I12. REV. underthe 03 2.6 66o. 76. hoursdui which 4. 62. II. which need not discussion.9.3 I 87 58 growthat all. but gains impetusas it POFUYEAS HOUR UP TO THAT grows. 107 5.5 48.6 37 17 659.5.0 433.I 2. of the cells thenpresent.60.. detainus in the present The outcomeof a typicalexperiment of this sort.0 lustrated by a concrete example.GROWTH OF POPULATIONS HOW POPULATIONS GROW 533 It is an observed which at thisstage f&ct.9 656. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions .8 I I8.8 64I.4 7 2-57. The figures amountof foundin an earlier and nutriment to the of columns wort. In short. which furnishes (b) (c) are shown graphii.3 93*4 8 350.8 0. Its reproduction.6 55.the populations 2. passinggraduallyinto a stage of GROWTH (==TIPER RATE HOUR (= PEROBCALCUTF CENTAGE RATE Of SERVED LATED rapid growth. For this 46. which finallyreaches a GROWTH) OF GROWTH) maximumof rapidity.the cells present at any given momentforming new cells investigation of the subject made by by budding.6 i6.0.8 662.2.4 62. or in otherwords the other technicaldifficulties.6 o 9. in place of direct counts.at each successive OF PRODUCE CENT OF TOTAL 2. 62. 594. is shown in table i.Growth ofa population of yeastcells(data from Carlson) tion growth may be described in the QUANTITY OF QUANTIYOF ASOLUTE ASLT YEAST PROYATQUANTITYO followingway.3 2.6 2..7 348. .0 speedof growing 7I.8 13 biologically. followsa regular In generaland everyday terms of common TABLE i manner sensethischaracteristic ofpopula.3 2. 8.I 4 67..9 9. This may best be ilIo 559. The population at first YEAST DUCED PER QUANTITYS IN PERHOUR HOU'RS grows slowly. NO. VOL. somc sort of size of the population.9y72-3 choose to stress.2. such as is afforded by the i.3 ways.3l5l6. I89. 4 This content downloaded on Wed.9 9 10c i6.8 14 I1.4 513.2.7 The yeastplantis an organism low in the i6 655.2. that the growthof populations of the most diverse organisms andcharacteristic course.3. An experimenton the Carison (3). Afterthis stage (b) (c) (a) (e) (d) of most rapid growth the population cent per increases ever more and more slowly.9.i is by the conditionsof the experiments. of the discussioninvolvesno theoretical implications whatever. processof cell division.4 64o.0o i8.5 growth of a population of yeast cells..9 and thenwane.3 we maywell take a relatively 34. Because of the thesecondcolumn theobserved (b) records minute size of the individualcells.7 44I. and of quantityof yeast.7 47. But this is a mattermerelyof detail.5.7 5o62.0 12.i scale of life. the data being taken froman QUAR.

(Data from Carlson. The figures values of the thiscolumnare the observed _ ~~~~~~~~~Asymprorr 665.and.that is the absolute amount of new yeast by the growthof the population produced in in each successivehour.) ratio dy/dt. they obviously form a curveshaped like a cocked hat or sugarloaf.. Thereafter as growth continuesthe absolute increment of new cells per unit of time decreases hour by hour until it almost hour. in proportion to the already attained size of the population. If the figures of column(d) are plotted. GRwT OP A POUAINO ES El /SO zso C1 16 1 1 1 10 14 1 18 14 FIo.as is correctly shownhere.whenthey shouldhave beenhours. the population has ceased to grow. As a naturalconsequenceof this trendof the first differences the relative growthrates given i column(e) are foundto be largestat the This content downloaded on Wed. as in figure2.M. vanishes. which maybe neglectedfor the present. for all practicalpurposes.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. TH. i. givesthe simpletimerateof growth. 3I. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . up to a maximumincrement per hour.which is attainedin this case somewhere about the 7thor 8th hour.gives the size of the populationat each successivehour equaas calculated froma mathematical (d) columa discussed tion to be later.0 600t MO~ $300 ~~~~~~~OR Fin. y denoting the size of population and t denotingtime in hours. through an error in proof-reading the abscissalunitswerestated to be days.the thirdcolumn(c). The last column(e) gives the relativerate of growth. low at the two ends and risingto a peak near the middle. in each timeinterval. THE GRowTH3OF A POPULATIONq CE-LL OF Y-BAST In the publication of this diagramin (8). In mathematical terminolin this column are the ogy the figures observed values of the double ratio _ dyforeach hour.

Austria. GROWTH INCRBMENTS OP YBAST POPULATION IN SUCCBssivz TImBINTERVALS Data from column(d) of table i France. so far as may be judged fromthe available census records. Java. here to review all the evidencethat this in the case of the world as a whole. is so. and of the fly Drosophila in their growthfollow this melanogaster. It has been shown populations of yeast. I shall not take the time the circles give the census counts (or. This has alreadybeen done in two estimates)of the populationsexistingat This content downloaded on Wed. 3). (d).Servia. Denmark. population growth. Scotland. These actually cases arepresented graphically such a wide varietyof organisms regarded are the United States (fig. 4). 5). to at least a first as a whole (fig. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . The phenomenon shown in column UnitedStatesofAmerica. is reached. 6). 8). Philippine Islands. Sweden grow.England and Wales.has beenfoundin recent In illustrationof this statementfour to be the way in which populations of here. New York City. and the creasestill the end of measurable years world as a whole. Although this relative rate of growthpresented in column(e) is the one most oftendiscussedby demographers in talkingabout populationgrowth. France (fig. growth BaltimoreCity. recentbooks (6. namelythat the absolute increment Belgium. Norway.in at least the following countries:Sweden. 80 K40 U) 2 FIG.it tells us much less clearly what is the real natureof the characteristic curveof such growth than does the simple time rate. with Hungary. and then de. characteristic curvewith greatprecision. of growthper unit of timeincreases time to a maximumvalue. that experimental of bacteria. or first derivative of the population growthcurvegiven in column(d). it has been demonstrated Furthermore statisticallythat populations of human beingshave grownaccordingto the same type of curve. 2.GROWTH OF POPULATIONS startof growthand to becomesmalleras the absolute size of the population increases. and the world as the characteristic. Italy. that it may now be fairly normal mode of (fig. In these diagrams approximation. Japan.. 4 6 8 /0 /2 /4 /6 /8 HouRs.

Q 5 X W5SEDEN I0 J3 - 780025 50 75 /19 FIG. In the case of the demographicunits listed above the census records do not UNITEDSTA TES /00 t75 NCE FRA 050 Uj) 40 CL 25 1800 20 FIG. THE OBSERVED AND CALCULATED GROWTH OF THE POPULATION OF THE UNITED STATES R YEA 30 j::20 6y ~~~~~~~ . 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 This content downloaded on Wed. the indigenous native populationof Algeria. THE 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.in which a the given dates.while the smoothcurves are the fitted theoretical curvesof population growth. And one cannotconduct with experiments human beings on this point. human groups the complete course exhibited by the yeast population just discussed. /750 7S /800 2$ 50 75 1900 25 5. But fortunatelyit has been possible to findone group of human beings. 4.if undisturbed. 40 60 80 /900 20 3.536 THE QUARTERLY REVIEW OF BIOLOGY sucha conclusion in somedegree probable. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . THE OBSERVED AND CALCULATED GROWTH OF THB POPULATION OF FRANCE R YEFA YEAR FIG. these having been carefully made by the French. as can be done with lower organisms. The available data only make cycle of population growth has been practicallycompletedduring the period for which census records are available.

birthrateslargerthan This content downloaded on Wed.ratesto the logistic curve. see also I9. and some of the mathematicalpropertiesof the logistic 8. as yet unpublished. first His pioneer work was forgotten. It is easilyshownthat ifbirthratesand death rates are assumedto remainconstant.OF POPULATIONS GROWTH 537 ofPopu. We have done a considerable amount of work. in size of populationwhich follows this particularcurve ratherthan some other THE LOGISTIC CURVE one.undervarious sequentstudents of the populationproblem. 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. Reed. and x = time of growth in hours. In i9Z0 the present writerand his 1600 colleague. For thewhole seriesof I9 observations the root mean square deviation resultsand the betweenthe experimental curveis only 3.is a valuable firststep. This demandsthat we shall push The equation to the curve which has the enquiry further. The covered by the Belgian mathematician. may be stated in simple form and are pertinent to the present discussion. $) 75 1900 6.and of the relation of birth rates and death consequentlyoverlooked by most sub. towards reaching an of the biology of the understanding process. zi1200 Z1000 S il"800 :Z 600 400 200 1750 75 /IO 25 FIG. because of their technical character.0. curve. THE YEAR OBSERVED AND CALCULATED GROWTH OF THE POPULATION O THB WORLD wherey = quantityof yeast.59. The possessionof such a curveas this. The equatheoretical curveis: tion of the fitted 66. The characteristic appearance. Certain aspects of it have also been discussed by Yule (z5) and Lotka (5).5355x e4-x896 (IS-). 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . Some of the results. in"0WORLD dependently hit upon the same equation Verhulst called his curvethe "logistic.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. and is illustrated in figure 7.5 I + . I shall not attempt to present the details of this analysis on this occasion. SO) in I838. What we want to know is how the biological forces of natality and postulates. Lowell J.mortality analyzedin my book TheBiology are so integrated and correlated here in theiraction as to lead to a finalresult lationGrowth (8).. further is a theoretical mathematical analysis Verhulst(i8. which is found by actual experienceto the course of populadescribeaccurately tion growth in a wide variety of organisms. at any values consistent with growthat all (i.e. without any knowledgeof Verhulst'spriorwork. but only a firststep. upon this subject. however." This usage we shall follow.

26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions .y X 44 l I t W 1 1Q o~~~~~~~~~~ t 1 W~~~~~~~~~~~~~~~t $MOIIIIk $11 NOWCOdOS This content downloaded on Wed.538 THE QUARTERLY REVIEW OF BIOLOGY _ _ A ~~~~~~~~~~~~~~~0 Xe I v -- - -- f * tt 8~~~~~~~~~~~~~~~~ i IVI ! 177N 1zC A_ _ A1 HtiH~~~~~~~~~~~~.

but the form is exponential. Nothing like the slowing down of growthaftera period. The absolute size of the definite universemay be small. it is tacitly. but implicitly. on a pinch. the test-tube or it maybe as largeas the earth. a b TIME FIG.mostof which could conceivably be inhabited.GROWTH OF POPULATIONS death rates) the population will grow accordingto an exponentialcurve. continueconstant that birthrates shall be constantregardless of the size of the populationalready attained.appears under this postulate.on to infinity. birth rates cannot In saying indefinitely. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . THE LOGISTIC CURVE AND ITS FIRST DERIVATIVE seen in the logistic curve. as in the case of which holds the yeast cells.which is 539 with this growth.byman.no matter whole group becomes. But in anyand every limit case there is ultimatelya definite to the size of the universein which any real populationlives. But plainly. But such an asfor any actual sumptionis unwarranted populationwhatever.with actual populationsof living organisms. All populationsof real organisms live in universes with LOGISTIC CURVEAND ITS FIRRST DERIVATIVE CURVE ASYMPTOTE K ~ ~ h O. assumed that the universein which the hypotheticalpopulation is growing ex- limits. pands proportionately so that each single elementalways has how large the plentyof room. 8.The rate of this progression may be slow or rapid according to the assumptionsmade. This consideration obviouslyalters the This content downloaded on Wed.

Furthermore ofwidelydiverse organisms. scribedand measured. that crowding.and arehuman without consideration of any other by the studyof experimental populations we are led rigorously of lower organismscan safelybe transvariableswhatever. etc. to the conclusionthat underthesepostu.Obthevariables. or any similar experiments on in the discussionof the worked out.physiologically reached beings. when it reaches a certain so widelyemployed forgenetic upon both of studies.and as we shall presently energy of my laboratory has and observational been devoted duringthe past five abundantexperimental years. has an adverse effect degree.lower organisms. 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. For it at once and knownconditions.Before undertakingto present some lying population growth. in fact. tion as a whole be observed. On the otherhand it mustalso be kept under carefullycontrolled management. natality and accountof theresults of theseexperiments mortality.. Its purpose the same in the two cases. 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. underseveral simplepostu.and the second. the two primary biological forcesunder.or The organismchiefly used has been the as it is technicallytermed. supportfor the view.viously in all such experiments the conmatical.540 THE QUARTERLY REVIEW OF BIOLOGY complexionof the case. They maybe is to observeat each stage the growthof or they may not. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . and mortality.density of fruit-fly Drosophila the form melanogaster. organisms concerned are less complex between the two first-order and psychologically than birthrate and deathrate. 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. as the growth continues beyond this can be particularly studied. between The net resultof this first. This content downloaded on Wed.density of population. does. It is to this mode hypothesis It is not an unreasonable a of attackupon the problem that muchof see there is the research priori. or mathe. if it describe if the same resultis. lates as to the quantitative relationships problemof humanpopulationsshould be neitherover-rated nor under-rated. and its bepoint they will get more and more havior under controlled conditions decrowded. as material population. composingit begin to get crowded. Not onlycan the follows that if the universesin which growthof such an experimental populalive are finite and inextensible. No conclusion ordervariabledensityof population. This assumption has been I wish to point out that the significance made and the theoreticalconsequences of these.with also moreuniform simple postulates as to the relations human populations live. To find out which is truedemandsspecial ad hoc the populationof some actual organisms.foundto appear in both experiEXPERIMENTAL POPULATIONS mental and human populations. Further the variables. attack upon the problem of ditions of life are not only simplerbut than any underwhich populationgrowthis to show that.ferred by simple inferenceto human lates the growth of population must populations. alternative small enoughto be capable of laboratory investigation.

aggregated of forces towardsincrease tending dition of aggregate wholeness are just as in numbers on the one hand. under diverse controlledcon.may be analyzedin experimental variables. To suit various theoristsit preeconomic and social factors are unim. these second. at any mortality suchas to makethegrowth of the oflivingthings. in develop.population rather follow the logisticcurve. As a matterof fact the populations of whatever organismsare. The elements end effect 2I-2.GROWTH OF POPULATIONS 54I in mind that the forcesof natality and the effectof these factors on human arebasic biological attributes of populationgrowthhas in thepast. although. natality and years the argument the very populations which have in fact are in a as as mortality. Whateverthe future may man today. by the host of economic growth.in the greatmajority a first-order variable in population of actual cases. menas well as rate. from it. Over and beyond faithin a priori logic as againstplain facts is the further one that this consideration of experience.sumablyoughtnot to. in humanpopulations. During the past few ing to the observedgrowthof a human thandeviate has been made that population are these two.account of" a lot of these second-order ditions. 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. shows plainly that human populations This content downloaded on Wed.in numbers on theotherhand. the past historyof the matter the majorityof observedcases at least.with natural increase by reproduction.been aggregates flies. populations of lower organisms.migration is theoretically the logisticcourse. This state. just truly they to a according or yeastcells.cannotpossiblyhave done with which the behavior of these so.aggregate wholes. upon population size of the their out behaviorarising of this con. The ultimate biological forces lead.of learninghas always shown. by that gesting pointsforstatisticalinvestigation wind-broken and spavined old stallion.and third-order variables. this growthis not sensiblyalteredfrom as we have seen. portant in their relation to population BenjaminFranklinlong ago pointedout growth.the logistic curvewith remarkable preciment should not be taken to mean that sion. excellentcase in point.0 it is impossible to detect in studyof humanpopulation growthis the the curveof growthany separateor disevidence that the steady onwardmarchof turbingeffect of immigration. because this curve does not "take variables. Wheeler. 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. The actual observed growthof and social eventswhich are supposedof thepopulation oftheUnitedStates follows logical necessity to affect it. The popuuals of one species as in those composed lation growthof the United States is an of individuals of another. Because grownwith greatprecision populationofflies of the much greaterease and precision logistic curve. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . This argument is rubbish. but in factit does.as compared important single elementin the biologi. and decrease true in populationscomposedof individ. logistic curvedoes "take accountof" all in their very nature. and behave in growthand other ways as in thesensethatit describes theintegrated such (cf. cally effective environment of civilized He was right. 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. All that is meantis that.4).

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.9 8o 30 5.8 2D o. The population can be carriedout.9 9 90 0.6 8.5 supply is used up. 3 6. There are two the thenexistingpopulationin pointsin connection with experiments thereafter of each bottle is carefully counted and this typewhich seemto need emphasisin work.and the individualscomposing the light of further to the bottleto carryon their that.7 8 7. to say that in bottles The resultis precisely here.1 IOO 10 IO I .3 logistic curve.4 290 ismadetoadd foodas the whichan attempt I.3 7. THE QUARTERLY REVIEW OF BIOLOGY have behavedin theirgrowthin the same way that experimentalpopulations of lower organisms do. as I have described in detail elsewhere (8). 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. This decreasegoes on at an experiments have alreadybeen accelerating melanogaster rate. Then quite The details of the techniqueof making suddenly thepopulationbeginsto decrease on population with the fly in numbers. and as is shown female-is added. At regular intervals graphicallyin figure9. and then an initial lates that birthrates decreaseand death groupofa fewflies-say one male and one rates increase uniformly at a constant linear rate. literallydo nothingfurther exceptto let nature take its course and to count the fliesat intervals. 2_0 2_. TABLE 2. and thereafter mentioned This content downloaded on Wed.0 4.7 3.3 I . 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.2. with truly remarkable faithfulness.8 3.3 6 6. The technical diffi6. The resulting growthof 2.6 6o 6.542.9 I3.5 7 5. This is shown in table 2.the foodconditions experiment of this type. The firstis recorded.0 8o 2_.9 40 70 4 cultiesof doing this satisfactorily with a i2L. 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.8 9.0 30 in large degree. 6o 8.7 40 I2-. What then happensis that the populationgrowsalong alogistic curvefora littlemorethanhalfof a comDROSOPHILA POPULATIONS plete cycle of such a curve. 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.6 5 50 Drosophila population are considerable.9 the population then follows a simple 0.2. The secondtypeof experiment is one in 0. It will suffice that which is to size definite amountsof food be expectedmathematically of measured on the postumaterial are placed. and need not be repeated vivorsleft.0 IO IOO I 3. until thereare no surDrosophila described(8).owing to the technical difficulties of it returned normallives until the next censuscount.9 50 caretheycan be overcome butby sufficient 70 8.4 3. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions .

while they were resident these newly hatched flies are counted food(9). At the end of threedays a censuscountis taken and the population to a new bottle. in any such bottle any daily. The observations of population size lie smoothlyon the logistic curve until the asymptote is nearly reached. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . This is accomplishedin the followingway. (FromPearl(8)) which I The thirdtypeof experiment. A2. and so on again containsnew fresh Meantime the lag bottles. 400 543 population.OF POPULATIONS GROWTH are definitely sub-optimal. The nutritional level of the bottle is not high. Supwith our pose a bottleA1 to be prepared A1. which transferred food. way for here reporton in a preliminary time. conis transferred new foodto thesameamount taining fresh as was presentinitially in A1. Whenever eggswhich werelaid flieshatchout from by the femalesof the adult population in that bottle. At the end of anotherthreedays a censuscount is again taken and the population again to anotherbottle.is so plannedas to have the the first food conditionsalways optimal both as to quality and quantity.. which the adult population are examined has left by transference. 350 300 R50 5 200 AsYMPTOTE 346. daily birthrate of the population. Also in which and to be seeded with an initial small each day the bottleis examined This content downloaded on Wed. A2. etc.and thesmoothcurveis thefitted areobserved The circles logistic. whichhas been (as births)and addedto the adultpopulanew synthetic to lead to lower tion in whateverbottle it happensthen shown experimentally ratesthanany to be. 16 /8 20 22 24 26 28 30 3 7 NOYVMBER 9 1/ 3 9. indefinitely. GROWTH oF AN EXPERIMENTAL POPULATION oF DROSOPHILA censuscounts.1 15C W so A I I I I I I I I I I 6 8 /0 OCTOBER /2 14 FIG. A3. Under these circumstances violent oscillationsof either birthratesor death rates do not occur. It only a little more than barely permits continued growth of the population. So then we get a recordof the birth deathratesandhigher other natural or laboratoryfood tried.

9 14.544 THE QUARTERLY REVIEW OF BIOLOGY this asymptotic level as long as the experimenterdesires.deathrate.4 2 I. I believethatthiscondition ofunstable equilibrium is.8 9. In fact thesewaves in the size of the population. A detailed account of these will be published.40 5.The general results reviewed. my laboratory regardingthe effectof Manyexperiments ofthistypehavebeen densityof populationupon natality and resultis thatthe mortality in Drosophila.0 I.2- IO.0 I. 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. I I2. 5 2.and the deaths recorded. populationfirst grows up to a maximum of these studiesmay be briefly or asymptotic Some years ago Pearl and Parker(ii) level.produced by oscillations in the birthand death rates. But showed that if a countwere made of the in thiscase the populationcan be keptat progeny (adult fliesor imagoes) produced 4. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . It has not so farbeen possible to deviseanymethod of holdingthesepopulationsin a steadystateat thelevel of the when thereis at all timesan asymptote.7 6.34 0. in partat least. The chief performed.6 curve can be completely accountedforon 2 I. OBSERVIED CALCULATED NUMBER OF PROGENCY PER FEMALE PER DAY I 2.5 47-75 relationsbetween three variables.8 4.6 This content downloaded on Wed.and (c) thereis complete registrationof both births and deaths.6 5. however.33 50 The basic postulatewas that increasing density of populationhas associatedwith populationas a whole.1 5.50 7. A striking result.1 5.2I. 8.8 2. 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.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.0 ingcertain simplepostulates the regarding I..2.I 6. 34.I 3.6 4.3 I. 90. 2.2 It has been stated above that the growth of a populationalong the logistic 10 3. it is hoped.6. 2._00 4 5 6 7 8 9 3 3.I mathematical theoretical grounds by makI5 2...32. The population it adverse changesin birthratesand death is dependent upon any given singlefood rates.5I 6.o2.02. 2.2-9 2-I. 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. just as in the second typeof experiment described above. Is thereexperimental evidencein supplyonly over a period of threedays.I I4.9 II. 9. 3.48 i6. THE EFFECTS OF DENSITY OF POPULATION the adult population is actually then resident. The population simply waves up and down about an averagesize.anddensity ofpopulation. are perhapsthe mostcharacteristic feature of of this particular populationexperiments type.70 I2.5 IO. 2. 2.6 io. birth 30 47-73 rate.I 2. abundance offresh food.62.5 3.74 4. with the optimalfood causallyconnected conditions.66 0. experiments some timein the courseof the nextyear. 18.

53 3525 d0-.32.8 z-0.32 4o.72. facts with extraobserved the describes denwith increasing theseexperiments. and uniform theresults densities ofpopulation..34 A-o. even in a Rarely.2 4 o.58 Qo. uniform air space.2-. 67 37-46 t0.8.31 37. and x denotesmean density and of the mated population (measured as a profound There is evidently fliesper bottle).0 I.-5I 40.GROWTHOF POPULATIONS amount in bottlesofuniform size.92. I7 II.2.o and the smooth The circlesgive the observations curve is the graph of the logarithmic equationdiscussedin the text. but with of food.o.3I 29. or physical per day declincs perfemale enyproduced obbetween closer one agreement get at first as density extremely increases.36 I9.I6 59. The total numberof flies(23.43 to.68 2. 42.50 80.4-2. 4o.6o 0o.00 75 85 44.49 39.65 53.36 30.54 .30 17.68 8.66 66.2. 5. varying werethoseshown in table 3. 6i h0. in therateofreproduction change regular that this equation It is at once apparent of underthe conditions of Drosophilag.oo8 x .2. I6. This content downloaded on Wed. O 2. give confidence The last column of table 3 gives the calculated ordinates of the equation. 36.66 37-47 t0o. These resultsare shown graphicallyin io.04 -t0.0.00 34-45 4o.The number sity does chemical experiment.95 MEANF JES P& FIG.4 16. I5 9.o.65 34.69 37.55 I5 2. by the method fitted to the observations of least squares.922) is large enough to progeny in the results.23 10 X o X 40 M m eoW 90 95 105 125 I50 .?IOo 2.85 37-. figure 22 545 rapidlyand then more and more slowly at higherdensities.0 34. ordinary precision.07 d0.60 =Io.93 DENSITY OPl WITHINCREASING OrDROSOPEILA TH:EMATED POPULATION Ix RATE REPrOtTcrioN 144-47 0 .88 o 17I7 A. logy = I.72. of progof population.77 3.5 35 45 55 65 2.36 94.46 .38 iii.6i 8 6. SHOWING CHANGES BOM OF 74. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions .0 4-0. I. wherey denotesnumberof progenyflies perday overa sixteenperfemale produced day period. 10.45 32.2.658 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 2 4 6 0 IO I2.

II. is upon fecundity (numberof Drosophila. 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. Here it is only necessaryto say that they show conclusively that the effect in populationsof primary ofdensity. 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. so farmenIn all of these experiments of densityupon birth tioned the effect 45 40 (4.This doubt has now been cleared up by an elaborate series of excarried out in my laboratory periments duringthe past year by Dr. of the Plainly the curveis the expression law relatingthese two variables. eggs laid).rate of and density. Elsewhere(8) evidencehas been presented showingthat the same kind of effect of densityis observablein populations of other organisms. The resultswill shortlybe published in detail. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions .546 THE QUARTERLY REVIEW OF BIOLOGY under different densities of population. I. So then it may be concludedthat in- servationand theorythan is here shown. The resultsof a large amountof experimentalwork regarding the effect of den- This content downloaded on Wed. Arata Terao. z). o20 5 /0 20 FIG. 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. io). 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. 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.

gardedas thedefinitive This content downloaded on Wed.on the one hand. and.and density and death rates.rangingfrombacteria and yeast to man. statistical. This is done in table 4.000 547 in figure ii. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . are found statisticallyto follow in theirgrowtha particulartype thelogistic.first. Experimental under lations of Drosophilamelanogaster controlledconditionsshows that in fact the relationsbetween densityofpopulation and birthrates.there is littlechangein meanduration oflife(or deathrates)with smallincreases in density. Only the broad outcomecan be presented here. and experimental study has brought to light the the genfollowingbroad factsregarding eral biologyof populationgrowth: i.while after death rates markedlyincreasewith increasing density up to an asymptotic limit. The figures for mean durationof life are necessarily determined by the values of the death rates at ages. A great deal more work needs to be done on the problem. The results are shown To summarize: A large amount of mathematical." It has been revisedfor publicationin the QUARTERLY REVIEW OF BIOLOGY. aftera relativelysmall initial rise. and this publicationis to be reform of thepaper. by small increases edly affected adversely in density at relatively low densities. ofcurve. Conference underthe title "The Biology of Population Growth. density of population. In condensed it was read at the WorldPopulation form in Geneva.OF POPULATIONS GROWTH sity of population upon mortality have recently been publishedin detail. One particular set of such postulatesis that it shall be that birthrates are markassumed. 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.which actuallyexistin suchpopulations. Other experiments have demonstrated that with extremely high densitiesof population the mean durations of life (or equally the death rates) approach an asymptote.and the investigations are being continuedin my laborapaperis to be regarded tory. The present only as a progressreport. while after a certain density is passed further increases produce only slight decreases in birth rates down to an asymptotic limit.birthratesand death rates. SUMMARY flies. In densities below 55. investigation of popu3. and the second-ordervariable. Mathematical investigation shows that a curve of this type is necessarily undercertainsimplepostulates generated betweenthe two as to the inter-relations first-order variables. The experimentsinvolved I3.. second. Anotherway of putting the same thingis to say that the death rates in these experiments increasedwith increasingdensityof population afterdensity 55 has been passed. graphically What the figures in table 4 show is that aftera densityof 55 fliesper one-ounce bottleis passed. I92-7. Populations ofthemost oforganisms diversekinds. which gives the meanduration oflifein days of Drosophila subjected to the indicated densities of population.that death affected rates are insignificantly by increasingdensityat relativelylow densia certain density is passed ties. 2. on the other hand. August 3I.the meandurationof life steadily decreases as density of population increases.

PENNIMAN. Pp. i92. Paris (Doin). ..2. (II) PEARL.II3-I2I. L. Roy. R. Deuxieme memoire sur la loi d'acde la population.5. I845. pp. Densityof populationand life duration in Drosophila. I-58. production Sci. theory PP.. 2. Vol. de Bruxelles. Ibid. (6) PEARL. On the influence gine-Leur Evolution...5. I-38. Science. I92. 83. 357-366.de l'Acad. 34-65.0) (g) PEARL..o.. PP. Vol. publ. Predicted Growth of New York and and its Environs). . 66-77. Nat.1) (IO) PEARL..3. Culturemedia for Drosophila. whichcontrolit. 307-32. PP.Corr. PP. 2-8o-i83. (2. B. studieson the durationof life. I92. i92. Jour. 88.. PP.88. YULE. Vol. Vital Statistics: A Memorial (I5) Volume of Selectionsfromthe Reports and Writingsof William Farr. Leur Ori- York (Harcourt. 6i. 653. PARKER. math. Biochem. Acad.2.C.. L.. Amer. The probable constants error ofcertain of the population growth curve. On the rate of PEARL. Vol. T. Metron. i8. . par A.. et phys. Jour.60.3- London. i885. der Hefevermehrung Ztschr. 2-89-3i8..Roy. D.4) (2. Soc. F. C. Vol..365-368. (17) Nat.. I92S. I-32. UDNY. 3I3-334. i92. Ibid.Bd. i92o0. The curve of population growth. M. pp. I92. Vol. 4. Pp. i92. I0-I7. Elementsof Physical Biology.R. I9II. L. Phil. and L. et Belles-Lett.4. Brace and Co... 64. III. (i) BROWNLEE.5.6. R. Pp. Stat. Skew growth curves. pp. (i6) (5) LOTrCA. Sci. Amer. R. Vol.). xiv + 2. 63.6.I2--i9.). Soc.. Humphreys. A. Ibid. and S. pp. its Environs.0. (2. ofthepopulationoftheUnitedStates growth of density of population upon rate of re- . Acad. d'accroissementde la population. I9I9.5. Vol.75-2. T.548 THE QUARTERLYREVIEW OF BIOLOGY LIST OF LITERATURE J. 433-440. .. I9I3(4) FARR. ii.3. Notes on the biology of a life table.. X. 2. 42. Quetelet. Proc. Vol. REED. Knopf. (3) CARLSON. 2. The growth of population This content downloaded on Wed. Recherches mathematiques sur la loi (7) Proc. New Les Societes d'Insectes.D. xii + 468. Vol. pp. (i8) P. Morph. 3. I847. Proc. I92.T. 6-I9. R. Vol. F.). I92.. Soc. W. J. W.5. . J. Nouv. 6o.) Social Life Amongthe Insects. Editedby Noel A. .B.3) . G. I92.2. pp. undGrosse (I4) UberGeschwindigkeit in Wiirze. Discussion. (2..4. i6-12. A. I92.. . I92. ALLEN. Vol. T. pp. Studiesin Human Biology. Pp. I92. pp. I92. pp. R. I92. PARKER. and the factors Stat.). New York (Alfred A. Ibid.7. R. Hyg. Noticesurla loi que la population suit dans son accroissement.37-140. and S. PP. (2. Nat. Baltimore (Williams and Wilkins Co. . On the mathematical theory of population growth. (19) (8) The Biology of Population Growth. 2. 2.... on and its influence medium A new synthetic densitiesof population. . Experimental XI.4. mem. at different fertility Amer... 8. Vol.. Jour. I838. 6. New York (Plan of New York . pp. 8. xxx + 460.pp. des Sci. Density and death-rate: Farr's law. J. Roy. Inc. MINER. pp. and W. PP. xxiv + 563. VERHULSt. The ant colony as an organism.S. M.L. Emergent evolution and the social.6. 26 Dec 2012 20:54:46 PM All use subject to JSTOR Terms and Conditions . Vol. D. A further note on the mathematical (X) of populationgrowth. representasince I790 and its mathematical (I3) tion. WHEELER.5) (I1) in Drosophila.. 82. (2. Baltimore (Williams and Wilkins Co. Ibid. I92. 57. PP. croissement PP. Jour.

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