INTEGRAL

ANALYSIS AND THE OF LIFE

by

PHENOMENA

Prof. F. G. DONNAN
(The Sir William Ramsay Laboratories of Inorganic and Physical Chemistry, University College, London, W.C. i)

If we concede that the scientific explanation of phenomena is ultimately based on a logical or mathematical formulation of relationships amongst measurable or observable quantities, a view which we may hold without reducing our scientific explanations to a purely positivistic formalism, then,. as Bom~ has shown so clearly, the mathematical formulation must not include things or sets of things which are essentially ("theoretically") incapable of exact measurement: Although I have expressed this somewhat crudely, every physicist will understand that I refer to the well-known limitation which the existence of the PI~A5"CK quantum of action h imposes on the ability of macroscopic measuring apparatus to determine, without essential interference, simultaneous exact measures of a 11 the space-time micro-parameters of an atomic system. Expressed somewhat differently, we may, perhaps, say that the existence of PLANCK'S h prevents us from extrapolating all the simultaneous observed values of these micro-parameters to their "true" limiting values corresponding to z e r o interference between the measuring (observing) apparatus and the observed object. A complete set of such simultaneous "true limiting values" has therefore no existence in the (experimentally verifiable) reality of an atomic system. In several recent publications Bo~II~ has applied somewhat analogous reasoning to l i v i n g systems (living organisms). It is now not a question of limitation owing to the existence of h, but of limitation owing to the special f u n c t i o n a l i t y o f l i f e . In other words, if we should attempt to measure simultaneously all the parameters necessary for an exact physical specification of a living organism, we shoutd thereby necessarily k i l l the organism (BoI~R's Exclusion Principle). The external object would still be there, but its internal and external functional relationships would now be different. W e might, indeed, call this new idea of Bo~R a p r i n c i p l e of functional Indeterminism.
Acta Biotheoretica I I I

F.G.

DONNAN

In the measurement of an inanimate system, we can set up as m a n y i d e n t i c a 1 samples as we require for the carrying out and comPletion of all the possible measurements. This repetition of identical samples enables us to rule out the factor of time in our technique of measurement. But a living organism l i v e s o n i n t i m e . W e cannot revert it in time, nor can we "set up" (create) a series of identical samples. It may be this peculiar time-linked (historical) character of a living organism which is the source of the novel principle enunciated by Bo~t~. In a living organism the past seems to reach out and grasp the future. The description of the "evolution" does not seem to be reducible to a purely differential time analysis, but to involve essentially also an integral time relationship, i.e. an integral time analysis. I f I may use somewhat philosophical language, there appears to exist a marked degree of h o 1 i s m with respect to time ("chronoh.olism"). In order to express these concepts in simple mathematical form, I shalI choose an ideally simple case, namely a system whose i n t e r n a 1 state is completely defined by a single measurable parameter c, and which is in relation with an external environment specified by a number a, /?, ,/,. ..... , of measurable environmental parameters. I shall call the curve (c, t) the "evolution-line" of the system (t = time). Let us consider, in the first place, the group of systems characterised by the differential equation of evolution dc d--t = ~ (c, ~, ~, 7 . . . . ) . . . . . (I) For the sake of (possibly) greater clearness and without essential loss of generality, I shall consider a specialised type of equation ( I ) , namely, dc - - k f (c) . . . . . . . . . (z) dt where k is a function of a, fl, ~, ...... ( " A u s a r t u n g " of the function ?). dc This equation states that the value of ~ at any given time t is completely determined by t h e measured value of c (the internal state of the system) and the values of a, fl, ,/ ...... (the environmental parameters) at the same moment of time t. Observing a given c at a given t, equation (2) allows us to predict the future state of the system (future evolution), provided we know how a, 13, ~/ ...... vary in the future. W e do not require to know the p a s t history of the system, i.e. how c and the environmental parameters have varied in the past. It is sufficient to know the forms of the functions f ( c ) and k (a, fi, y ...... ), and to know that at the time t the internal parameter possesses, i.e. has arrived at, the value c. I f we set up a

INTEGRAL

ANALYSIS

AND

THE

PHENOMENA

OF

LIFE

series of samples of the type of systems considered, all observed to have: the same value of c at a given time t, then all the samples will behave in the same way in the future, i.e. they will all possess identical evolution-lines, provided, of course, that the environment varies in the same way in each case. So far as we can measure them, all the samples will be i d e n t i c a I in nature and behaviour. T h e y do not " r e m e m b e r " their past histories. Their evolution lines may have been different in the past (different initial values of the internal state-parameter and different time-histories of the environmental parameters ~ , p , 7 ...... ) , b u t w h e n t h e y i n t e r s e c t they coalesce in a single evolution-line, provided they are subjected to the same environmental conditions in the future. Their individualities vanish. I shall call these systems u n h i s t o r i c a l systems, and m y intention is to suggest that they possess the time-character of what we call non-living or inanimate systems. It must not be forgotten, however, that, for the sake of simplicity, I have chosen an ideally simple class of such systems (single state-parameter systems). I proceed now- to consider another class of systems, the internal state of each system being, as before, defined by a single measurable parameter c. And, as before, each system is in relation w i t h a set of external environmental parameters a, fi, y ...... But in this new class of systems the differential equation of evolution is no longer expressed by an ordinary differential equation. Instead, there must be employed an i n t e g r o - d i f f e r e n t i a 1 equation. F o r this purpose I shall use the equation.
t

dxc dt

/~lfl

(s -~ fk~f2 (c) dt . . . . . . .
]

(3)

where hi, and k2 are functions of a, /3, 7 . . . . . . . , and ,~ is a c e r t a i n definite and finite period o f p a s t t i m e . The functions fa, f2, kl, and k2, together with the quantity z, charaeterise the species of systems now considered. I wish to say at this point that I do not attach any particular importance to the special form of equation (3), except that it m u s t contain the integral term (or some similar integral term). In the form which I have given, it will be noticed that, provided f2 (c) is finite in the interval ( t - - z, t), equation (3) approaches equation ( I ) in the limit as k2 and (or) z become smaller and smaller. W e may express this by writing (3) ~ (2) when k2'and (or) z - , o. Equation (3) differs, therefore, from a functional point of view, only quantitatively from equation (2). The fundamental characteristic of (3) is that it contains the h i s t o -

4 t

F.G.

DONNAN

rical

term

~h2 f2 (c) dr, which for the sake of brevity may be

!

denoted by the symbol H. Hence the value of dr- at any given time t is n o t completely determined by the measured internal state of the system and the values of the parameters a,/~ ~,,. ..... at the same moment of time t. Observing now a given c at a given t we cannot predict the future state o f the system (future evolution) unless we know'the value of H, that is to say, unless we know the p a s t h i s t o r y of the system, i.e. how the environmental parameters, and therefore kl, k2, and c have varied with time in the historica] period z. W e might call this a principle of h i s t o r i c a 1 I n d e t e r m i n i s m . Different individuals of this species of systems, even if they were supposed to possess the same value of c at the time t, would not in general show identical behaviour should their evolution-lines arrive at the same (c, t) point. The various individual evolution-lines would in general diverge after intersection, e v e n if subject to the same environmental conditions in the future. The "individualities" of the systems would not vanish, as in the cases previously discussed. W e cannot therefore "set up" a series of evolving samples, all having the same observed value of c at a given time t and all showing the same behaviour i n the future under identical conditions (identical values of a, t?, 7, .,.), unless the special past histories of the systems have been such as to give them all the s a m e H-values at the specified (c, t)-point. I f z be regarded as a constant quantity, a system would have to evolve for a minimum period of time z before its behaviour could possess the f u 11 historical character corresponding to equation (3). W e could not, therefore, suddenly set up or "create" a system with this full historical character. Nevertheless, it would always be within .our power to t r a i n or c o n d i t i o n any individual by suitable variation of the environmental parameters during a n y given period z. O u r training or conditioning would, by changing the value o f H, alter the future behaviour (evo!ution-line) of the individual. In particular, by so altering the parameters a, fi, ~,, ... that k 2 always possessed a very small value, we might be able to d e g e n e r a t e the individual by reducing H to a very small value. The behaviour of the individual would then approximate to that of an inanimate unhistorical system. Conversely, we might be able to make H very large (highly conditioned "very alive" system). I propose to call the species of systems whose evolution is described by equation (3) h i s t o r i c a 1 s y s t e m s, and my intention is to suggest that they possess the character of what we call animate or living systems, at least insofar as the aspect of their time-evolution is

INTEGRAL

ANALYSIS

AND

THE

PHENOMENA

OF

LIFE

concerned. Thus the integral term is a measure of the c hr o n o h o 1 i s t i c character of an animate system. The greater the relative value of H, the greater will be the "animate" character of the system in its relation with time. It is particularly the famous Italian mathematician VOLTERRA who has developed the theory of integro-differential equations 1). Indeed, in this connection I think that the mathematicians have employed the adjective h e r e d i t a r y where I have used the adjective h i s t o r i c a l . As regards the mathematical form of my remarks, I lay no claim to any originality. My sole object has been to suggest that some form of integral or integrodifferential analysis will be necessary for a mathematical description of the phenomena of life. In the working out of this suggestion I have chosen ideally simple systems characterised by a single (internal) state-parameter, so that attention could be concentrated on the pure time-integration (chronoholism). There must also exist in any real systems another type of holism, namely s p a t i a l holism, requiring an integral or integraldifferential analysis extending through space, i.e. to a multipIicity of internal state-parameters conceived as co-existent in space at any given time. WEYL (I932) has pointed out that this type of holism is already existent in the modern quantum mechanics of atomic and molecular systems. I would suggest that the special organisation of a living system will require a further development of such spatial (and multiple) integraldifferential analysis. In the end, of course, both types of integral analysis must be fused in the mathematical description of a living organism as a special type of world-tube in space-time. The evolution-lines which I have mentioned in my discussion represent, in fact, ideally simple world-lines in an idealised two-dimensional state-time geometry (what one might, perhaps, call a two-dimensional thermodynamic state-time geometry). A living organism represents a system characterised by a very great number of functionally linked parameters, both internai and external. The nearest approach to any concrete type of integral analysis of such a system is that given by the complex of inter-linked nomograms, whereby L. J. HENDERSON (I928) has so wonderfully represented the bloodrespiration relationship-system of certain warm-blooded animals. Perhaps one might call HE?,TDERSON'S nomographic analysis an integral thermodynamic (mu!tiple state-parameter) analysis. I regard it as one of the finest achievements of modern quantitative biology, and the pioneer

r) Reference must also be made to the important work of BOREL and P~ca~D, although I have not had an opportunity of consulting their relevant publications.

F.G.

DONNAN

precursor, i n various respects, of the biological integral analysis of the future. It will be necessary now to consider some special matters. If we re-write equation (3) in the form dc

dt

--F(c,~,r

....

),

then the essential point is that the f u n c t i o n a 1 f o r m F changes with time. In fact, equation (3) shows how this gradual change of functionality may be related, by means of functional forms which do not change with time, to the special history of the system. W e may say that the historical e n g r a m H (to borrow an expression from the "engraphic" theory of S E ~ o ~ ) is an imprint on functionality (functional form). In the case of an inanimate system, there is no historical term /-/-, and therefore the functional form F is invariant with respect to time. Can, or must, we form any physical picture of this change of functionality with time? It would be natural to suppose that the term h r represents the change (or buildingup) of "something" in the system. It may be that this something is of the nature of organisation, and that the change of organisation is reflected in the change of the functional form F, and explicitly represented in equation (3) by means of the constant functional forms f~ and /"2 together with the integral term H. From this point of view, equation (3) would express the historical building-up (or change) of organisation. It would be natural to ask whether we can form any physical picture of this change of organisation with time. The essentially b i o 1 o g i c a 1 character of a living system might consist, however, in the impossibility of constructing any physical picture or model to "explain" this time-effect 1). A living organism may stand in some relation to what we call time that is not expressible by means of our present physical concepts. The probabilities of the micro-parameters of a definite physical system are generally regarded as time-less; in fact, they define the nature of the system. Can we define the nature of a living organism in this way? There is another matter to which I must refer. Life, as we know it, does not consist in the evolution of a single individual (system). The drama of life is played by a vast succession of separate individuals, giving us the picture of a special sort of discontinuity in continuity. Equation (3) would then have to be written I) In this connection reference might be made to EUGENIORIGNANO's theory of a nervous energy endowed with the energetic property of specific accumulation. Cf. Scientia, "The Phenomena of Life", p. 317, May 1929.

INTEGRAL

ANALYSIS

AND t

THE

PHENOMENA

OF

LIFE

7
(4)

d~ = k l f ( c ) +

t--'l"

k 2 f 2 (c) d t + H 2 . . . . . .

where ~- represents the period of time that has elapsed since the "birth" of the individual, i.e. t - - r is the date of birth. Hz is a second historical term representing the "racial history". Equation (4) would then apply to the life of a particular separate individual. In the language of PArLOr'S famous theory, the term f-/2 would represent the inherited "instinctive" reflexes, whilst the term H1, given
t

by the equation 2-[1 ~-=

]'k 2 f 2
t--'r

(c) dr, would represent the conditioned

reflexes acquired by the individual during the period t - - r ~ t of individual life. The "initial" evolution of the individual at birth would be "described By the equation dc d-t : k, f~ (c) + H 2 . . . . . . . (5) Returning to a consideration of equations (2) and (3), it will be noted that equation (2) is a differential equation of the first order. Examples of such equations are those governing chemical reacti6n-velocities or the radioactive decay of radioactive materials. The state-parameter c which I have considered is therefore a statistical (thermodynamic) m a c r o parameter. In such cases it is sufficient to know the value of c at a given time t in order to calculate its future value in a specified environment. In the dynamics of a particle, the motion is governed by a differential equation of the s e c o n d order. In this case the differential equation connects forces which are functions of the coordinates of the particle and accelerations which are second order differentials with respect to the time. dc T h e value of c (i.e. the value of the coordinate) and of d-t (i.el the value of the velocity) must both be known at a given time in order to make a dc prediction of the future possible9 In quantum mechanics either c or d-t may be measured with any required degree of accuracy, but both can be known at the same time with only limited accuracy. This restricts the possibility of prediction tO a certain extent (the well-known quantummechanical principle of indeterminism). The method of employing equation (3) in specified environmental conditions of the future may be explained in the following manner. From the knowledge of c during the time interval t - - z to t the behaviour of c in the infinitesimal time interval from t to t q- dt can be predicted by making

F.G.

DONNAN

use of the integro-differential equation (3)- From the values of c in the time interval t Z c d t - - z to t + d t t h e behaviour of c from t ~ - d t to t + 2 dt can be predicted in a similar way. Continuing this process, and afterwards going to the limit dt = o, the future can be predicted. By making measurements of c in the immediate neighbourhood of a given time point t~, we could determine with any required degree of dc accuracy the values of c and ~ at tl, and therefore the value of H at t 1. But this would not enable us to calculate therefrom the values of h r during a specified future interval t 1 -~ t2. SClZR6DIt,~CE~ (1935) has pointed out that, even in the older dynamics of particles, prediction of the future involves a certain amount of chronoholism. For, if at any time t we specify the particles by means of their positions and velocities, then the observational knowledge of these velocities will require measurements of position in the immediate neighbourhood of the time point t. W e might call this a differential chronoholism in comparison with the integral chronoholism expressed by equation (3). The previous discussion may be readily extended to simple types of multiple-parameter, systems. Let us consider a homogeneous single-phase system, meaning by this expression a system where any specified parameter possesses at any moment t the same value in every part of the system. If the state-parameters be denoted by c 1, c~ . . . . . G , we have, instead of equation (3), the system of n integro-differential equations dc__Ai =kifi dt (cl, c 2 , . . , c,) + [ ] ( i Fi (q, c2,. 9 9 c~) dt, (6)

where ~ denotes any one of the n integers I, 2 . . . . n, and ki and E~i are functions of the environmental parameters a, /3, y, ..., corresponding to each p a r a m e t e r ci. The evolution-line of the syskem is a curve fin an (1~ + I)-dimensional space, any point in this space being represented by the multiple quantity (Cl, c 2 , . . - c~, t). Thus the state of the system at time t is represented by the multiple quantity (q, c2. . . . c,~). Just as in the case of single-parameter systems, the individuality of the history of any particular system is determined by the values of the functions c~(t), B (t), Y (t) ... for each value of t in the period t - - Z - ~ t , and by the boundary values of the state parameters at t - - z . Denoting the multiple quantity (c~, ce, ca . . . . c~ ) by C, we may write equation (6) in the form
t

dci dt

/~j, (c) + [

F~ (C) dr.

(7)

INTEGRAL

ANALYSIS

AND

THE

PHENOMENA

OF

LIFE

In a similar manner we may consider a multiple-phase system in the case where each phase is homogeneous. The state of the system is now represented by the multiple quantity which specifies the value of every parameter in every phase. Denote this multiple quantity by C. The evolutionline of the system is a curve in a (2; n~ -~-I)-dimensional space, where
\ F

denotes the number of parameters defining the state of the p-th phase. The evolution of the system is determined by the system of X ~t~ integrodifferential equations
r P

dci'~----- ki,~ f~,~ (C) + ~ K~,~ Fi,~ (C) . . . . dt t--.X

where (i, p) refers to or species of systems is specify :(I) (2) (3) (4)

(8)

the i-th parameter in the p-th phase. Any group characterised by the four multiple quantities which All All All All the the the the functional functional functional functional forms ki, forms fi, forms Ki,~ forms Yi,

respectively, together with the value of z. Apart from boundary conditions, the individual behaviour of any specified individual of the group or species is determined in each case by the particular values of the environmental parameters ~, /3, 7, ... as functions of time. I cannot conclude without expressing my gratitude to Dr EDI:ARD TELLER for the very valuable discussions I had with him during April 1935, that is to say, during a preliminary study of the theme of this paper. Since he has not seen the final amplified draft for publication, he cannot be held responsible for any'errors I have committed. December, 1935. NOTE ADDED AT TIME OF CORRECTING PROOFS Since writing this paper, I have had an opportunity of reading the remarkable contributions made by Professor ADOLF MEYEI~ to the principle of holism. On page 5~ of his monograph entitled ,,Ideen und Ideale der biologischen Erkenntnis" (Leipzig, Barth, I934), he writes ,,Ich persGnlich glaube, dass die neue biologisch erforderliche Mathematik in Richtung eines Ausbaus der Gruppentheorie und der Integralgleichungen liegen wird. Differentialgleichungen bildeten das wichtigste mathematische Riistzeug der theoretischen Physiker, Integralgleichungen werden dasselbe fiir die theoretische Biologie besorgen." In this remarkable passage

IO

F.G.

DONNAN

P r o f e s s o r ADOLF MEYER f o r e s h a d o w s ( f o r the first time, so f a r as I k n o w ) the necessity o f e m p l o y i n g i n t e g r a l analysis in the m a t h e m a t i c a l f o r m u l a t i o n of the b e h a v i o u r of living o r g a n i s m s . I t h i n k also t h a t his p r o p h e c y as r e g a r d s g r o u p t h e o r y is, equally r e m a r k a b l e a n d valuable. T.o all w h o a r e i n t e r e s t e d in the d e v e l o p m e n t o f t h e o r e t i c a l b i o l o g y (and, I m i g h t also add, the d e v e l o p m e n t o f a c o m p r e h e n s i v e p h i l o s o p h y of all n a t u r a l science), the m o n o g r a p h w h i c h I have m e n t i o n e d , t o g e t h e r w i t h P r o f e s s o r ADOLF METER'S book, , , K r i s e n e p o c h e n u n d W e n d e p u n k t e des biologischen D e n k e n s " ( F i s c h e r , Jena, I935) , can be w a r m l y r e c o m m e n d e d . REFERENCES HENI)ERSOSr, LAWRENCE, .]-., 1928. "Biood. A Study in General Physiology". Yale University Press. SCHR6DI~GER, E., 1935. "Science and the Human Temperament". London, George Allen & Unwin, Ltd. WEYL, IffERMANN, 1932. "The Open World". Yale University Press. ZUSAIVIMENFASSUNG Die Integralanalyse und die Lebenserscheinungen

Der Beschreibung der zeitlichen Entwicklung lebender Systeme kann eine reine Differentialanalyse nicht geniigen. In solchen F~illen muss man sich an Stelle der gewghnlichen Differentialgleichungen der integraldifferentiellen, bezw. der Integralgleichungen bedienen. Zur leichteren Veranschaulichung der mathematisehen Darstellung betrachtet Ver:fasser zuerst diejenigen Systeme, deren innerer Zustand sieh durch ein einziges Parameter c bestimmen l~isst. Die zeitliche Entwicklung eines leblosen Systems dieser Klasse werde dutch die Differentialgleiehung ~ ) - dc~
kf(c) ...

(I) dargestellt, wo t = Zeit, und k eine Funktion der ~iusseren Parameter ~, p, y . . . . ist. Im Faile eines jeden Systems sind die ~iusseren Parameter a, p , ,/ . . . . charakteristische individuelle Funktionen der Zeit. Misst than c am Zeitpunkt t, so liisst sieh die Zukunft des Systems vermittelst der Gleichung (I) bereehnen, wenn die zukiinftigen Werte der ~iusseren Parameter gegeben sind. Solche Systeme nennt Verfasser n i c h t - h i s t o r i s e h e Systeme. lm Falle der lebenden Systeme werde die zeitliche Entwicklung durch die Integral-differentialgleichung
t

aD = k, A (c) +

ae

k2 A (c) at

. . . . . . . .

(2)

dargestellt, ~vo fl und f~ Funktionen des Zustandsparameters c, und kl und ks Funktionen der ~iusseren Parameter sin& ])as Zeitintegral erstreckt sich auf das vergangene Zeitinterval t - - z --~ t, wo z einen positiven endliehen konstanten Wert besitzt. Diese Systeme nennt Verfasser h i s t o r i s c h e S y s t e me. Setzt man H fiir das darin enthaltene Integralglied, so verwandelt sich die Gleichung (2) in: de
a-) = kl f i (c) + H . . . . . . . . . . (3)

INTEGRAL ANALYSIS AND THE PHENOMENA OF LIFE

II

Misst man nun c am Zeitpunkt t, so kann man in diesem Falt die Zukunft des Systems ohne Kenntnis des historischen Glieds /4 nicht berechnen. Das Vorkommen des historischen Integralglieds t4 nennt V e r f a s s e r C h r o n o h o 1 i s m u s. Man kann deshalb die historischen Systeme auch c h r o n o h o l i s t i s c h e Systeme nennen. Die biologische Interpretation dieser Betrachtungsweise wird yore V e r f a s s e r diskutiert, und die Gleidhung (2) auf Systeme mit erblichen Eigenschaften ausgedehnt. Es werden zuletzt Systeme mit vielen Zustandsparametern in gewissen einfachen F/illen kurz behandelt.

R1~SUMs
L'AnaIyse int6grale et les ph6nom6nes de la vie L'analyse diff6rentielle ne suffit pas, selon l'avis de de l'auteur, pour la description de l'6volution des syst~mes vivants au cours des temps. I1 faut utiliser dans ces cas les 6quations int6gro-diff6rentielles ou int6grales. P o u r faeiliter le traitement math6matique, on pent consid6rer d'abord les syst~mes dont l'6tat int6rieur est d6fini par un seul param~tre c. L'6volution d'un syst&me inanim6 de cette c l a s s e se laisse d6crire par l'6quation diff6rentielle dc
a-? ----- k / (c), . . . . . . . . . . . (~)

oh t = temps, et k est une fonction d6termin6e des param~tres ext6rieurs a, fl, ~', ... Si l'on connait c ~ un temps donn6 t, l'6tat du syst~me 5. un temps futur t' se laisse pr6voir au moyen de l'6quation (I), quand les valeurs des paramfitres ext6rieurs clans l'intervalle f ~ t' sont fix6es. L'auteur appelle les syst6mes de cette classe des syst~mes n o n - h i s t o r i q u e s . P o u r Ies syst&nes vivants, il ~aut se servir, selon l'avis de l'auteur, des 6quations int6gro-diff6rentielles. P o u r les syst~mes d6finis par un seul param~tre c, on peut se servir, par exemple, de l'6quation t
_dc __-- klfl dt (c) -+- f k ~ f 2 (c) dr, J . . . . . . . . (2)

off kl et k2 sont des fonctions d6termin6es des param6tres ext6rieurs. L'int6grale s'6tend sur l'intervalle t - - z---+ t ,,~ z e s t positif et constant. En indiquant le terme int6gral par le symbole H, on peut 6crire (2) dc dD -----k~ f l (c) + AT . . . . . . . . . . . (3) La connaissance de c ~ un temps donn6 t ne laisse pas pr6voir l'6tat du systfme un temps futur t' sans connaissance du terme historique /4, quand m6me les param~tres a, /~, 3', ..., seraient des fonctions d6termin6es du temps dans l'intervalle t---~/. L'autenr appelle l'existence du terme int6gral historique Jr-/ le c h r o n o h o 1 i s m e. On peut appeler les systhmes de cette classe des s y s t b m e s h i s t o r i q u e s o u chronoholistiques. L'interpr6tation biologique de cette fa~on de tralter les syst6mes vivants est discut6e par l'auteur, qui 6tend l'6quation (2) pour le faire repr6senter aussi les propridt6s h6r6ditaires des ~tres vivants. Quelques cas de systhmes dont l'6tat int6rieur est d4fini par plusieui-s param6tres et plusieurs phases sont trait6s brigvement.