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of Biology, Vol. 32, No. 3 (Sep., 1957), pp. 217-231 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/2815257 . Accessed: 26/12/2012 21:20

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of the organism.but in fact hardlyexplored question. Thereare. per square meterof body surface. and to some answer to metabolism the seemingly trivial. the chemicalcomposition or the enzymecontentof the cells-we excretion. Bergmann and Leuckart. activities.in spite of the fact that the animals under comparison belong to different orders. factordetermining heart or respiratory we take total metabolism. of even a simple What we call growth complexphenomenon organism is a tremendously cytological. "Why does an organismgrow at all. It appears with dogs of different that metabolic rate per kilogramdecreases. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions . that is. after a certain time. The comparisonof metabolic rates in mammals led Rubner to the contentionthat warm-blooded animals produce daily about 1000 Cal. physiologidiversity cal mechanisms. 3 September.. A classical example is provided by Rubner's experiments size (Table 1). Heat outputtakesplace temperature thebodysurface.metabolismis calculated per unit of body surface. It goes back over more than a hundredyears to the time when Sarrus and Rameaux. and Richet noticed that weightspecificmetabolicrate. 37?C. vary always will findthat theycharacteristically with body size. thesamenumber of caloriesmust be producedper unit surfacein orderto maintainthe body temperature constant. byRubnerin terms All warm-blooded animals heat theirbodies to a of ca. To give 217 This content downloaded on Wed. approximatelyconstant values appear. _ HE work reviewed in this paper is just one example:pulse rate in mammalsclosely to the Y powerof body weightover corresponds seven orders of magnitude.in manyphysiological the absolutesize of the body is a mostimportant Whether the rate of processes. does its growth come to a stop?" QUANTITATIVE RELATIONS BETWEEN BODY SIZE AND METABOLIC RATE T BYLUDWIGVON BERTALANFFY INTRODUCTION and University Los Angeles Mt. If. through Hence. 1949). however. viewpoints.and are adapted to all sorts of climate and ways ofliving(Fig. NO. fromthe biochemical. considerable difficulties it may be In orderto begin this investigation. Adolph. and morphological certainaspects that are amenableto quantitative analysis.VOL.and so forth(cf. rate. the intensity of as measuredby oxygenconsumption metabolism or calorieproduction ofbodyweight. 1).and such an approach appears to lead to some insight into the connectionsbetween and growth. however. per kilogram decreases with increasingbody size. emphasizedthat. Sinai Hospital. their metabolism organisms. physiological.This is the originof the famous surfacerule. ofSouthern California.from a dwarf bat weighingsome four grams. I957 THE QUARTERLY REVIEW of BIOLOGY QUANTITATIVE LAWS IN METABOLISM AND GROWTH beaimed at establishingconnections aspectsofliving tweentwofundamental and growth. There are. 32. to an elephant of 2000 kilograms. adaptations to certainenvironments. Biological Research. The relationbetweenmetabolicrate and body size belongsto the classical topicsof physiology. and why. which was explained ofhomeothermy.however. this being true even thoughthe the organisms comparedin such respectsshow a in their tremendous anatomy.

7 19.). in the series of mammals. 500kq IOQOV9 200O kg BODY WEIGHT FIG. Brody. Weight Cal. Kleiber. Dimensionally. mammals are almost solely taken into account (e.F. This volume is equal to stroke volume (S) X frequency (F). i. The Dubois is: S = kW0425 formula X L0 725.8 61. in kg. however. production per kg.In the case of man.but a simplemathematical applied. sq. Hence: S.218 1000 THE QUARTERLY REVIEW OF BIOLOGY ~~~~80O IL L co DD OVS~~~~~~~MQMTO-' P o ED6Eh'0G RABBIT = 3 C X 4W" !50Q ioOs - -~~~~~~~~~~~~~_T 509 I g kg SW I tk!g -A SS X ctS0O .39 5Sg log EUOSELjEP 50k(s l0kg . the W! rule. If two bodies are reasonablysimilarin shape. In a rough first approximation. cW0 725(0 The relationbetweenmetabolicrate and body size can be studiedeitherintraspecifically. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions . by animalsofthesamespeciesand different comparing body size. However..the Dubois formulais identical with the surface rule.2 34. in particular.Here thesomewhat thyroid more complicated Dubois formula is applied.7 30. body weight)to the elefactorin the controlof pulse frequency.. pulse basal may be assumed that the volume of blood transported per minute is proportional to the basal metabolic as the oxygen consumed must be transported by the blood. Hence.8 1191 1047 1141 1082 984 in measuring the outer surfaces of animals device can be exactly.per production Cal. 1947.e.6 40. 1. and otherdifferences absolute body size is the ecologicalo simplirication in a rangefromthe dwarfbat (4 g.3 45.g.2 23.5 57.28. 1950). S may be taken as proportional to body weight (W).in orderto diagnose disorders and the like.3 44. In considering the quantitativerelationbetween metabolic rate and body size.e. by comparing adult animals of different species. A grave objection can be raised against the surfacerule as foundin textbooks of physiology. homeothermic vertebrates and. body surface 85. and = = C'W a =-.2 1909 1073 11. We are at present mainly concerned with intraspecific comparison. i.0 17. presupposing geometrical similarity. the surfaceareas of geometrically similarbodies can be obtained by multiplying the 23 power of the weightby a suitableconstant.4 3. or interspecifically. The metabolic rate follows interspecifically. and therefore its square has the dimension of a surface. lengthL = cW'.physiological. = CW. phant (2000 kg. ALLomETRic DZEPENDENCE OF PULS5E FEREQuENcY ONBODY WEIGHTIN MAMMLS It rate. the gross over figmreshows that the allometry Notwithstanding whichneglectsanatomical. this can 33) = bW be written: S = kWO-420. their surfacescan be expressedas a 23 powerofweight.1 6. As. the determination of the basal metabolism is a clinicalroutine. 1945.This is seen in the ofMeeh: well-known formula S =bW (1) The surface rule of metabolism accordingly states that the basal metabolic rate is proportional to the /% powerof the weight. m. ModifiedafterBertalanffy TABLE 1 Metabolism in dogs AfterRubner (1902).the case ofmammals is by no meanssimplebut rather This content downloaded on Wed. dominating (1951a). sincethecubicrootofthevolume or weightis a lineardimension.. The constantof pulse frequency. Krebs.'.

Oxygen consumption and certaininvertebrates. On the otherhand.etc. in an animal of double size is simplydoubled. metabolic whereM is the metabolic rate per unit time. theconstant per unit rule. 15 33 50 100 160 Weight 3. types.5 54. If a = 1 or the slope is 450. its oxygenconsumption the surface metabolic rate is proportionalto weight. of thatof the total animal.0 49. physiological.2 7. biochemical. values are taken. clams. weight. many familiar conclusions and explanatory fall flatif not onlymammalsbut also hypotheses are vertebrates and invertebrates poikilothermic necessary It is therefore takeninto consideration. the obtaina straight a. to considerthe problemon the broaderbasis of A considerablepart of comparativephysiology. factor in the relation between body size and intraspecifically as well as interspecifically.et seq. to individuals of different evolutionary data.development. Dixippus morosus. metabolic rates.2 Cmm. is intricate.but the explana.the equationbecomes: problem Here the The second type is quite different. in the author's thisworkhas been carriedthrough laboratories./hr.2 53. that is. the experimental in 1941b.one more In orderto understand is necessary.covering tion given by Rubner is too restricted.in weight-specific Correspondingly.0 5. 3. but remainsconstantper unit surface. we certaininvertebrates. themetabolic ratefollows metabolic rate in the sowbug.Other groups belonging to this poikilothermic tion. to a surface RepreThat is to say. M =bW (4) metabolic rate is proportionalnot to surface so oxygenconsumption area.2 15. growing in thefollowing way: written metabolicrate is proportional In thefirst type. three different ways of This is a special case of the so-called allometric metabolic fornula (Huxley.C02/hr. vertebrates of a wide application. the As can be seen. Ifa = 3.decreasewithincreasing and the slope of thelogarithmic plot is negative. comparingimagos of different 1.Subsequently 1 > a > 23. (2) metabolictypesexistwithrespectto the relation rate and body size. Direct proportionality of metabolic rate to weight we can summarize is found in growinginsect larvae and hemimeAfter these preliminaries. stick.): (Bertalanffy. Armadillidium.8 51.and a and b are constants.as we shall see presently. line the slope of whichindicates and ascaris. there are many classes Finally. Thus we come to the statement that several M = bWG. The surface rulealso holdsforpoikilothermic Table 3 shows metabolic rates in the walking The rule is. With weightdecreaseswith increasingbody size. if it will be seen that sowbugand companyreveal If weight-specific metabolicrate per unit weightis plottedinstead quite a bit about human growthas a central ofphysiology. against body weight double-logarithmically. and rate. as well as interspecifically. amount can be distinguished. an animal fourtimesas large is quadrupled. sizes or to be that is. but to weightitself. 200 174 144 112 94 Perg.that is. Perunitsurface (WI)/ 48. metabolic annelidssuch as the earthworm.The dependformula mathematical ence of the metabolicrate of an animal on body in the equation: size can be expressed TABLE 2 CO2production pallasii ofArmadillidium (Temperature 21?C. and as was emphasized. related species. Table 2 presentsone example.6 hr. results in the followingway tabolous insects. for in wide range. After 219 in mg. theseresults. all body sizes and the entire animals thereis no thermoregula.2 11. an intermediary case obtains.per gramand hour appears to be constantover a therefore.QUANTITATIVE LAWS IN METABOLISM AND GROWTH Moreover. as a generalrule. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions . and thus the latter cannot be the basic type are land snails of the familyHelicidae. on body size foran enormous dependence allometry. 2. of animals in which the surfacerule does not hold. in the thirdtypemetabolicrates are This content downloaded on Wed. if metabolic rate is plotted sentatives of this type include fishesbut also such as crustaceans. to intraspecific of morphological. o? log W (3) log M = log b + or the 23 powerof theweight. between In view of what was said previously. This formulacan further animalswithin one species.) MUller (1943b). 1932) which expresses the dependenceof the metabolicrate on body size this classification applying.three W the body weight.

250 236 243 252 245 242 TABLE 4 Oxygen consumption ofPlanorbis sp. Recent investigations of the Ludwiglaboratory (Ludwig. the ofmetabolic rate to bodysize.What can be shown.henceits validityin fishand certaininanimateclasses.9 25. There is no doubt that energyexpense for thermoregulation forms a considerable part of the total metabolism in homeothermic animals. thedecreaseof metabolic rate with increasingbody size may be due to intrinsic in themetabolism ofthecellsofsmaller differences based upon cellular or upon organismic factors. So does basal metabolic ratein theinterspecific comparison of mammals. and pulse rate. for example. different or recall ascaris. pulse rate decreases approximately proportional to the % power of the weight(Fig. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions . Anotherinterpretation assumes that the surfaceruleis based upon theanatomy and physiology of the circulatory system.and in factis moreaccurately established.9 5. The most familiar one has already been mentioned. 1956. rule? (Cf.the clams. Table 4 gives data and filatworms fortheramshorn snail.the degree of capillarization.and showsthatitsmetabolic rate decreases with respect to its weight. 02/hr.0 30./hr. The supply of oxygen and nutritive materialsto the tissuesis naturally a function of the intensity of the blood current. first. it may be regulated by factors present and active onlyin theorganism as a whole. As has already been indicated. the frequencyof heart beat. namely. 1956) give some supportto the hypothesis proposed by Ludwig and by Bertalanffy (1951a. and Pirozynski.but withrespectto its surface increases area.if adult specimens of corresponding species are plotted (Brody.e. Kleiber. 1956. Perg. This content downloaded on Wed.8206. The latterdependson factors such as the size and stroke volume of the heart. thealternative whether of metabolicrate on body size is the dependence That is to say. 252f. We must admit that we withfurther references). Bertathe surface form. in interspecific comparison"fromthe mouse to the elephant".what is at the basis of the relationbetweenmetabolicrate and of its most familiar body size and. the surfaceof tracheasin insect larvae developsproportional to body volume.2 154.2 7. There seemsto be. Weight Cmm. Kienle and Ludwig. in cold-bloodedvertebrates and even in certain invertebrates wherethereis no thermoregulation. and the like. 1945. 1947.Remember. 1951a.4 69 65 56 Perg. 30-3558-6290-100140 190-200 2. 1). Bertalanffy 1953.0 28. is that however." i. there are rather strict quantitativerelationsbetween body size. thermoregulation. in mg. where the circulatory system is completely fromthat foundin vertebrates. Thus. and larger individuals.as was shownby Sattel (1956) in Bombyx mori. Per unit surface 22.3 3.A few minor need elucidation. 02/hr.this explanation cannot be generalsince the surfacerule also applies. mentioned are typicaland characThe relations of the species concerned. Let us start with the organismic hypotheses.hence the proportionality of metabolicrate to weight. 8 130 250 450 630 850 Weight 2.5 between proportionality intermediate to weight to surfacearea. hemodynamics a general explanation. p.220 THIE QUARTERLY REVIEW OF BIOLOGY TABLE 3 Oxygen consumption ofDixippus morosus (Temperature 20?C.6 60. do notknow. the diameterof the blood vessels. cannot offer However.7 113. Kleiber. but in generalit discrepancies can be said that the "metabolictype. (WI)/hr.3 9.1 52 48 27. Table 5 gives a teristic survey of available observations. To this type and proportionality belongsuchpond snailsas Planorbisand Lymnaea like Planaria.) After Bertalanffy and Muller(1943).However.1 26.) that the "metabolictypes" are connected withtypes of respiratory apparatus. lanffy. in mg.Gill-breathing animalsappearto follow the surface rule./hr. (Temperature 23?C.) After Muller (1943a). or else.6 Cmm. INTERPRETATIONS OF THE SIZE DEPENDENCE OF METABOLISM We now come to the question. metabolic rate. Sattel. which has no blood circulationat allanimalswhosemetabolicrate nevertheless follows the surface rule.is a physiorelation of the species or group of logical characteristic speciesconcerned. On the otherhand.. the explanations usuallygivenare insufficient. in particular.in which case it should also be foundin the respiration of tissuestaken out of the organism. 1947).

1943b Will. 1943 Bertalanffy Fusser and Kruger. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions . Planorbiscorneus Planorbiscorneus Isidora proteus Pulmonata and Operculata. and Maller.]?) Weight Weight Surface Surface Surface Surface Surface Intermediate Surface Probably surface Surface? Weight? Intermediate Surface Weight Weight Weight Surface Surface Surface This content downloaded on Wed.1928 Kramer. 1952 Ellenby. Eisenia foetida MOLLUSCA Lamellibranchiata Anodonta cygnaea Dreissensiapolymorpha Prosobranchia Lithoglyphus.1951 1952b Krywienczyk. W (weight) or intermediate PLATYHELMINTHES Dugesia gonocephala NEMATHELMINTHES Ascaris lumbricoides ANNELIDA Lumbricus sp. Brand. 1943a Kittel. Chilotrema.1934 Weymouthet al.1919 1950 Ludwig and Krywienczyk.TABLE 5 Relationbetween metabolic rateand bodysize Species Reference to proportional Respiration W2'8(surface). 1952 Muller. 1930 Wolsky.15 species intra-andinterspecific Helicidae and Cepaea (interHelix. 1944 Thomas. Cyprinus.intra-and interspecific molitor Tenebrio PISCES Lebistes reticulatus Various species (Scorpaena.. 1952b Krywienczyk. 1943 Bertalanffy Kruger. 1943 Bertalanffy and Muller. 1941 and Muller. vivipara Pulmonata Lymnaeastagnalis Lymnaeastagnalis Lymnaeaacuricularia Planorbissp. v. 1948 Liebsch. 1943 Bertalanffy 1948 Jan6aroik. 1943b Kruger. 1943 Bertalanffy Fusser and Kruger. Bertalanffy Muller. 1943 Bertalanffy Jost. 1952a Krywienczyk. 1953 and Krywienczyk. 1952 Will.1952 Weinland. 1943b Will. 1951 Kalmus. specific) Cepaea vindobonensis CRUSTACEA Branchiopoda Daphnia pulex Artemiasalina Isopoda Asellus aquaticus Asellus aquaticus Armadillidium pallasi Porcellioscaber Oniscusasellus Ligia oceanica Decapoda Astacus astacus torrentiuin Potamobius Pugettiaproducta Homarusvulgaris INSECTA Hemimetabola Dixippus morosus Holometabola Various species. Nolan and Mann. and Muller.1934 221 Intermediate Surface Weight Surface? Surface Surface Surface Intermediate Intermediate Weight? Intermediate Intermediate Intermediate Intermediate Surface (high tenperature[30?C.) REPTILIA Lacerta and Muller. 1954 Muller.1951 1952b Krywienczyk.1940 Muller.Abramis. 1929 and Muller. Paludina fasciata and P. etc.

however. Wey. is due to a corresponding is measured brain by Vernberg and Gray (1953). 2 illustrates. in interspecific are found in comparativemetab.Bertalanffy. are relatively whichis particularly imporlarge animals. tion of glutathione (Gregoryand Goss.l 02/mg. are irregular weight tionedhas been stimulatedby the investigations the same speciesand different The ques. satisfactory is not parallelin the variousorgansand. of cytochrome This content downloaded on Wed.What one may expect Martin and Fuhrmann. is different in intraStill anotherexplanationof the surfacerule is as betweenrat tissuesfrom comparisons. factors usually contemplateddo not offer a explanation.. 1955). We have alreadysaid that the organismic observers mouth. have established(Kleiber. lawsto be explained on growth oftissuerespiration is a So we have to assume factorswhich.animals of different withincreasing composition by callyactive" organssuchas theviscera. in the various organs consistent tive estimate(Bertalanffy to account with. can be illustrated It is by the action of thyroxin. main organsof the rat have been investigated and Pirozynski (1951. however. a highpercentage of body more oxygenand are respon. olism as classifiedin the metabolictypes men. but which do not In interspecific fromthe mouse show up in the isolated tissueused forWarburg species of different sizes.declinein thediaphragm. 1953) decrease of Qo. (Bertalanffy to the horse. 1943). ofmetabolism and Pirozynski. chronichormonalconditions c (Rosenthaland are manifestedby significant Patru'sev.1937). 1953). dry weight/hr. 1941. considerableamount of work has recentlybeen content From this it would appear that genetic. injectionof thyroxin thesurface way. apparatus. based upon anatomical or chemicalchanges in specific body size and age. Similarly.a decreasewithincreasing in spite of many efforts In a corresponding made. 1952). respiration as testes by Homma (1953).is less thanwouldcorrespond into the animalin vivo. On the otherhand. with increasing determination 1951. one of average Qo2withincreasing from body size is found of innerorgansis verydifferent growth that it in brain. 1950. As Fig. Field. and a marked is no systematic So there 1951a). a decrease of Qo. lung. of cytochrome changes of the oxidase (Kunkel tissue Qo. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions .222 THE QUARTERLY REVIEW OF BIOLOGY the presence and Campbell. and largerin small as comparedto Bertalanffy etc. and heart.administration of thyroxin to a tissue in vitro. So it can be assumed that they skeletalmusculature. weight-specific metabolic rate with increasing ofbasal metabolic fortheactual variations of body size.thebrain. to thevarioustissuesor are absent. . These results have been essentially in terms Now we come to the interpretations This amountsto sayingthat confirmedby other workers and with other factors. (cf. oftwo typesof respiratory The picture. controversial of the case. that is.1953). and Kleiber 1942.Fried and as Qo.partlystimulatedby our in the enzymatic differences just as we may hencespecies-specific.comparisons. the tissues...within tionofthe size dependence of regulate the respiration one. 1933. and responsiblefor the decrease of the shows that this factoris not sufficient rate. Differences. This decrease. "Metaboli. a slight decline in and so it is improbable organto the other.liver. but the statementsto follow the intact organism. nectedwithrespiration. in theteleost rule.and Pirozynski. in the heart muscle of the guinea with increasing decrease in the pig by Wollenberger and Jehl(1952). and kidney.and done along these lines.tant because it forms consumerelatively and Estwick metabolic mass. was studied by Bertalanffy sible for the higher weight-specific no significant decline rate in smallerorganisms. A quantita.able to reproducesatisfactorily such as in the concentra.Bertalanffy. and in rat Tissuerespiration oftissues. Seven body size. intracellular metabolic rate materials: in growingchicken by Crandall and the decrease of weight-specific size. determinedwith the Warburg apparatus.1951).activity and Qo. However. as a number of Schmidt-Nielsen. nobodyhas been this effect by an body size was found in enzymaticsystemscon. but ruleor the 3 powerruleofmetabolism.withrespect hereafter. can yield the simplerelationof the surfacerule skeletal muscle. as expressedin the surface Smith (1952). to easy to induce an increase of metabolismby as a generalrule. of succinodehydrase and cases wouldresultfrom Intermediate malicodehydrase (Fried and Tipton. A Tipton (1953) did not find a decrease in the of respiratory enzymes. own work now to be presented. ranging and Pirozynski. body size is generallyfound. oftissues also say that the interest in animals of however.the relative (1953). as has been shownwith tissuesfrom Drabkin.the sum total of whichis the metabappear to be a fairpresentation comparison of mammalian olism of the entire animal. Krebs.

in the sense defined. can be considered out by thefacts. v and K are constants of anabolism and catabolism respecm and n indicatethat tively. Now as thereare different metabolictypes. TIssuE RESPIRATION OF TIE WHITE RAT IN RELATION TO BODY WEIGHT Qo2 = . so long as building up preto take the metabolictype.at present. Indeed. the Ottawa study was startedwith this Let us startwitha ratherobviousdeliberation. BODY WEi6rT IN G. in consequence of a and the situationwould be much more generaltheory of affairs. the originalpaper. so-calledmetabolictypes can be distinguished by virtue of the relation between the metabolicrate and the body size. definite and strictconnection betweenmetabolic The writer does not feelhappy about thisstate types and growth types. among the various animal classes. of the anabolismand of metabolismin general thus catabolismof the buildingmaterialsof the body.there are also different growth typeswhich are distinguishedby the course of growthas expressedin In words: The change of body weight W is given by the difference betweenthe processesof building up and breakingdown. even this cautious attitude leads to equal. After Only regression and Pirozynski Bertalanffy (1953)./hr. vails over breaking down. Animal growth which.and the exponents the latterare proportional to some power of the bodyweight W. we considerit This content downloaded on Wed. size-dependence remains rather We have. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions .~~~~~~~~~~~~~~z 6 t--Y?US 4 31 a 1o 20 30 40 50 60 eo MO no l_ 300 400 J or VARious ORGANS FIG. based. There will be growth unsatisfactory.drywt. ofgrowth whichestablishes rational if a straightforward satisfactory relationbetween quantitative laws of growthand indicates the the decrease of the weight-specific metabolic physiologicalmechanismupon which growthis rate and the tissue respiration could be found.1954). expectation. 2. problem METABOLIC TYPES AND GROWTH TYPES It has already been stated that.It appears dwarf lack somatotrophin mice.QUANTITATIVE LAWS IN METABOLISM AND GROWTH 223 15 7.dl 02/mg. was not borne firstindicatedby Putter (1920). Obviously the growthof any organismis of an enormous whether complexity. a resultof a counteraction of The explanationof the surfacerule and of the synthesis and destruction. steady state if and when both processes are However. animals and in pituitary thegrowth hypophysectomized curvesoftheseveralspecies. for individual data and statisticalevaluation cf. We may expressthisin a generalformula: inferences certainremarkable with respectto the -KWn. dWIdt = nWWm (5) of growth.unfortunately.which (Bertalanffy that we have been successfulin establishing a and Estwick. lines are shown. the organism reachesa as an empiricaldatum of the species concerned.

componentsneeds. 1949). rule.So we may put. that of it is directly proportional to weight. 1951a). So far as the rate of allometricregression the formation line has a slope of 23. 1953. oftenin a quite unexas demonstratedby the isotope techniques. organism. The experimental results indicate levels. W = {87/K . and degradative processes insensitive to smallerdeviationsof the exponent grossresultofsynthetic the law of allometry.what is called the basal metabolicrate blocks such as amino acids.according to the surface highrate (cf. So we may assume. We shall not bother with the mathematical have shownthat elaboration. is. but the growing processes. The catabolic lism. 3 gives metabolism itself maintains theanimalorganism and growth in the small 1947). there we can state quite definitely that a is a lawfulconnection Nevertheless. froma biochemical. of course. facts (cf. hand being replaced by way of resynthesis in the case of the surfacerule the of new cells.1956.W2 -KW.and thisprediction Biochemically. mathematicalconsiderations an overallformula (Bertalanffy.the continuous buildingmaterialas it takes place in any living growth type of an animal from its metabolic thismeans the turnover type. n fromunity.whatever age.measuredas oxygenconsumption. F. D. it means the renewal of cells. pectedway. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions .If bodyweight. rule. But this assumption therateofall physiological The solution approximation in a first ofequation5 (n = 1) is (Bertalanffy. As will be reand membered.Thus we can predict the processesmean. sugars. general The case is somewhat different withrespectto what has been found in the dependenceof the anabolism. for example. the solufirst approximationand based upon various tion of the growth equation gives theoretical This content downloaded on Wed. Bertalanify. Our equationstates that the 1941b) show that our basic equation is rather lies in the following. that.and morphological physiological.the outcomeof innumerable and on theotherhand energy which. anabcertain organismobeys. and to a and so forth. Remember. Both can be taken into account as organismundergoeschanges at the biochemical. In a first is proportional type. the law of growth unexpectedly assumes the and Leblond.224 THE QUARTERLY REVIEW OF BIOLOGY morphological.oftenat an 23 power of weight. the surface olism. curvesare concerned. Not only this. building metabolism basal or resting Of course.so faras higher cellular. However. in fact. Accordingly. betweenrespiration. to 1. as a far as the growth catabolism is concerned. ofourbasic equation expressedas a power functionof body mass. Lebistes reticulatus. has provedto be correct and particularly materials of proteins. respiration to the found in many tissues and organs. let us say. On the other or any otheraspect.phosphates. Leblond and Walker. Leblond and Stevens. can be expressedas a functionof the -' the dependenceof anabolismon body weight. The synthesisof high-molecular cell of the intact animal.the justification and thesimplemodelit implies hand. as Cytologically. sucha simple.and on the other in the log-logor allometric plot. because at least mucheasier.[vq/K. as well as cells being con.and growth which works out in the folof the lowingway. on the one hand. aquariumfish. large extentunknownprocessesof intermediary in aerobic animals. is provided by oxidative metabolism.but show immediatelythe results. in a largenumber of building of cases. Storey form: and Leblond. chemicalcomponents are presented tinuallywornout or degraded. Leblond school is given in Bertalanffy.1951. that the rate of metabolism its size or developmental The exponentn in our basic equation denotes entireanimal. physiological physiological. limiting factors. This makesthesolution is justified.thegrowth laws forthe organism in question loss of follow automatically. a dW/dt . in 1941b): hitherto can be expressed investigated processes allometric or power formulas (Adolph. (7) table of the rates of cell renewalas foundby the 1957). The isotopeand othertechniques in a so-called Fig. that the rate of these processescan be siderableloss of generality. Bertalanffy following 1948.Wo (1-m)]e-(1_-tn)Kt}1-m (6) The intrinsiccomplexityof the phenomenon concerneddoes not preclude it from following withWo = weight at timet = 0. dynamic or steady state (Schoenheimer. animalsare concerned. that is. Metabolicrates. law. powerofits respective form thatvalue whichis experimentally we insert to define theprocesses foundforthe size dependence We have nowmoreclosely of resting metaboappearing in our basic equation. withoutany confollows withinthe organism n equal the exponent that is.

for the surfacedependenceof respiration surplus remaining for growth decreases. for example. 3. Guppy(Lebistes reticutatus).such as the gill-rakers said that in certain animals. increasesin size. 1920. 1957). Hence. maywellbe adaptable curves(b) in the Metabolicrate (a) and growtlh to otherpopulations)is beyondthe scope of the Growth curvesfor. length growth.commun. hensivetheoretical model of the dynamics of exhas beendeveloped.--40 ---about one-third of the finalweight. Discussionof this populationmodel FIG. 1954.. surface-proportional anabolism prevails work.QUANTITATIVE LAWS IN METABOLISM AND GROWTH 225 200 __ _ main characteristics. we have to insert1 for the exponent m in C'. This is themostcommon form ofgrowth curves. its surfacesincrease Let us see whathappensin thiscase (Fig.but its volume and mass with the third will give a line witha slope of 45?. calculated to equation (7).). and theyhave been adopted 120 6-30 in applied biology. growthis sigmoid. Beverton and Time in weeks Holt. l. over weight-proportional catabolism. The approximatelywith the second power of the log-log rate againstbody weight plot of metabolic length. Fisheries and Food at -0 ?' 2 I 0 Lowestoft (England). withthe possible exception A compreofthehake (Wimpenny. the same appears to be true for assimilating Now we come to the second type. After Bertalanffy and Millthat examination of the variousgrowth functions according proposed led to the conclusionthat "von Ber..surface. ploitedfish populations where-0 0 in growth of the speciesconcerned is represented 12 10 6 0 4 2 8 by equation 7 (Beverton. with certainrestrictions. .the a) -0 curves for weightgrowthand linear growthare 4o 0~ The curve of weight characteristically different.with a point of inflexion at . so long as the animal is tionof mathematical analysiscan be foundin this small. respiration does not follow the surface If a body. classes and also. talanify's growth equationis the mostsatisfactory been developed"(Bever. d: present review.the more the et al. eventuallya steady state will be reached where The quantity ofplankton consumed by thesardine anabolism and catabolism balance each other. It appears that the "Bertalanffy growth equaE E 00 b) 125----_It tion" is widelyappliedin international fisheries. of shape. and the A relation similarto that stated by Bertalanffy animal grows. IHE FIRST METABOLIC AND GROWTH TYPE (which. in a numberof invertebrate foundin fish. This is what we actually find Weight In mg. apartfrom fisheries.It shouldbe mentioned. curves with the following First. Ampledata as well as descrip.weightgrowth. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions .weightis continuallyshiftedin disfavorof the ofanythathave hitherto ton and Holt. and and growth comesto an end. in mammals. has been foundto fitthe commercially exploited fish of speciesstudiedby the Fisheries Laboratory 80 4~~~~the Ministry of Agriculture.Consequently. however. 160 40 -The validityof these growthequations has been shown in many examples (Putter.). and was found by Yoshida (1956) in food intake.The curveof -e00 30 lineargrowth is a decayingexponential withouta . growthrates are decreasingand growth eventuallyattains a steady state. We have and the gut. to weightitself.. organs. 4). pers. On the other power.ler (1943). in This characteristic course of growthis easily insects.withnot too muchchange rule but ratheris proportional understood. 1951a). Secondly. Berta_ 140 5 3 lanffy.c. 1934. the ratio between surface and hand. is proportional to the square of body length.j -- This content downloaded on Wed.20 60 80 100 200 300 500 800 30 40 turningpoint. experimentally.The largerit grows.

in land snails.Again we can calculate what growth curvesare theoretically to be expected. However. The same metabolicand growthtype also applies to hemimetabolousinsectslike the walking-stick. The curve of weight growthdoes not differ very much fromthat in 20 - - a)~~ 0 0-0 10 -j - -A 5--20 30 40 WeightIn mg. E8 = '- --V4 {> get a comthe growthequation. THE THIRD METABOLIC AND GROWTH TYPE the faster the largertheanimalbecomes. The morecatabolismincreases.43 -?- - > b) 2--f a) 0 8jII7 6 = 30 40 o 20 40 60 80 60 80 100 200 300 Weight in mg. After whathappenls.ofMetabolic snail (Planorbis shell)in the ramshorn sp. the more does anabolism. rate (a) and linear growth (b) (diameter steady state is reached.226 80 60 THE QUARTERLY REVIEW OF BIOLOGY Of course. and thereupon cm e 0 _ _ curveand a secondgrowth growth pletelydifferent 4 type.growth 4 12 24 8 0 16 20 rates will not decrease but always increase. where there is no apparent metamorphosis. a value 23 < m < 1 into If we insert the basic equation. at the same pace. it appears that the growth should followa thirdtype. whichalso belongto thistype. THE SECOND Timqe in hours la) (Drosophi (exponent(b) culrve rate(a) and growth Metabolic and Muller ial) in insectlarvae. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions .Again. we have described a thirdmetabolic type.exponential growth is intercepted by seasonalcycles. AfterBertalanify (1943).This seems to be a para.In contrast 0 run catabolism. It is metamorphosiswhich abruptly intercepts the exponentialincrease. Therefore. Finally. Bertalanffy butis exactly stateofaffairs.one where metabolic rate is intermediate betweensurfaceand weightproportionality.growthis stopped here by an altogether different mechanism. This content downloaded on Wed. 4. doxrical and Muller (1943).both being weight-proportional. and whichis exemplified by pond snails. 500 700 AND GROWnTYasPE METABOLIC FIG. and no Growthis not limitedbut exponential. 5.even causing a decrease in body weight as largeamountsof tissueare broken down in order to develop the imago.and Time in weeks it grows.). FIG.anabolismand to the first type. (Tenebrio) 60 80 100 200 300 5 E 1: .an insect larva does not grow to any indefinite size. but the hormonal mechanismsresponsiblefor development appear to be similar.

applies the surface modified form of the Dubois K which rulein the somewhat pare the values of the catabolic constant mammalian curves. BerSo in different 1957).with inflexionat ca./ that the first experiments could and the second part much flatter. part of the allometric withN15.Table 6 it may be said that.and founda value of 1.fish.such (1949) calculated protein turnoverfrom their detail. the overallallometric value found was 1.attaining. types of growth. no steadystate attained. discussions of the theory.however.obvious that the theoryrepre. This content downloaded on Wed./kg. of metabolism. body weight/day. theconstants of the first In a number by experiment. surface line has a slope of about 23. (Bertalanify It is. werecalculatedfromthe growth pattern followsthe characteristic roughly as directly determined growth values of proteinturnover growth typediscussed. there are complications. 1953). witha degreeofcorrespondence In detail. type. the determination (cf.thatis. line is steeper. hand.Correspondingly. In more Eleven years later. preGROWTH IN MAMMALS AND MAN cedingsexual maturation. is verified ourprediction animalclasses thereare different Harms. The breaktakesplace correspond suchprediction. Respiration intermediate a steady state. stated by Rubnerformammany surfacerule was first just outlined.theerror If the basal or restingmetabolismin the rat is considerable. that is. have been verified there and somewhat paradoxically. on the other are relativelyfew good data suitable for this is admittedlyoversimplified in thephysiological determinations typeof analysis. a comparative physiology. ofcases.Lineargrowth. Duspiva. as a first be considered of appear to belong to our firsttype.and weight-propor(b) Weight growthcurve: sigmoid.As we have found in and Pirozynski.withthe standard formula. mammals inflexion a steady state.otherinvestigations and that withfurther 1952. thefirst Again.1955. it in 1938 the authorcalculatedthe turnover the protein from the growth curve of man. 5). of is different. however. II.cf. and it was already mentionedthat the of 1957). talanffy.165 g.basal metabolismin man.QUANTITATIVE LAWS IN METABOLISM AND GROWTH 227 TABLE 6 Metabolictypesand growth types Metabolictype Growth type Examples I. 6) is measuredover the entirelife span. Linzbach. and. metabolic types and different A case in point is growthin mammals. similar breaks appear in quite a sentsa first approximation. Only clinicallyimportantcase.than would Only a sound theory kg. where the physiology namely.similar tionality to I(b). and may the viewpoint givesa surveyof examplesinvestigated. surfacerule of metabolismapplies.but growth or seasonal by metamorphosis intercepted cycles. Sprinson and Rittenberg regression thereis a breakin thisline. Zeuthen.From agreeing with theoreticalexpectation. 13 of final weight. mammals draftfora new chapterin in a firstand crude approximation. Bertalanffy verifiedempirically one further examplewill be given. Indeed. the growth. Respiration weight-proportional Linear and weightgrowthcurves exponen. Orthoptera. to a slope of 23.1955. (Fig. 1955. of growth From the theory consequences can be derived which have been mals. (b) Weight growthcurve: sigmoid. For example. curve: attainingwithin. The appears that as a crudeoverallapproximation ruleobtains.Insect larvae. Helicidae tial. of proteinturnover rate of (Fig.. however.a steady state. have permitted at a body weightof about 100 g. We can com. body weight/day. flexion surface.Planorbidae between (a) Linear growth III. model Unfortunately sincethe theoretical whichis quite striking. developmentrevisions and the introduction however. complicating factorswill be necessary(forrecent as its curveis S-shapedwith an inflexion.3g. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions . Respirationsurface-proportional (a) Linear growthcurve: attainingwithout Lamellibranchs. 1955.

to the or RELATivE GRowTHI typicalpatternof growth.that weight growth----. 1957). analysis are the hormonalbalance. simplyby takingdifE 400 YmuS Wt1n *th r g. .before A correspond-growth puberty.the curve is enormously protracted.This growth curveofman. 1000 _ the shapes of theirgrowthcurves are the same. 6). scales fortimeand body size. however. beginning is t fn 5 50 The first puberty. when pubertytakes place an at in othermammalsand early age. merical fitthatcan be obtained withthe theoretical whose growth formulas. however. 6.Animalsrun cominginto play of the sex hormonesentails a theirdevelopmental through periodspeedily. This is demonstrated by and pathological cases.as the factorforhis uniquenessin nature.howgrowthtype.If. is one of the basic factors in the evolution thereare others(Fig. the rat. is At the same time. body weight.Since our growth formulas to a apply large number of species.is very different. as it does at the number of physiologicalcharacteristics. body weight. however.in the values of the constants. pituitarydysfunction. 7.After Bertalanffy Racine and Piozynski 1953) (1953). Length growth calculated according to equafirsttype. However. since laboratory dietshave increased thegrowth in This is to modern place at the timeof sexual maturation.whichdoes not CP 100 much alter the picture. with the qualification. curveis different from all others. the excellent nuno morethan a first be considered approximation. GROWTH OF THE ALBINO RAT meansof our formulas So it wouldappear thatmammalsbelongto the Donaldson'sdata (d1). DISCONTINUITIES IN THlE ALBiNo RAT is heralded in thegrowth cyclesoflowermammals. The latter. in still apes. 800 _ and the same curve can be used to represent the 600 growthof various species. follows the general pattern.e. the involutionof the thymus.Althoughthis change FIG. c200 (Bertalanify. 4 40 In infancyand part. it an important that these breaks and shiftsare found. shows twogrowth cycles. 8 shows ferent the growthcurves of a fish and a mouse. of ap. criticalpoint of around 100 g. an 200 300 400 0 100 Time in days excellentfit of the empiricalgrowthcurves by can be obtained(Fig. the growthcurve of the rat was analyzed long before the physiological E __E studiesjust mentioned 1938. such as pubertaspraecox in givenin the original papers. \P This content downloaded on Wed. There is small wonder of man. and certainly human developmentwhich.the quantitative expression of the retardation of ofliverand thymus. The result was of the rat followsthe first foundthat the growth too R100 _______ change.If these growthcycles are taken into account. Onlyregression linesareshown in as it were._ -200 _ _ _ _ on rat growth is a discussionof recentliterature given in Bertalanffy. with a characteristic again at the ever. (1952. 7). I I I 11II 1II1II1 30 400~ 10 15 20 30 40 60 100 200 childhood.. Body weighti'n g. 8e0 ib. proximately curve.thereis one organism After Bertalanffy (1941b). a breakat approximately 100g.228 1500 _ THE QUARTERLY REVIEW OF BIOLOGY .is of courseconnected data and statistical individual thefigure..abnormal in thegrowth of theentire is found ingdiscontinuity withchangesin animal(see Fig. The singular growth curveofman is a same age and body size: in the allometric growth of the liver. similar to that of the rat.onlyin man does it lead to a singular discontinuities shape of the !FAII appearat a bodyweight 100g. curve of man is entered. according to Bolk tissuerespiration (1926). however. 1951a. _ _ _ _ _ __ _300 300 On the other hand. with the tion(7).The figure shows. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions . Donaldfrom thefirst to thesecondcycletaking son'sdataaretoday transition notconsidered tobe optimal. FIG. Fig.the growth it appears to be unique.and deep-reachingchange in the entire metabolic pattern. A new growthcycle is added.i. verQor2 n0 with The second part of the curve. 7). The calculation two growth cycles must be distinguished. also shows a 300 growth cycle in detailedanalysis. as it were.

Westphal. 10: 181-213. This content downloaded on Wed. Untersuchungen keit des Wachstums. soon they reach puberty and the adult stage.Nat.Bern.and is thusenabled to learnand to collect experience. 26 Dec 2012 21:20:04 PM All use subject to JSTOR Terms and Conditions . Biologie. Experientia. of of Canada. 1953." growth the "first following whilegenerally with curve connected show breaksin the growth herald The data and conceptspresented puberty. Biol. H. Lond. 1940. Untersuchungen die Gesetzlichkeit des Wachstums. 1941a.. and C. Science. Arch.or one intermediatebetween surface and weight The various theoriesregarding proportionality. Mammalian and human growth. F. Rec. VON. Untersuchungen und des Wachstums. U. 115: 515541. tiberdie Gesetzlichkeit .is givena longperiodof youth.0 c _ _ _ ~ 0 o and the theNational Cancer search Institute.The theory typesand growth and verteby examplestaken frominvertebrate brate classes.. Wachstumsgradienten metabolische Gradienten bei Planarien. Man* Mouse x 5 5 lo SUMMARY years 40 37 5 40 _ _ 0- o0 20 20 ~~ _____ lO weeks Abramis bramao 5 ~~0 0 years______ 7V FIG. Untersuchungen uber die Gesetzlichkeit des Wachstums. a new chapter of physiology. Probleme einer dynamischenMorphologie. Theoretische Wachstum. . Untersuchungen keit des Wachstums. betweenmetabolic explanationof the connection is illustrated types. uber die Gesetzlich. Org. are discussed.. bers.. VII. of metabolism the size dependence of the relation of with particularconsideration to to body size. 131: 613-653. P. 1. Aufl. permits advancedby theauthor. D. there are growth by different "growthtypes" distinguished A generaltheory curvesof the speciesconcerned. III. 85: 111-117. 1951a. Corresponding tissue respiration the "metabolic types" mentioned. three "metabolic types. 1934. physiology LIST OF LITERATURE E.QUANTITATIVE 100 LAWS IN METABOLISM AND GROWTH 229 _80 80~~~~~~~~~~ -______ ________x z- 0 < 3: (. 163: 156-158. Gen. Bertalanffy. 1953. Thus the characteristic humangrowth curve is a prerequisite for that mental development and civilizationwhich so sharply distinall otherbeings...von W.." i. LEBLOND. or to weight. Bd. CO1PARISON THE GROWTH CURVES O0F OF A FISH. . physiological constitutions 109: 579-585. Aufl. Francke. 1938. L. EntwickMech. V. 4: 255-269. 1948. ofanimalgrowth.and 1957. 61: 510-532. Stoffwechsel. 1942. uber BERTALANEBY. 1942. Nature.. type. 15: 1-22. EntwickMeck. Borntraeger. Man. -.. gen. Quantitative relationsin the of mammals. Arch. AND MAN After Bertalanffy (1951a). Anat. iiber die Gesetzlich.Partofthiswork was aided by research grants from the NationalRe- between connections Workaimedat establishing In thevarious is reviewed. Vieweg. 140: 81-89. Org. A quantitative theory of organicgrowth. and growth metabolism animal classes. Biol. Zbl. formsof dependenceof metabolicrate on body of size can be distinguished:proportionality metabolicrate to surfacearea. II. 1941b. Stoffwechseltypen und Wachstumstypen.A survey Council Research Humanities is givenin in general of animalgrowth the problem 1951a. F.Braunschweig.MOUSE. IL . Metabolic types and growth types. Amer. 15: 295-311. QuantitativeBeziehungen zwischen Darmoberflacheund KBrpergrossebei Planaria maculata. uber die Gesetzlichkeit . IV. I. Biol. Untersuchungen des Wachstums.the comparative ofgrowth. Allgemeine Grundlagen der Theorie: Mathematische und physiologische Gesetzlichkeiten des Wachstumsbei Wassertieren. 1949. Problems of organic growth. Biophysikdes Fliessgleichgewiclsts. on the otherhand. Hum.8. Das organische Wachstum und seine Gesetzmassigkeiten.e. Council. The BERTALANFFY. 2. Berlin. guishesman from ACKNOWLEDGMENT Time in days Thisessayis baseduponwork carried through in the author's laboratories at theUniversity ofViennaand theUniversity ofOttawa(Canada). .. 1949.. Biol. ADOLPH. 1951b. continuousrenewal of the two types of alveolar cells in the lung of the rat.

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