On the Making of a System Theory of Life: Paul A Weiss and Ludwig von Bertalanffy's Conceptual...

Volume 82, No. 4 December 2007THE QUARTERLY REVIEW OF BIOLOGY

349

The Quarterly Review of Biology, December 2007, Vol. 82, No. 4

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ON THE MAKING OF A SYSTEM THEORY OF LIFE:PAUL A WEISS AND LUDWIG VON BERTALANFFY’S

CONCEPTUAL CONNECTION

Manfred Drack

Department of Theoretical Biology and Department of Neurobiologyand Cognition Research, University of Vienna

1090 Vienna, Austria

e-mail: [email protected]

Wilfried Apfalter

Department of Philosophy and Department of Neurobiologyand Cognition Research, University of Vienna

1090 Vienna, Austria


David Pouvreau

School of Advanced Studies in Social Sciences, Koyre Center75231 Paris, France


keywordsorganismic biology, theoretical biology, Biologische Versuchsanstalt,

systems biology

abstractIn this article, we review how two eminent Viennese system thinkers, Paul A Weiss and Ludwig

von Bertalanffy, began to develop their own perspectives toward a system theory of life in the 1920s.Their work is especially rooted in experimental biology as performed at the Biologische Versuchsanstalt,as well as in philosophy, and they converge in basic concepts. We underline the conceptual connectionsof their thinking, among them the organism as an organized system, hierarchical organization, andprimary activity. With their system thinking, both biologists shared a strong desire to overcome whatthey viewed as a “mechanistic” approach in biology. Their interpretations are relevant to the renaissanceof system thinking in biology—“systems biology.” Unless otherwise noted, all translations are our own.

350 Volume 82THE QUARTERLY REVIEW OF BIOLOGY

Introduction

IN TERMS OF publications, “systems biol-ogy” is currently an exploding field (see,

e.g., Chong and Ray 2002; Grant 2003). Note,however, that system thinking is not at all newin biology. In the sciences of life, it can betraced back at least to the German “teleo-me-chanicist” tradition inaugurated by Imman-uel Kant’s Critics of Judgement, of which Johan-nes Muller and Karl von Baer were the mainproponents (Kant 1789; Lenoir 1982). Moregenerally, integrative (or “holistic”) thinkingis not a “discovery” of the 20th century. It hasa rather long history going back to ancientGreek thinkers: Aristotle already claimed that“the whole is of necessity prior to the part”(Aristotle 1966:1253a20). Between the 14thand 16th centuries, it was a major intellectualtrend, which strongly reemerged at the turnof the 19th century in German Romanticismand post-Kantian idealism. Among others,Nicholas of Cusa, Gottfried Wilhelm Leibniz,and Johann Wolfgang von Goethe are consid-ered “precursors” of modern systems think-ing (Bapp 1921; Bertalanffy 1926a; Harring-ton 1996). One of the leading figures, whorecurrently referred to those thinkers, is thephilosopher and theoretical biologist Ludwigvon Bertalanffy (1901–1972), mostly knownfor his project on a “general system theory”(GST). Unsurprisingly, he is still regularlyquoted—although mostly laconically—by ad-vocates of “systems biology” in the early 21stcentury (e.g., Chong and Ray 2002).

Pursuing the historical roots of biologicalsystems theory inevitably leads to early 20thcentury Vienna. Ludwig von Bertalanffy livedthere and was influenced by biologists HansLeo Przibram (1874–1944), the founder of alaboratory for experimental biology, and hisstudent Paul Alfred Weiss (1898–1989), bothof whom were also based in Vienna. We willmore closely examine the convergences ofthe experimental and theoretical works ofWeiss—who allegedly claimed that “he was re-sponsible for Bertalanffy’s initial appreciationof the systems nature of development andbehavior” (Haraway 1976:38)—and Berta-lanffy’s “system theory” of life; we extract ba-sic concepts that were already developed longbefore the recent emphasis on “systems biol-

ogy.” The early writings of Weiss and Berta-lanffy can be fruitful for modern research:they already developed many concepts thatcan shed light on certain contemporary de-bates and perhaps only need to be adapted tocurrent intentions.

The purpose of this article is therefore two-fold. We first provide a historical overview ofthe works of these system thinkers, and thenreflect the early system conceptions on recentdevelopments in biology.

Context of Early “Viennese”System Theory

The fin de siecle and the following yearswere characterized by innovations in manyfields, among them science, arts, and poli-tics. In particular, the developments in phys-ics and psychology, together with problemsin biology, provided an impetus that led to asophisticated system conception of organ-isms.

An important revolution took place inphysics, which many scholars from other dis-ciplines liked to cite. A statement by MaxPlanck, inspired by the new quantum me-chanics and referring to the false dictum thatall physical processes can be displayed as astringing together [Aneinanderreihung] of lo-cal processes, summarizes the relevance forsystem conceptions: “The physical world isnot simply a sum of spatial and temporal sin-gle worlds running one besides the others,and many phenomena escape [entziehen sich]the understanding when one does not con-sider a physical object [Gebilde] as a whole”(Planck 1929:17).

At about the same time, a conceptual shiftoccurred in another field, namely psychology,which was also extended to the realm of phys-ics. Based on the conceptions of the Austrianphilosopher Christian von Ehrenfels, Gestalttheory was developed (Ash 1995); it empha-sized the claim that an entity’s total charac-teristics cannot be discovered by simply sum-ming up its partial characteristics. WolfgangKohler, one of this theory’s leading figures,derived concepts based on physical systemsas known from thermodynamics and elec-trostatics. He considered Ehrenfels’s two cri-teria (“oversummativity,” i.e., the whole is

December 2007 351WEISS AND BERTALANFFY’S CONCEPTUAL CONNECTION

more than the sum of the parts; and “trans-position,” e.g., a melody can be played in adifferent pitch, but stays the same) as con-stitutive for Gestalt (Kohler 1920). Similarly,Bertalanffy (1928:69–70) characterized aGestalt as comprising properties that cannotbe found by simple addition of the compo-nents’ properties and that furthermore dis-appear when the Gestalt is destroyed; he usedthe term “Gestalt” synonymously to “systemicstate” [Systemzustand] (Bertalanffy 1929c:89).

Bertalanffy witnessed a controversy amongbiologists at the turn of the century between“mechanicism” and “vitalism” (for contempo-rary analysis of this controversy see, in partic-ular, Haldane 1884; Loeb 1906; Schaxel 1919;Needham 1928; Woodger 1929; Bertalanffy1932; M Hartmann 1925; and M Hartmann1937; for a recent discussion see Pouvreau2005b). “Vitalism” can have two meanings, ametaphysical or a methodological one. Meta-physically, it asserts that biological phenomenacannot be explained without the action of anonspatial principle harmonizing the matterand energies involved in living phenomena.Methodologically, it is not antinaturalistic andonly asserts that, at least provisionally, biologyshould have its own categories, methods, andlaws. As for the term “mechanicism,” it wassometimes loosely referred at that time tothe opposed view that should be termed“physicalism” in order to avoid confusion;that is, the idea that the sole concepts, meth-ods, and laws of physics and chemistry en-able biological phenomena to be grasped.But this was not the only meaning. More gen-erally and precisely, “mechanicism” seems(and seemed) to refer to a complex group ofmore or less coherently related positions. Themore fundamental position is the “analytico-summative” approach to biological phenom-ena: its basis is the (methodological or meta-physical) postulate that any entity can beanalyzed in parts whose properties can bestudied in isolation from others without in-convenience (the relationships among theparts being “external,” not “constitutive”).Through decomposition into “independent”causal chains and their “linear” composition,the properties of the whole are then sup-posed to be derivable from the knowledgethus acquired. “Mechanicism” would then be

represented in a biology that combines the“analytico-summative” approach with one orseveral of the following positions: physical-ism, determinism (each state is univocally de-rivable from previous states), and “reactivism”(the changes in the behavior of an entity areascribable to the sole action of its environ-ment). An extreme example is Jacques Loeb’stheory of tropism, which combines all ofthese items.

A paradigmatic variant of mechanicism is a“machine-like” model applied to the organ-ism. This model can be seen as a successor ofDescartes’s “bete machine,” which explainslife merely by means of physics and chemistry.Bertalanffy (1932:55) saw this Cartesian meta-phor as one of the most unfortunate concep-tions in the history of science and philosophy.Such a machine theory was opposed by manyscientists. Some biologists in the late 19thcentury were considering a type of living mol-ecule—e.g., Darwin’s “gemmulae,” Pfluger’s“living proteins,” or Haeckel’s “Plastidulen”(Penzlin 2000:434)—although most physiol-ogists rejected vitalism. Two findings were im-portant for rejecting vitalism (see Lenoir1982; Penzlin 2000): the first law of thermo-dynamics and successful physical-chemicalanalyses of vital functions. This was coupledwith the Darwinian selectionist scheme ofevolution, which is “analytico-summative” (se-lection and summation of single modifica-tions) and “reactivist” (the phylogenetic ad-aptation of the organism is a mere reactionto the environment). In contrast, after exper-iments with early development stages of seaurchins, Driesch formulated a “neo-vitalistic”concept: he viewed the embryo as an “har-monic-equipotential” whole, i.e., a whole ofcells in the embryo to which it does not mat-ter whether some cells are removed or dis-placed, in which each event is finalized by a“wholeness causality” [Ganzheitskausalitat]manifesting the operation of an immaterialagent, the “entelechy” (a term used byDriesch, as well as by Aristotle and Leibnizbefore him). A crucial experimental findingfor his argument was when, in the two-cellstage of development, he separated the cellsand found that each would eventually form awhole larva (Driesch 1899; Penzlin 2000).


institutions of experimental biologyAnother debate in biology at that time was

on methodology. Since the 1850s, a lively dis-cussion arose between proponents of two sci-entific approaches in biology: should mereobservation or experimentation be employed(Nyhart 1995; Querner 2000)? Arguing forobservation, traditional morphologists suchas Ernst Haeckel saw no sense in artificial, vi-olent, experimental studies of developmentbecause they considered it to be methodolog-ically sufficient to thoroughly draw conclu-sions from observing nature’s own diversityand genuine transformations. In their view,an experiment alters the events that normallyoccur during developmental processes and,thus, provides no causal explanation. Arguingfor experimentation, scientists such as Her-mann von Helmholtz, Sigmund Exner, andWilhelm Roux favored well-constructed ana-lytical experiments from which they hoped toderive causal explanations.

Apart from older institutions such as theStazione Zoologica in Naples (founded in1872) or the Marine Biological Laboratory inWoods Hole, Massachusetts (founded in1888), a novel, avant-garde Viennese fin desiecle institution became famous for its ex-perimental research on developmental andevolutionary processes. This Biologische Ver-suchsanstalt (Institute for Experimental Bi-ology), or popularly “Prater Vivarium” (asthere were other vivaria in Vienna, too), wassituated in a Neo-Renaissance building at theso-called Prater, Vienna’s popular recreationand entertainment area (including parks, for-est land, and fields) at the beginning of thePrater Hauptallee (Prater main avenue; Fig-ure 1). Equipped with automatically regu-lated installations for controlling humidityand temperature, the Biologische Versuch-sanstalt was an innovative research facilityboth methodologically and technologically. Itwas privately established as a new type of bio-logical research institution in 1902, andopened in 1903, devoted exclusively to exper-imental research. It was founded by Przibram,along with his botanist friends Leopold vonPortheim and Wilhelm Figdor (Reiter 1999;Hofer 2002; Gliboff 2005).

The “Prater Vivarium” was designed “to

tackle all big questions of biology” (Przibram1908–1909:234) by transdisciplinarily apply-ing similar experiments to both plants andanimals (e.g., with transplantation, breeding,processes of selection, orientation, and polar-ity in development). To perform those bio-logical experiments, methods from physicsand chemistry as well as physiology were ap-plied to living organisms. Compared to phys-iological research, the “Vivarium” biologistsneeded long-term experiments to deal withquestions of embryogenesis, regeneration,and evolution. This called for methods thatallowed investigations over a series of gener-ations.

There were several departments at the Biol-ogische Versuchsanstalt: one each for zool-ogy, botany, and physiology of plants. Forsome time, there were also physiology andphysics/chemistry departments. WolfgangPauli Sr., a colloid chemist and close friendof Ernst Mach, headed the latter. AlthoughPrzibram, Paul Kammerer, and Figdor had al-ready received their Habilitation (grand uni-versity teaching license) at the University ofVienna, the “Prater Vivarium” researchers—in many cases, liberal or left-oriented, open-minded individuals with a Jewish background(Reiter 1999; Coen 2006)—were subject to ef-ficacious disadvantages orchestrated by estab-lished, and often anti-Semitic, academicsfrom the university. However, effective Janu-ary 1, 1914, the successively enlarged andhighly productive Biologische Versuchsan-stalt formally became a part of the KaiserlicheAkademie der Wissenschaften (ImperialAcademy of Sciences) in Vienna. At the sametime, the three founders were appointed asunsalaried heads of their departments, andPaul Kammerer was appointed Adjunkt (civilservant), which, at that time, was the only po-sition with a salary.

There were also strong international activ-ities and lively networking. About one-third ofthe positions were reserved for scholars fromabroad. Those guests included people such asKarl von Frisch (Germany), Erich von Holst(Germany), A E Hopkins (U.S.), and JosephH Woodger (UK); Woodger translated Ber-talanffy’s Kritische Theorie der Formbildung (seeBertalanffy 1928; see also Pouvreau 2005b fordiscussion about their relationship). From


Figure 1. The Biologische VersuchsanstaltThe “Prater Vivarium” is depicted in the left foreground as seen from the Riesenrad, Vienna’s giant ferris

wheel. � Osterreichische Nationalbibliothek/Wien L 39.274-C.

1920 to 1934, 39 of the 109 scientists (practi-cum students or research personnel) of theBiologische Versuchsanstalt were women(Rentetzi 2004:368). One example is AlmaMahler, who worked as an assistant in PaulKammerer’s laboratory (Everdell 1997:325).Due to a series of public, high-level lectures—“big show-lectures” (Kammerer to Hugo Iltison July 6, 1910, cited in Gliboff 2006:530)—the “Prater Vivarium” was quite popular in Vi-enna (Hofer 2002).

the visionary of the biologischeversuchsanstalt

One of the founders of the “Prater Vivar-ium,” Hans Leo Przibram, studied zoology inVienna under the guidance of Berthold Hat-schek, a well-known comparative morpholo-gist with a special focus on vertebrate bauplane(body plans) and himself a student ofHaeckel and Rudolf Leuckart. After studies inLeuckart’s laboratory (with its highly attrac-tive microscopic equipment) in Leipzig, and

after graduating from the University of Vi-enna in 1899, Przibram went to Strasbourgin order to study at the laboratory of physio-logical chemist Franz Hofmeister (Reiter1999:591), which was very avant-garde at thattime. In 1904, he received a Habilitation “forzoology with special consideration of exper-imental morphology” at the University of Vi-enna. He started his multivolume work Ex-perimental-Zoologie (Przibram 1907) with thefirst volume on embryogenesis, researchingissues that were later also addressed by Ber-talanffy (e.g., 1937:91–99; 1942:289–291),such as growth rates and allometric growth.Although Hatschek followed the scientificstyle of Haeckel, Przibram was influenced byLoeb (Reiter 1999) and strove to put intopractice Roux’s program—integrating analyt-ical approaches from mechanical anatomyand evolutionary morphology (Nyhart 1995).The goal was to find the specific “forces” thatact upon the development of organisms.Loeb and Roux (also a student of Haeckel)


Figure 2. Hans Leo Przibram (1874–1944)� Osterreichische Nationalbibliothek/Wien NB

510.429-B.

both tried to find a physical, chemical, andphysiological basis for ontogenesis. In partic-ular, Roux conceived Darwin’s principle of se-lection as also implying internal selection.When mentioning a “struggle of parts withinthe organism” (Roux 1881), he especiallyhighlighted the “struggle” for nutrients andspace. Bertalanffy, although in deep disagree-ment with the Darwinian selectionist scheme,later (the first time in Bertalanffy 1940b:56)refers to this “struggle of the parts,” in whichhe sees an illustration of the metaphysical(Heraclitean and Cusean) principle of har-mony through contradiction, when develop-ing his theory of allometric growth. Arguingfrom Roux’s point of view and consideringliving beings as highly active entities inti-mately related to their environments, the bi-ologists of the Biologische Versuchsanstaltsystematically questioned external selectionas the crucial drive directing evolution(Gliboff 2005). This approach vaulted the“Prater Vivarium” staff directly into highlycontroversial central issues of developmen-tal and evolutionary biology. Contemporaryevolutionary developmental biology (EDB),or “evo-devo,” examines this role of devel-opment in evolution and in the evolution ofdevelopmental mechanisms (Hall et al. 2003;Muller and Newman 2003). Wagner and Laub-ichler (2004) recognize the Vivarium programas a precursor of modern “eco-evo-devo.”Describing the laboratory style at the Biologis-che Versuchsanstalt, Przibram programmati-cally declared: “No specialization, [but rather]a generalization of obtained experience is ourgoal” (Przibram 1908–1909:234). With exper-imental research, not only on one specificgroup of organisms, but rather on variousplants and animals, he aimed at finding gen-eral answers to the “big” questions in biology.

Przibram proposed to ascribe “the Gestalt,developing under the influence of the innerforces, to a state of equilibrium [Gleichge-wichtszustand],” so that “by viewing organicform as a state of equilibrium . . . a compen-sation of disturbance [Storungsausgleich] willbecome accessible to mathematical treat-ment” (Przibram 1923:22–23). In this con-text, note the influence of the German phys-iologist and psychologist Karl Ewald Heringon Przibram. Przibram quotes Hering, who in

1888 wrote: “An organism is a system, whichis able to sustain its constitution [Beschaffen-heit] (chemical thermal state) against influ-ences from outside, and which shows a dy-namic state of equilibrium of considerablestability” (Przibram 1906:222). The synony-mous usage of the terms “Gestalt” and “form”is obvious in Przibram. He refers to the influ-ence of the whole on the parts and how thiscan be dealt with mathematically. This no-tion—as discussed below—became importantin Weiss’s concepts, although he was less in-volved with mathematical details. Przibramcommittedly discusses the possibility of apply-ing mathematics to biology in general and tomorphogenesis in particular. He also consid-ers laws of higher order in cases where thedetailed underlying single processes are notknown. “Anyway, the influence of the ‘whole’on the ‘part’ turns out to be accessible tomathematical processing, grounded on ourideas from exact natural history [Naturlehre]”(Przibram 1923:23). Thus, he proposes a“mathematical morphology” by which the pa-rameters of the developing forms (e.g., pro-


portions) should be described, also in relationto physical measures such as surface tension.The investigation should reveal “constants”that explain similar forms in various species(Przibram 1923:23–25). Przibram also dis-cusses Frederic Houssay’s project of a “mor-phologie dynamique” (Przibram 1923:9–11);the fundamental idea here is to unite mor-phology and physiology, an improvementcompared to D’Arcy W Thompson’s approachof mathematical morphology. Thompson’stheory, or rather inductive method, which haslittle predictive value, is likewise criticizedby Przibram (1923:13–14) and Bertalanffy(1937). Przibram’s outlook toward a “math-ematical morphology” is very similar to thenotion on which Bertalanffy (1932:105) laterbuilds his concept of “higher order statistics”(i.e., describing the average behavior ofhigher units of cells or organisms, regardlessof physicochemical single events). This issimilarly already expressed by Przibram. Hisexample is the first formulation regardingthe course of organic global growth (preciselythe field where Bertalanffy later brings hismain contribution to mathematical biology,see Pouvreau 2005b; Pouvreau and Drack2007): “It is, however, not always necessary toknow the contribution or the nature of theinvolved components of morphogenesis[Formbildung] to arrive at mathematically for-mulated and theoretically usable laws” (Przi-bram 1923:25). With regard to the statisticalview on nature since Ludwig Boltzmann, Przi-bram is influenced by his contemporary,physicist Franz Exner (Coen 2006). Przibramenvisions and already initiates “a mathemati-cal theory of the organic on the basis of quan-titative inquiry, of tying up with geometry,physics, chemistry, and of sharp definitions ofbiological terms.” He expects this “mathe-matical biology” to serve “as a safeguard ofhumankind against the forces of nature andnamely as a bulwark against prejudice, super-stition and untenable speculation” (Przibram1923:64).

Furthermore, Przibram also opposes theidea of a mere passive organism. When talk-ing about “‘self-construction’ [Selbstgestalt-ung] of organisms” (Przibram 1923:21), heemphasizes the pivotal agency (i.e., inherentactivity) of living beings. In this view, living

organisms are not mere playthings of envi-ronmental forces.

Paul Weiss’s System ConceptionsIn 1918, Weiss took up mechanical engi-

neering, physics, and mathematics at Vi-enna’s Technische Hochschule, now calledVienna University of Technology. After oneyear of training in engineering, he decided tostudy biology at the University of Vienna (forWeiss’s background and professional career,see Brauckmann 2003). By 1920, he had al-ready managed to receive one of the highlycoveted permanent positions at the “Vivar-ium” laboratories, where he later conductedwork for his doctoral thesis (Weiss 1922;Hofer 2000:149). After Paul Kammerer re-tired and left Vienna, Weiss, as his immediatesuccessor, was appointed Adjunkt in early1924. But he left the “Vivarium” in 1927 whenhe applied for dispense of his working duties(letter of March 25, 1928 and protocol no. 44of January 31, 1929, Vivarium papers, K2) inVienna in order to conduct research in Mon-aco, Paris, and Berlin. He definitively resignedfrom his salaried Adjunkt position in 1929. In1931, he was appointed Sterling Fellow at YaleUniversity. He went to the University of Chi-cago in 1933 and became an American citizenin 1939 (Haraway 1976; Brauckmann 2003).

The “Prater Vivarium” became especiallyimportant for Weiss and his later systemthinking; so much so that at the Alpbach sym-posium in 1968, Bertalanffy, when discussingWeiss’s talk, stated that, to his knowledge,Weiss was the first to introduce the term “sys-tem” in biology in 1924 (Koestler and Smy-thies 1969). As a matter of fact, Bertalanffy(e.g., 1932:87) knew very well that the philos-opher Nicolai Hartmann (1912), with whomhe was very much in line, had already held“system” as the fundamental category of bi-ology as well as of other fields (Pouvreau2005a). The concept of the “harmonic equi-potential” system as applied to the biologicalorganism was already the core of Driesch’sthinking since the end of the 19th century(Driesch 1899). But even Driesch was not thefirst one to refer to the organism as a system;as mentioned above, Hering did so beforehim.


Figure 3. Paul Alfred Weiss (1898–1989)� Courtesy of the Rockefeller University Archives.

weiss’s early butterfly experimentsWeiss received his doctoral degree from the

University of Vienna, supervised by Przibramand Hatschek, for an experimental study onthe normal positions of butterflies in re-sponse to light and gravity, which greatly in-fluenced his system conceptions. His experi-ments were designed to determine if and howmuch the normal position of butterflies is re-lated to the direction and intensity of lightand gravity. He therefore used one or twolight sources and changed the angle of a wallon which the animals rested from vertical tooverhanging horizontal. This experimentalapproach to the influence of external factorson organisms typifies the research approachat the “Prater Vivarium.” On the one hand,this yields knowledge about external factors;by knowing the influence of these factors andabstracting from them, it also yields insightinto the core of events—which he terms “spe-cific vital” (Weiss 1922:1, 8–9)—that are notinfluenced by outside conditions.

In the experiments, he first investigated theinfluence of the single factors by excludingthe others. In order to isolate the influenceof light, he allowed the animals to attach onlyto the ceiling. The direction of gravity wasnormal to the light vectors and, therefore,played no role in the orientation of the but-terflies. The influence of gravity was varied bychanging the plane angle of the substratefrom vertical to overhanging horizontal, thusaltering the relevant component of gravity.

A result of Weiss’s study was that the posi-tions of the butterflies could not be preciselypredicted by knowing the vectors of light andgravity alone. Nevertheless, he demonstratedthat the vectors do play a role. When only onevector was applied, the animals responded toit; when both were applied, the positions ofthe animals were influenced by a combina-tion of gravity and light. When altering theangle of the plane with constant light, therewas a greater influence of gravity in the ver-tical position, while light became more im-portant if the component of the gravitationalforce decreases. This allowed a rough esti-mate of the resting position:

The experiments totally confirmed theo-retical conclusions. It could be demon-strated that knowing the efficacy of sin-gle factors within a complex allowedunequivocal conclusions to be drawnabout the efficacy of the entirety fromwhich we previously isolated single parts;all positions [of the butterflies] turnedout to be demonstrable as coming off theentirety of those partial effects, an exclu-sive efficacy of one single factor wasnever observed (Weiss 1922:19).

The experiment with two light sources andnegligible gravity showed that the butterfly’sposition cannot be predicted based solely onlight direction and intensity. The organism’s“history” apparently plays a role. By takinginto account the individual’s preexperiencesbefore taking the normal position, Weiss con-cluded that some kind of memory must playan important role. Nonetheless, the devia-tions from the calculated positions seemed tofollow definite laws, and the butterflies ori-ented themselves toward the light source towhich they were closer when crawling to their


final position (Weiss 1922:8; 1925:223). Whendiscussing the symmetrical position in rela-tion to two light sources, Weiss claimed that,after disturbance, an equilibrium state isagain found by the animals. “It is always a re-action of the affected system from within itself,which after each disturbance causes the re-turn to a state of unique definiteness” (Weiss1922:20, our emphasis).

new approachesin developmental biology

With his early work in developmental biol-ogy, Weiss focused on regeneration and trans-plantation in vertebrates. Those studiesproved certain prevailing concepts and dis-cussions to be inaccurate (e.g., the debate onpreformism versus epigenesis). Weiss alsotried to overcome the debate on mosaic ver-sus regulative development, which was also in-fluenced by Hans Spemann’s “organizer ef-fect.” He argued that one cannot distinguishbetween them based on a strict timeline (suchas the eight cell stage); rather, every devel-oping organism at some time includes cellsthat are determined and cannot develop inother ways. Weiss’s studies in developmentalbiology also use the system concept. Heclearly states that ontogenesis cannot be un-derstood by examining single isolated aspectsonly. Again, he refers to Gestalt theory anduses it to describe the occurrences found inthe organism (Weiss 1926).

The developmental biologist Weiss notesthat, in the organism, spatial heterogeneity isaccompanied by substantial heterogeneity;thus, spatial form must not be viewed as sepa-rated from the variations in the involved ma-terials or substances. With the morphologicalstructure, certain functions are determinedand everything that appears as distinct differ-entiation was, in a former phase, a latent abil-ity. Biological form is defined as “the typicallocalization, i.e., arrangement and allocationof different sub-processes within the respec-tive material systems.” Accordingly, alreadythe fact “that at this location these and onlythese and at that location those and onlythose processes are introduced is for us‘form’; not only that those different processesactually create a certain spatial Gestalt” (Weiss1926:5). Developmental biology again har-

bors the concept of inseparable systems withan argument derived from the dynamic pro-cesses involved:

If in the developmental process [Gestalt-ungsprozess] each part would simply beconnected to its neighboring parts with-out taking a specific subordinationwithin the whole system, then the small-est deviation of a partial process at thebeginning would at the end lead to mas-sive discrepancies. This contradicts theobservation (Weiss 1926:7).

Weiss’s criticism of misleading terms re-flects his opposition to the concepts on whichthey are based. Terms like “equipotent” or“omnipotent” seem improper to him becausethe “material parts” of the early germ have noability or potential to build up the whole inan organizational perspective. Rather, theparts should be called “nullipotent” becausethe specificity of their development is deter-mined by their position, and differentiationis dominated by the factors prevailing at a cer-tain position (Weiss 1926:10–11, 13). In thiscontext, he derived a field theory for devel-opmental biology comprising five aspects(Weiss 1926; Weiss 1928:1570; listed as “fieldlaws” in Bertalanffy 1932:328):

1. Every field has axial structure, whichleads to hetero-polarity [Heteropolaritat]in at least one axis.

2. When material is separated from a field-bearing system, the field is contained inthe remainder in its typical structure.

3. When unorganized but organizable ma-terial meets a field area, then it is in-cluded.

4. The field components are different ac-cording to their directions. When twofields are brought together they can addup or result in a mixed field dependingon their orientation.

5. A field has the tendency to take up andincorporate equivalent fields from itssurroundings.

He programmatically declared: “All fieldlaws can be conceived as special cases of gen-eral system laws” (Weiss 1926:26). A field as-cribes to every point in space a certain quality,direction, and intensity, which is very similar


to the use of this term in physics. A wholeresponds to influences from the outside as awhole by inner reactions of the componentsin such a way that the former state is re-gained. The special cases in developing or-ganisms are “morphogenetic organizationfields.” The field concept is considered to becompletely in line with Gestalt psychology(Weiss 1926:23). During the developmentalprocesses in an organism, various subfieldscan come into existence that govern certainareas of the material parts within the whole,subsequently leading to an “autonomisation”of those entities (Weiss 1926:31).

Note that Weiss was not the first to talkabout fields in developmental biology. Boveri(1910) already postulated a concept ofmorphogenetic fields (see Gilbert et al.1996:359), and in the same year, Gurwitschstarted to use the field concept, which even-tually led him to refer to an embryonic field[embryonales Feld] (Gurwitsch 1922). Spe-mann, who discovered the effect of embry-onic induction and introduced the organizerconcept, also used the term “field.” However,Weiss is considered to be the originator of“field theory” in developmental biology (Har-away 1976:177)—he provided a theoreticalbasis for the concept (Gilbert et al. 1996:359).

far-reaching conclusionsWeiss’s results from his butterfly experi-

ments and further theoretical conclusionsfrom his thesis were first published in Ger-man. The article was already written in 1922(Weiss 1925:248), that is, before he came intocontact with Bertalanffy—they very likely meteach other in 1924 for the first time. At anyrate, Bertalanffy regarded this work as an im-portant contribution. This is reflected in thefact that an English translation was publishedin the General Systems Yearbook, the book seriescoedited by Bertalanffy (Weiss 1959). Weissexplained:

That 80-page article stated the basicpremises and principles of a holistic sys-tems-theory which I could derive co-gently from my own studies of animal be-havior and from cognate trends such as“Gestalt” Psychology so as to define in de-tail the scientific characteristics by which

a singled-out fraction of nature can right-fully be accorded the designation of “sys-tem” (Weiss 1977:18).

Weiss concludes that his results “proved to-tally incompatible with the mechanistic doc-trines—or rather, rationalizations—of animalbehavior prevailing at that time” (Weiss1977:17). With such doctrines, he refers es-pecially to Loeb’s tropism theory, by whichthe behavior of an animal is reduced to amere physicochemical reaction triggered byfactors exclusively coming from outside theorganism, and where the mechanisms thatgovern orientation toward light and gravityare the same in plants and animals. Weiss(1925:171, 177–178) rejects this “fatal mis-take,” which consists of identifying similarphenomena [Erscheinungsformen] as the resultof similar mechanisms. According to Weiss,the reduction of biological problems to merephysical and chemical aspects fails to touchthe core of the problem of life.

Besides animal behavior, a field in which heperformed only a few studies, Weiss is alsosimilarly critical of developmental research,where his main interest lies, and to whichLoeb also made contributions (Weiss 1922).

a conceptual apparatusWeiss points out the importance of theo-

retical considerations, but he uncriticallyholds that the description of facts does notchange, whereas theories, which are formu-lated to integrate the findings into a largercontext, may change (Weiss 1925:168–169).In contrast to the “reductionist” attitude,Weiss states that it is more promising and ec-onomical to first determine the laws in a cer-tain research field such as biology, beforeseparating it into physics and chemistry likeLoeb did. Accordingly, it is important that bi-ology develops its own scientific conceptualapparatus [Begriffsbildung] (Weiss 1922:2, 8).The basic terms of a conceptual apparatusfrom Weiss’s perspective are listed below.

Even his early work contains the basis of ahierarchical order necessary to allow actions(such as locomotion) of an animal as a whole.This concept of hierarchy is also applied inhis work in developmental biology. Accord-ingly, with each level (be it a physical or chem-


ical reaction, a single muscle fiber contrac-tion, a contraction of the muscle, a movementof the organ, or a move by the whole organ-ism), new features appear that cannot be ex-plained simply by constituents of lower levelsalone. By splitting the whole into lower-levelelements through quantitative analysis, thefeatures of the whole may become lost evenmethodologically. This would also be true forinorganic entities such as molecules or atoms.Although a quantitative approach should beapplied, it can never lead to a complete ac-count of a complex entity, because when suchan entity is fragmented, some characteristicsare lost. “Higher complexes” can be viewed asunits for which certain laws can be foundwithout reducing them to their components(Weiss 1925:173–174).

Another possible term in a conceptual ap-paratus is the “system.” In his chapter on “lawsof systems,” Weiss proposes an early systemdefinition: “As a system we want to defineeach complex that, when parts of it are mod-ified, displays an effort to stay constant withregard to its outside” (Weiss 1925:181, 183).In other words, the state of a system shouldbe stable within and distinct from the outsideconditions. Therefore, Weiss defines a systemby its behavioral feature of reaction. This no-tion was not new and can be found in Her-ing, as mentioned previously; in GustavFechners’s “principle of tendency towardsstability,” according to which every naturalsystem, closed or open to external influ-ences, tends toward relative stationary states(Heidelberger 2004); and in Henri Le Cha-telier’s principle for chemical reactions,where the change in one of the parametersleads the others to modify in a direction thatcompensates this change and enables the sys-tem to return to an equilibrium state. AsWeiss interprets it, a constant system state inthe whole, when parts are altered, could onlybe achieved by inverse changes in its parts,which is done by the system itself (system re-action). He continues:

The appearances of regulation and ad-aptation are typical system reactions ofthe organism. They are so widespreadthat their mere existence suffices tomake the organism recognizable as a sys-tem (Weiss 1925:185).

Weiss distinguishes three different “systemreactions.” In the first one, an outside factorapplied to the system results in a reaction ofa single part, such as in simple reflexes. In thesecond, more than one part of the system isaffected by an outside factor. And in the finalreaction, an outside factor does not trigger areaction of single parts of the system, but ofthe whole system, where the reaction of thewhole is not a result of the reactions of theparts: it is the other way around, and the re-actions of the parts are a result of the overallreaction (Weiss 1925:175–181). The third re-action—which he illustrates by examplesfrom physics and Kohler’s Gestalt theory—isimportant for his argument that an organismis not merely a conglomerate of cells, mole-cules, or atoms. This notion also allows lawsto be found without knowing all of the detailsof the involved elementary processes, al-though the latter are important as well. Weissemphasizes that the third case has no rigidmechanism. Rather, the parts show some plas-ticity, while the reaction of the whole is lawful.The physicist’s descriptions of processesbased on “parts and their relations” is verymuch in line with that argument (Weiss1925:186–187).

With regard to dynamic processes and Ber-talanffy’s concept of open systems in “fluxequilibrium” [Fließgleichgewicht], which was in-fluenced by several authors, it is interestingthat Weiss—when discussing system reac-tions—picks up Hering’s “schema of metab-olism.” Equal rates of assimilation and dissim-ilation yield a metabolic equilibrium. Thesame idea is emphasized in 1920 by AugustPutter, another organismic thinker, in his vol-ume on growth equations, which had a sig-nificant influence on Bertalanffy’s own the-ory of growth: the end of growth correspondsto this equilibrium (see Pouvreau 2005b). Forexample, if dissimilation is accelerated by anoutside change, then following the laws ofchemical kinetics—which Weiss calls “systemlaws of chemistry”—assimilation is also accel-erated (Weiss 1925:193–194). He argues thatif this concept is viewed beyond the chemicalcontext, it can serve as system law for livingorganisms.

The historical approach is also fundamen-tal for biology. Weiss discusses teleology on


the one hand and chance on the other, buthe is not satisfied with either concept andclaims that something such as an arch con-sisting of single bricks can never come intobeing. A mere mechanistic “causal analysis,”however, cannot solve such a problem either.The solution of the contradicting perspec-tives is seen in the historical development ofvarious processes where, from the very begin-ning, the parts are subordinated to the whole(Weiss 1925:190–191). The problem of rec-onciling the system perspective and the his-torical perspective was also mentioned byJean Piaget when referring to the centralproblem of every structuralism: “have the to-talities through composition always beencomposed, but how or by whom, or have theyfirst been (and are they still?) on the way ofcomposition?” (Piaget 1968:10). The impor-tance of this problem is also emphasized byLeo Apostel (1970:167): According to him,structuralism is so dominated by the notionof equilibrium that it does not succeed to in-tegrate genesis and history; the problem iscentral both to structuralism and system the-ory.

epistemological considerationsWeiss is deeply influenced by Mach; for

example, he refers to the antireductionistprogram of the positivist thinker (Hofer2000:151). Although Weiss contradicts theidea that biological activities can be explainedby physics and chemistry alone, without fur-ther requirements, he leaves open the ques-tion whether or not physical or chemicalterms could replace biological ones in the fu-ture. He does claim that the laws of complexissues will never be fully replaced (Weiss1925:170).

With regard to prevailing positions of re-ducing biology, Weiss suggests that, notwith-standing progress within biophysics and bio-chemistry, the latter are only concerned withparts. The laws concerning the relationshipsamong all parts are still the realm of biologi-cal research. But they must not remain mere“rules,” they have to become “laws.” The bio-logical laws should be as strict as those ofphysics and chemistry, and should be able tounite several subdisciplines within biologywithout losing autonomy. This is the outlook

that Weiss (1926:43) provides. The funda-mental distinction between “rules” and “laws”had already been emphasized by Roux andPrizbram in biology, but also by physicistssuch as Walter Nernst (Przibram 1923; Nernst1922); and it was a recurrent theme in sub-sequent reflections on theoretical biology,particularly by Max Hartmann (1925) andBertalanffy (1928). It is another expression ofthe Kantian distinction between “natural sci-ence” as “a whole of knowledge ordered byprinciples” a priori, of which “the certainty isapodictic,” and “natural history” as a “descrip-tion of nature,” of which the certainty is onlyempirical, and that is knowledge only in aninappropriate sense (Kant 1786).

We agree with Khittel when he states thatWeiss’s epistemological background is rathersimple, and that he uncritically combines the-ory and experiment (Khittel 2000:160). Weissis satisfied with posing questions to naturethat are answered by the experiment anddoes not go deeper into epistemology.

The experimental biologist Weiss never de-veloped a “system theory.” Rather, from theinsight he obtained by his experiments andobservations, he described what such a theorymust comprise and where the problems liewith the “analytico-summative” approach, aswell as its abstraction and shortcomings. Thissheds light on Weiss’s statement that he onlyhad “marginal contacts with the field [ofgeneral systems theory]” in his later life(Weiss 1977:18). Weiss was only minimallyconcerned with systems beyond the singleorganism, but he does consider them occa-sionally—for example, at the Alpbach sym-posium (Koestler and Smythies 1969); he alsorefers to communities and societies whentalking about systems (Weiss 1970b:9). One ofhis last descriptions of systems broadens thescope even more: “a system is an empirical en-tity which in our experience has sufficiently durableidentity to be defined on a macroscale as a whole,although from our knowledge of the parts alone intowhich we can physically or mentally fractionate theentity on the microscale, we could never retrieve therule of order discernible in the intact macrosystem”(Weiss 1977:55). Furthermore, he states thatthis holds true throughout the hierarchy ofnatural systems, from stellar bodies, organ-isms, and molecules to subatomic units. As


Figure 4. Ludwig von Bertalanffy (1901–1972)� Osterreichische Nationalbibliothek/Wien NB

533.771-B.

the following quotation demonstrates, Weissadvocated, in his late writings, system think-ing on a broad and deep basis:

It is an urgent task for the future to raiseman’s sights, his thinking and his acting,from his preoccupation with segregatedthings, phenomena, and processes, togreater familiarity and concern withtheir natural connectedness, to the “totalcontext.” To endow the epistemologicalfoundations for such a turn of outlookwith the credentials of validation by mod-ern scientific experience, is thus a majorstep toward that goal (Weiss 1977:19).

Ludwig von Bertalanffy’sOrganismic Concepts

Ludwig von Bertalanffy, born to a family oflower nobility, was (from an early age) onlyinterested in the works of Jean-Baptiste de La-marck, Charles Darwin, Karl Marx, and Os-wald Spengler (Hofer 1996; Pouvreau 2006).In his youth, Bertalanffy lived on his mother’sestate outside of Vienna next to Paul Kam-merer, one of the most prominent “Prater Vi-varium” experimenters. Being a neighborand friend of the family, Kammerer oftentalked to the young Ludwig about biologicalmatters. Thus, Bertalanffy, even before hisstudent years, was probably well aware of thespecific style in which research was conductedat the Biologische Versuchsanstalt laborato-ries.

At the end of World War I, the Bertalanffyfamily temporarily left Vienna and moved toZell am See (Salzburg, Austria) because of fi-nancial losses. Ludwig started his universitystudies, focusing on history of art and philos-ophy (especially metaphysics, logics, and phi-losophy of religion), in Innsbruck (Tyrol,Austria). To a lesser extent, he was alsotrained there in botany by the experimentalmorphologist Emil Heinricher and plantphysiologists Adolf Sperlich and Adolf Wag-ner—the latter being one of the main advo-cates of a “psycho-vitalistic” philosophy of life.In 1924, Bertalanffy returned to Vienna,where he still focused on history of art andphilosophy (especially philosophy of knowl-edge). As for biology, his only courses there

(in 1925) can be labeled as “metaphysics oflife” and dealt especially with Henri Bergson’sthinking. Indeed, the student Bertalanffy wasessentially what one might call a well-in-formed autodidact for biological matters. Heearned his PhD in 1926 with a dissertation onGustav Fechner’s metaphysics of organizationlevels, which already stressed and discussedsome contemporary ontological controver-sies in biology. Between 1926 and 1932, hedevoted most of his studies to philosophy ofbiology, justifying and developing a project of“theoretical biology.” He received his Habili-tation in 1934, which was in “theoretical biol-ogy.” He lived in Vienna until 1937, develop-ing both his theoretical perspectives andsome experimental applications in the fieldof animal organic growth. A Rockefellerscholarship was granted to him for one year.He worked in the United States from October1937 on, especially in Chicago (with NicolasRashevsky and his coworkers) and in the Ma-rine Biology Laboratory in Woods Hole (Mas-sachusetts). After Hitler’s Anschluss of Austria


in March 1938, Bertalanffy tried to find a wayto stay in the United States. But conditionswere unfavorable in that regard, with prioritybeing given to persecuted scientists: he wasforced to return to Austria in October 1938.The dark period of Bertalanffy’s life thenstarted. Despite his deep disagreements withthe essential parts of Nazi ideological com-mitments, he became a member of the NaziParty in November 1938. It was a means tofacilitate his promotion as a professor at theUniversity of Vienna, which he was awardedin September 1940. During the war, in par-allel to the elaboration of his theory of or-ganic growth, he closely linked his “organis-mic” philosophy of biology to the totalitarianideology in general and the Fuhrerprinzip inparticular. Bertalanffy lived in Vienna until1948, when he decided to leave permanently.He was impatient regarding his career oppor-tunities in post-1945 Austria because of un-favorable outcomes of his personal “denazi-fication” procedure and because of theenmities induced by his opportunistic behav-ior during the war (Hofer 1996; Pouvreau2006). He then went through Switzerland andEngland to Canada.

Bertalanffy was influenced by many people(see Pouvreau and Drack 2007). From thephilosopher’s side, it was by no means onlythe Viennese Circle but, among others, alsoincluded neo-Kantian thinkers such as HansVaihinger and Ernst Cassirer. His organismicbiology was largely influenced by JuliusSchaxel, Emil Ungerer, Nicolai Hartmann,Hans Spemann, and Friedrich Alverdes, butto some extent also by the “Prater Vivarium”and Paul Weiss.

bertalanffy meets weissWhen still a student, Bertalanffy met the

recently graduated Weiss and they discussedsystem issues in biology, Weiss with an exper-imental and Bertalanffy with a philosophicalbackground. Five years after Bertalanffy’sdeath, the 79-year old Weiss remembers:

Those days that a sparkling Viennese stu-dent, a little more than three years myjunior, approached me for a meeting—Ludwig von Bertalanffy. We met in cof-feehouses and “milked” each other. I

soon found that his thinking and minemoved on the same wave-length—hiscoming from philosophical speculation,mine from logical evaluation of practicalexperience. And so it remained for halfa century, each of us hewing his separatepath according to his predilection. Thatis, I kept on as the empirical experimen-tal explorer, interpreter, and integrator,for whom the “system” concept re-mained simply a silent intellectual guideand helper in the conceptual ordering ofexperience, while he, more given to ex-trapolations and broad generalizations,and bent on encompassing the cosmosof human knowledge, made the theory[general system theory] itself and the ap-plicability of it to many areas of humanaffairs his prime concern. Does not thisconfluence here, once again, prove the“hybrid vigor” of the merger of ideasthat, coming from a common source,have converged upon common ground,albeit by separate routes—the one offer-ing a distillate of a life of study of livingsystems, the other the extensive elabo-ration of an intuitive philosophical ide-ology, tested in its pertinence to humanevolution? (Weiss 1977:18–19).

During the last three months of Berta-lanffy’s initial U.S. visit ( July through Septem-ber 1938), he worked with Weiss at WoodsHole (Hammond 2003; Pouvreau 2006). Thiswas their last meeting until 1952. In autumnof that year, they met again at the Universityof Austin (Texas), where Weiss was working.At that time, Bertalanffy was on a tour, mak-ing about 15 conference presentations in dif-ferent parts of the country; he came to Austinat the invitation of the biochemist ChaunceyD Leake. Weiss probably played a role in thisinvitation (Pouvreau 2006:76). Years after go-ing abroad, Bertalanffy and Weiss met againin Austria at the small alpine village of Alp-bach, Tyrol, on the occasion of a special sym-posium devoted to systems thinking, orga-nized by Arthur Koestler in 1968 (Koestlerand Smythies 1969).

Bertalanffy’s written estate (Bertalanffy pa-pers), which was found in 2004, yields no evi-dence of communication between Weiss and


Bertalanffy after 1945. As for the period be-fore 1945, no letters to or from Bertalanffyare conserved in this estate, because his homewas burned at the end of World War II.

early works of bertalanffyIn the course of his studies, Bertalanffy fo-

cused on philosophy, having Robert Reinin-ger—the leader of neo-Kantianism in Austriaand president of the Viennese PhilosophicalSociety (1922–1938)—and Moritz Schlickamong his teachers. Bertalanffy recognizedhis student days in Vienna as a time of highintellectual intensity, triggering many devel-opments that later became important for him(Bertalanffy 1967). Concerning the positivistideas of the “Vienna Circle” around Schlick,Bertalanffy logically regards empiricism as anerroneous doctrine. He holds that empiricaldata have to be ordered by laws, because “onecannot base and develop any science solelyon experience and induction” (Bertalanffy1927a:660). Referring to Kant (1787:A51/B75), he liked to say that, though theory with-out experience is mere intellectual play, ex-perience without theory is blind (Bertalanffy2003:101). Furthermore, when Bertalanffyoutlines the difference between science andmetaphysics, mythical thinking, and mysti-cism, he argues that a dialogue between themis necessary. For example, he writes: “Themost noble task of science is that mysticismdoes not bring us back to the night of igno-rance and faith, but that it becomes itself ameans for the deepest understanding of na-ture” (Bertalanffy 1927b:264). Convergenceswith positivism can nonetheless also befound: especially a taste for logical analysis ofconcepts, the rejection of substantialism (seeMach 1906 and 1933), and the will to save thedignity of science in the face of the rise ofirrationalism—a characteristic trend in theinterwar German and Austrian context.

Bertalanffy’s doctoral thesis (Bertalanffy1926a) was concerned with intriguing ques-tions: Can units of higher order (integrating,from Fechner’s point of view, the living or-ganism) be methodically investigated in anabstract manner? Can they be made empiri-cally plausible? To what degree can an induc-tive metaphysics be based on a generalized

concept of organization, as opposed to spec-ulative metaphysics? His dissertation alreadycovers the stratified levels of biology, psychol-ogy, and sociology, and he emphasizes the“perpetual recurrence of the same in all levelsof integration [Integrationsstufen]” (Berta-lanffy 1926a:49). To illustrate this, he employsthe model of the atom as a small planetarysystem, reviving the ancient doctrine of thehomology of macrocosmos and microcosmos.Note, however, that this model was already re-garded as erroneous at that time in physics,after having been understood as a useful fic-tion.

After graduating, he wrote many articles inthe field of theoretical biology. In 1928, hepublished his “critical theory of morphogen-esis” (Bertalanffy 1928), which acquaintedhim with the Berlin Society for ScientificPhilosophy and Hans Reichenbach—a coun-terpart to the Vienna Circle—and openedcontacts with Gestalt psychologists (Hofer2000:154). In that work, Bertalanffy regardsthe development of an organism out of agerm as a “paradigm” [Paradigma] illustratingthe need for an “organismic” and “theoreti-cal” biology. He therefore analyzes the vari-ous theories of development [Formbildung],among them the approaches of Przibram andWeiss (Bertalanffy 1928). Przibram’s analogyof a germ and a growing crystal is criticizedbecause the first one consists of inhomoge-neous parts, while the second is homoge-neous, although both have a “vectorial poten-tial” and tend to regenerate if disturbed. Alsocriticized is the lack of differentiation be-tween life and the inorganic realm (Berta-lanffy 1928:166–175). The “field theory” inWeiss’s developmental biology is categorizedby Bertalanffy (1928:189–190) as an organis-mic theory (see below). Bertalanffy con-cludes from the analyzed theories of devel-opment that “wholeness [Ganzheit], Gestalt, isthe primary attribute of life” (Bertalanffy1928:225). He uses wholeness [Ganzheit] andGestalt as synonyms (Bertalanffy 1929c:89).

In May 1934, Ludwig von Bertalanffy wasthe first academic at the University of Viennato receive a Habilitation in “theoretical biol-ogy.” The basis for this licensing procedurewas the first volume of his book on what hecalled Theoretische Biologie, which had already


been published two years earlier (Bertalanffy1932).

theoretical biology andepistemology

At a young age, Bertalanffy was probablyintroduced to the ongoing controversies inbiology. The neo-Lamarckian Kammerer hasalready been mentioned above. Kammerer’simpact is clear in Bertalanffy’s early criticismof selectionist or mutationist theories of evo-lution (Bertalanffy 1929a). But very early on,he distanced himself from the different kindsof neo-Lamarckism that were flourishing inthe first two decades of the 20th century as aconsequence of the inner weaknesses of Dar-winism (related in particular to its apparentcontradictions with the Mendelian theory ofheredity). He therefore criticized Kammerer(Bertalanffy 1928:95) not only for his “La-marckist” speculations, but also for his al-leged “theory free” empiricism. As a conse-quence of his early contact with the(psycho-)vitalist botanist Adolf Wagner, Ber-talanffy very likely had firsthand knowledgeregarding the criticism of the Darwinianschool and about attacks on biological “me-chanicism” (Reiter 1999; Hofer 2000; Pou-vreau 2006). Like Weiss, he was also verymuch influenced by Kohler, whom he re-garded as a “precursor” of his “general sys-temology” (Bertalanffy 1945:5; 1949b:115).Kohler, who wanted to extend Gestalt theorytoward biological regulation, coined the termSystemlehre (Kohler 1927), which Bertalanffyalso used and that we translate as “systemol-ogy” (Pouvreau and Drack 2007). Kohler’saim was to show that self-regulation by meansof dynamic interactions is a common abilityin both physical and organic systems.

An important figure regarding the need fora theoretical basis for biology was Schaxel,who in 1919 started a book series (Abhandlun-gen zur theoretischen Biologie; Treaties in Theoreti-cal Biology) dedicated to implementing theo-retical biology. It is not by chance that worksof Przibram (1923), Weiss (1926), and Ber-talanffy (1928) were published in that series.In the same year, he published an essay onthe basics of theorization [Theorienbildung] inbiology (Schaxel 1919). He also popularized

the term “organismisch” (organismic), whichwas coined by the entomologist LudwigRhumbler in 1906 (Schaxel 1919:203). Thisterm was used by Bertalanffy in order to char-acterize his own fundamental conceptions;the influences of both Schaxel and Ungerer(1922) are clear and explicit in the genesis ofhis ideas and his associated vocabulary.

Spemann’s works (1924) on “organizers ofdevelopment” also influenced Bertalanffy.The dependency on the positions of the partscan only be understood when considering theorganism as a whole, including processes ofdifferentiation and hierarchization.

Bertalanffy’s first book, in which he em-phasized the need for a theoretical biology,appeared in 1928. His motivation for goinginto biology is much related to the fact thatbiological categories (especially wholenessand organization) were very often and un-critically used in areas beyond biology, andoften served as a basis for dogmatic ideolo-gies.

The science of life has nowadays to a cer-tain extent become a crossroad, in whichthe contemporary intellectual develop-ments converge. The biological theorieshave acquired a tremendous ideological[weltanschauliche], yes even public andpolitical significance. . . . The conditionof biology, problematic in many respects,has led to the situation that the “philos-ophies of life” were until now by nomeans satisfactory from the scientific asmuch as the practical point of view; wesee all the more clearly the importanceof the theoretical clarification of biology(Bertalanffy 1930:4–5).

Another consideration demanding a soundepistemological basis in biology was its meredescriptive or experimental character, which,according to Bertalanffy, had to be overcome.He argues in the same line as Weiss in so faras both think that biology has to be autono-mous, and that life cannot be reduced to therealm of physics and chemistry alone, as “me-chanicists” (or rather, “physicalists”) wouldlike. There are two main justifications for this.First, biology needs to be seen with differentmethodological perspectives: physicochemi-cal, ganzheitliche (integrative or “holistic” in a


broad sense) or organismic, teleological, andhistorical (e.g., Bertalanffy 1928:88). The “an-alytico-summative” approach expressed bythe physical and chemical investigation of iso-lated processes leaves aside organization,which is a core problem of biology. Second,Bertalanffy regarded the conflict between“mechanicism” and “vitalism” as an essentiallymetaphysical one, which cannot be solvedempirically—this can also be seen as a criti-cism of empiricism. A new approach wastherefore necessary. Bertalanffy called it “or-ganismic biology” (Bertalanffy 1932).

In starting his theoretical program, he wasconvinced that biology’s right to exist as a sci-ence requires a theoretical background thathad not been established yet, although manyso-called “theories” were around. For him, bi-ology of this time was little more than somekind of natural history in a precritical (“pre-Copernican”—note the allusion to Kant)stage (Bertalanffy 1928:1–3, 51–56, 90–100;1932:1–9, 20–35). Empirical data have to beordered by laws, and Bertalanffy distin-guished between “empirical rules” and laws,a distinction that empiricists usually do notmake—as mentioned, Weiss did so but not inan elaborated way—and that consequentlyleads to confusion.

The rule is derived inductively from ex-perience, therefore does not have any in-ner necessity, is always valid only for spe-cial cases and can anytime be refuted byopposite facts. On the contrary, the lawis a logical relation between conceptualconstructions; it is therefore deductiblefrom upper [ubergeordnete] laws and en-ables the derivation of lower laws; it hasas such a logical necessity in concor-dance with its upper premises; it is not amere statement of probability, but has acompelling, apodictic logical value onceits premises are accepted (Bertalanffy1928:91).

Another confusion involves mixing de-scription, which is “the mere reporting offacts,” and explanation, which “means thelogical subsuming of the particular under thegeneral, the ordering of the facts standing inour way in more general relations” (Berta-lanffy 1932:23–24).

The role of the experiment is overesti-mated by empiricists and, from a logical pointof view, not justified. The law does not belongto the empirical realm. Furthermore, it can-not be induced because, in such a procedure,the “disturbances” in reality have to be knownbefore the law is derived from empirical data,which is logically impossible (Bertalanffy1928:93). Laws therefore have to be derived“hypothetico-deductively,” with experimentsbeing an incentive and instance of proof(Bertalanffy 1932:27–28). A final mistake ofempiricists is the illusion that a “hypothesisfree” science could possibly exist. All the ac-tivities of science contain a certain theoreticalmomentum. Scientists who are unaware ofthat instance might easily become victims ofmetaphysical or ideological bias.

According to Bertalanffy, biology must alsobecome an exact nomothetic science (Berta-lanffy 1928:55). On the one hand, a theory ofknowledge and methodology of biology is re-quired. On the other, general principles andlaws of life for deriving particular laws are alsopart of a theoretical biology, within which thedifferent fields of biology should also be syn-thesized and unified in a coherent system.Theoretical biology has to plan experimentsand strive for the conceptual mastery of thephenomena. Furthermore, the unstructuredaccumulation of empirical data and the spe-cialization into various fields, which obscuresthe links among the problems, must be over-come (Bertalanffy 1932:32–34). Nevertheless,establishing “exact laws” does not mean thatbiology has to be reduced, by “causal analy-sis,” to physics and chemistry. The essentialand sufficient character of a law is “to linkunivocally a definite initial state with a defi-nite final one,” thereby “it is not indispens-able” for such a determination “that all inter-mediary processes are known” (Bertalanffy1930–1931:400). This is similar to Prizbram’sposition but, of course, also to Boltzmann’sstatistical approach in thermodynamics towhich Bertalanffy repeatedly refers, or to thecontemporary approaches in quantum me-chanics.

Bertalanffy alleges that his “organismic” bi-ology, where the Ganzheit (wholeness) is heldas a fundamental characteristic of biologicalentities, is beyond “mechanicism” and “vital-


ism.” His “organismic program” aims at de-termining the corresponding laws that arecomplementary (in Niels Bohr’s sense) to theanalytico-summative approach. Furthermore,the concept of teleology is rehabilitated: notin a vitalistic way (in the metaphysical sense),but as a logical form of the maintenance ofthe whole. The basic assumption that theworld is stratified (N Hartmann 1912) is usedin order to connect causality and finality inconnection with the concept of dynamic equi-librium: “What in the whole denotes a causalequilibrium process, appears for the part as ateleological event” (Bertalanffy 1929b:390;1929c:102). His “de-anthropomorphization”of teleology—it might be less confusing toemploy the word “teleonomy” instead, whichwas introduced by Pittendrigh (1958)—anduse of a systemic category is in line with otherauthors such as Ungerer, Schaxel, Weiss, andNicolai Hartmann.

Just like the concept of energy is theform of expression for the causal law rul-ing the inorganic world, the concept oforganism is the form of expression forthe finalist [finale] perspective, which wehave to apply to the organism besides thecausal one. The characteristic of the or-ganism is first that it is more than thesum of its parts, and second that the singleprocesses are ordered for the maintenanceof the whole (Bertalanffy 1928:74).

Important in this context is the idea thatan “equifinal” state of the whole may bereached independently of the initial states ofthe parts and in the way they reach it. Thisconception is crucial in developmental biol-ogy and clearly contradicts a “mechanicist” at-titude.

The epistemology of “organismic biology”is based on the conception of the organismas a system, and can thus be called a “systemtheory” of life. The term Systemtheorie [“systemtheory”] first appears in Bertalanffy’s writingsin 1930–1931 (p 391), although he does notexplicitly define what is meant by system be-fore 1945—where a system is described as acomplex of elements in interaction (Berta-lanffy 1945). Mathematics also plays an im-portant role. Inspired by thermodynamics, heaims at the determination of “higher order

statistics,” where the causalities driving thesingle part are left aside, “exact laws” are nev-ertheless derived without reduction to thephysicist and chemist realm. Clearly, Berta-lanffy’s thinking and the attempts of Przibramconverge. With his growth equations, Berta-lanffy (1934, 1941) later shows how a mathe-matical approach in that direction can be im-plemented in the field of the dynamics ofmorphogenesis (Pouvreau 2005a,b).

It should also be mentioned that Bertalanffyrejects emergentism and holism in their origi-nal sense (Lloyd Morgan 1923; Smuts 1926).He considers them as belonging to metaphys-ics and as sterile in science—both implying anontological irreducibility of the whole to itsparts and relationships, even when they are ex-haustively known (radical qualitative novelty).He also easily resists critiques such as those ofPhillips (1976). Although that latter critique—toward a simple-minded Hegelian attitude ofholistic thinkers—focuses on the late works ofboth Weiss and Bertalanffy, it should be dis-cussed here because it also provides insightinto their early works. Bertalanffy’s positionmight be interpreted as echoing Georg W FHegel’s “doctrine of internal relations” (ac-cording to which the whole relational struc-ture is constitutive of each “part”) as well asholism, which is not the case. In contrast toPhillips’s critique, Bertalanffy’s statements arepurely epistemological. Weiss was probablytoo pragmatic and too much an experimen-talist to incorporate any Hegelian attitude.But the chief argument brought forwardagainst “Hegelian thinking”—namely, againstthe idea that the parts gain certain propertieswhen combined to a whole—cannot be sus-tained for the two biologists. Bertalanffy un-derstood the “whole” as the sum of its “parts”and their relations, and thought that this is allthat can be tackled by science. The only prob-lem lay in the epistemological difficultiesposed by “organized complexity”—as WarrenWeaver (1948) calls it—which sets into prac-tice the impossibility to satisfy the requiredcondition of knowing all the parts and theirrelations in order to know the whole:

The properties and modes of action ofthe higher levels are not explainable bythe summation of the properties andmodes of action of their components asstudied only in isolation. But if we know all


the components brought together andall the relations existing between them, thenthe higher levels are derivable fromtheir components (Bertalanffy 1932:99;1949a:140).

The system laws thus deal with the relation-ships among the parts: the existence of suchlaws relies on the idea that the relationalstructure may change, while the parts stay “in-trinsically” the same (i.e., as studied in isola-tion from the whole), so that the whole hasproperties of its own. Only in that sense isemergence tangible by science, and not in thementioned metaphysical sense (Bertalanffy1932:99). A later statement by Weiss is inter-esting in this context, which is very much inthe same line. In the phrase “the whole ismore than the sum of its parts,” Weiss inter-prets “more” not as an algebraic term, but re-fers to the “more” information that is neededto describe the whole compared to the parts.With this conception, Weiss bridges the gapbetween reductionism and holism (Weiss1970a:20).

Bertalanffy is also very much in line withWeiss (see above) when he states that bio-logical system laws are not once and foreverirreducible to physics, although Weiss refersto biological terms that in the future mightbe replaced, but he holds that the laws ofcomplex issues will never be fully replaced byphysics and chemistry (Weiss 1925:170).

To the question whether the concepts ofphysics are at present sufficient for sci-entific biology, the answer is no. . . . Tothe question whether the [purely biolog-ically applied] concepts can anyway bereplaced by physical ones, the answer is:wait and see (Bertalanffy 1932:113–114following Woodger).

If the system laws of the organism cannottoday be formulated in a physico-chemi-cal way, then a biological version of thoselaws is justified. The question whether anultimate reduction of the biological sys-tem laws is possible . . . is of secondaryimportance (Bertalanffy 1932:115).

organismic principlesBertalanffy formulates two general organ-

ismic “principles” or “working hypotheses”

(Bertalanffy 1932:331), which should moreappropriately be regarded as conceptualmodels of biological organization. These“principles” had already been previouslymentioned by others, but Bertalanffy unifiesthem and extends their scope from the singleorganism to biological organizations in gen-eral (from cell to biocoenosis), thereby open-ing the way from organicism to systemism.The first one is the “principle” of the orga-nized system as an “open system” in “fluxequilibrium” [Fließgleichgewicht] (Bertalanffy1929c:87; 1932:83–84, 116, 197; 1940a:521;1940b:43). This equilibrium cannot be com-pared to chemical equilibria because the latterare characterized by a minimum of free en-ergy. In contrast, the organism is an open sys-tem that maintains itself through a continuousflux of matter and energy, by assimilation anddissimilation, distant from true equilibrium,and is able to supply work. Metabolism is thusan essential property of the organism.

The second “principle,” also first enunci-ated in 1929, is the “striving of the organicGestalt for a maximum of formness [Gestaltet-heit]” (Bertalanffy 1929c:104). This is thoughtto play a role in ontogenesis as well as in phy-logenesis, and later becomes the “principle ofhierarchization” or “principle of progressiveorganization (or individualization)” (Berta-lanffy 1932:269–274, 300–320). The dynamicinteractions in the system give rise to order.The principle is very much related to epige-netic phenomena and comprises the devel-opment from an initial equipotential state(with maximum regulation abilities) over seg-regation processes. This is followed by differ-entiation and specialization, where some sortof centralization, with “leading parts” thatcontrol the development of other subsystems,can also occur. Characteristic is the inherenttrend toward an ever-increasing complexity—a trend that he later (Bertalanffy 1949a) calls“anamorphosis,” after Richard Woltereck hadcoined the term in 1940.

Besides the “principles” of wholeness anddynamics, a third one, namely that of primaryactivity of the organism, becomes explicitonly later when Bertalanffy’s organismicthinking has matured (Bertalanffy 1937:14,133–134). It was nonetheless implicit in manyof his writings between 1927 and 1932, and


also has ancient roots. It must be viewed inopposition to a mere passive concept of or-ganisms that merely react to the outsideworld. The concepts are not new, but werealready forwarded by researchers of Lebens-philosophie (“philosophy of life”), which wasan intellectual fashion in the first two decadesof the 20th century (Pouvreau and Drack2007). Weiss and Przibram also argue alongthe lines of primary activity, but Bertalanffydoes not refer to them in this regard. Berta-lanffy largely cites Weiss when discussing thetheory of the morphogenetic fields (e.g., Ber-talanffy 1928), although he at least onceclearly writes about system theory with regardto animal behavior without mentioning Weiss(Bertalanffy 1932:57–58). There, he refers tothe Erstarrung (congealment) into a machineas a secondary process and the Fahigkeit zumGanzen (ability toward being a whole) as pri-mary to the plasticity and regulation abilitiesof the most rigid, machine-like reflexes andinstincts of the organisms. But Weiss’s theoryof animal behavior is never explicitly men-tioned. Bertalanffy (1937:136–142) providesdetailed criticism of Loeb’s theory, but onlyFriedrich Alverdes is mentioned, not Weiss.In the 1920s and 1930s, Alverdes was the mainGerman advocate of an organismic approachto animal behavior (individual, but especiallysocial); his entire body of work is dedicatedto it.

beyond weiss’s conceptionsNote that Bertalanffy’s concerns went be-

yond the convergences with Weiss’s concepts.The program of his “organismic biology”comprises four dimensions. It is (1) a scien-tific program for opening the way toward con-structing exact theories of biological systemsof different levels. It also is (2) a natural phi-losophy of the biological world, where the twoprinciples mentioned above “probably havein the organic realm a similar basic impor-tance as the physical fundamental laws for in-organic nature” (Bertalanffy 1932:282). Es-sential for this philosophy is the stratifiedcharacter of reality and the autonomy of itslevels, where fundamental system laws can befound at each level—a scheme already dis-cussed in his doctoral thesis (Bertalanffy1926a). Furthermore, the dimension of (3) a

basis for developing an inductive metaphysicshas to be mentioned. Metaphysics can, ac-cording to Bertalanffy (1932:283), find a solidfundament in “organismic” biology for re-newing answers to old problems. Finally,there is (4) an ideological dimension, insofaras “organismic” biology is supposed to lay thefoundations of an alternative worldview be-yond “mechanicism”—a foundation in whichthe Ganzheit [wholeness] plays a central role(Pouvreau and Drack 2007).

The fact that Bertalanffy’s “organismic biol-ogy” was not an empty program is exemplifiedby his theory of organic growth, which eventoday remains a central reference in that field.In this theory, global growth and relative (al-lometric) growth are tackled not only by find-ing growth constants, but also by linkinggrowth with anabolism and catabolism. Thisstep had not been made before, and combinesthe open system concept with equifinality (seePouvreau 2005a,b; Pouvreau and Drack 2007).As already described above, this is in line with“laws of higher order” and “mathematical mor-phology” as mentioned by Przibram.

Conceptual ConnectionThe fields of interest of Bertalanffy and the

“Vivarium” biologists overlapped consider-ably. The “Vivarium” tackled some of the ba-sic questions in biology experimentally andBertalanffy strived to set up a theoretical basisof biology. Unsurprisingly, they argue in simi-lar fields, such as developmental biology,which comprises fundamental biologicalquestions; indeed, observations from embry-ology also influenced Bertalanffy’s “systemtheory of life.” The research style at the “Vi-varium” clearly offered one, although not theonly one, introductory approach to issues of“system theory.” When Weiss and Bertalanffymet in 1920s Vienna, their discussions verycertainly centered around concepts that laterconverged in Bertalanffy’s “system theory.”Bertalanffy, however, was also well acquaintedearly on with the works of Przibram, and firstmentioned Przibram in his first paper in thefield of biology (Bertalanffy 1926b).

Bertalanffy’s concepts in his “system theory”are not all congruent with those of Weiss. Nev-ertheless, some ideas converge, and Weiss’swork probably provided motivation. Other-


wise, Bertalanffy would not have mentionedhim at the Alpbach symposium as a key figureregarding the system approach in biology.Weiss was one of the first researchers who notonly described but effectively grasped the or-ganism as a system. The experiments that heperformed support the system approach of theorganism and, thereby, also Bertalanffy’s “or-ganismic biology.” Like Bertalanffy, Weiss ad-dresses “system theory” later on in his career.However, the development of both thinkerswas more parallel, and the influence of oneon the other was only minor.

The conceptual connections betweenWeiss and Bertalanffy can be summarized asfollows: Both think dynamically and in hier-archical terms, and both consider “whole-ness” as a key conception for understandingliving systems or organisms as systems, as op-posed to a “mechanicist” approach. In this re-gard, the principle of primary activity is cru-cial for both. The idea of “conservation” isalso fundamental to their system concepts.Weiss’s tentative definition of system and for-mulation of “system laws” also has a certaingenerality, which is paralleled in Bertalanffy’slater effort toward a “general system theory.”Both also argue for biology as an autonomousdiscipline with its own conceptual apparatus,which cannot be reduced to physics andchemistry; and for biologists to find laws of“higher order.” Nevertheless, Bertalanffy wasmuch more interested in establishing a theo-retical biology than Weiss and, therefore, hisepistemological attempt was more sophisti-cated.

Potential Contribution to FutureSystem Approaches in Biology

As the conceptual connection betweenWeiss and Bertalanffy arises from both exper-imental and philosophical considerations,these can serve as a useful framework for fur-ther research regarding biological systems. Inrecent years, “systems biology” (see Kitano2002) and “biocomplexity” research (seeMichener et al. 2001) have appeared as newfields, often in order to provide quantitative(computer) models for complex biologicalphenomena.

Although there is currently no widely ac-cepted definition (Kirschner 2005), “systems

biology” can be described as the study of theinteractions among the components of bio-logical systems and the consequent functionand behavior of the system as a system (i.e.,how a biological whole behaves over time).This already shows that “systems biology” goesbeyond a mere “analytico-summative” ap-proach. Still, quite often concepts only fromengineering or cybernetics are used to modelbiological systems; for example, when feed-back loops are introduced or when robust-ness is investigated. These concepts are oftenderived from physics and do not comprisecentral characteristics of life. Woese (2004)criticizes this engineering approach, whichhe already observes in the Human GenomeProject, a pivotal kickoff impulse for “systemsbiology.” Some scientists aim to explain lifewithin “systems biology” on the basis of mo-lecular interactions with emphasis on genesand proteins. Furthermore, the question ofhow the phenotype is generated from the ge-notype is important within “systems biology,”as is the correspondence between moleculesand physiology at a larger scale.

Kirschner (2005) notes that the perfor-mance of a protein is not only determined bya gene, but that it depends on the context inwhich it occurs, and this context is history de-pendent. The historical dimension of all liv-ing systems can be detected in the work ofWeiss and Bertalanffy as well.

The concept of hierarchy was already cen-tral to Weiss and Bertalanffy. Thus, they werefar removed from what Mesarovic and Sreen-ath (2006) criticized as a “flat earth perspec-tive in systems biology.”

In addition to the concept of control, asstemming from engineering feedback sys-tems, the importance of coordination in liv-ing systems has recently been pointed out byMesarovic and Sreenath (2006:34): “Coordi-nation . . . provides motivation/incentives tothe parts to function in such ways that theycontribute to the overall system’s goal whileat the same time they pursue their local ob-jectives.” This parallels the concepts of“wholeness” and “conservation” of Weiss andBertalanffy.

An engaged warning has been utteredagainst a possible new “regression into over-simplification” (Newman 2003:12) that might


prevail in “systems biology.” Here, the con-cepts of Weiss and Bertalanffy might helpavoid the risk of “mechanicism.” As shown,some of these concepts have already been in-corporated or encouraged to be integrated in“systems biology.” Primary activity, however, isnot an issue in “systems biology” today, butwas acknowledged as important by Weiss andBertalanffy.

Although skeptical about earlier system at-tempts in biology, O’Malley and Dupre(2005) emphasize that further philosophicalefforts have to be made within “systems biol-ogy.” This is what they call “systems-theoreticbiology,” as opposed to “pragmatic systems bi-ology,” which is concerned with interactionsof molecules on a large scale.

As we have demonstrated, Bertalanffy al-ready contributed much to epistemologicalissues in his early German publications on or-ganismic biology that approached the organ-ism as a system. The data-driven approachfound in modern “systems biology” is oftenseen as a necessary first step. Theory from(not only) Bertalanffy’s point of view, how-ever, clearly plays an important role especiallyat the start of a new scientific endeavor: theo-

retical work must be combined with experi-mental efforts at every stage. This is preciselywhat he intended with his theoretical biologybased on an organismic approach.

As Kirschner (2005) remarks, “systems bi-ology” is not a branch of physics. In the viewof Weiss and Bertalanffy, it was self-evidentthat biology is an autonomous discipline andhas to work with its own conceptual appara-tus. This prompted them to develop a systemtheory of life.

acknowledgments

This research was supported by Austrian Science Fund(FWF) Grant P18149-G04 to Rupert Riedl (1925–2005), who unfortunately passed away shortly after theresearch project was initiated. Riedl was one of Ber-talanffy’s students in 1940s post-war Vienna and lateron actively emphasized the importance of “Systemtheo-rie” [system theory] and “Langsschnitttheorien” (longi-tudinal-section theories)—not only in biology, but alsoin other sciences and in humanities as well (e.g., Riedl1978, 2000). His emphasis on the importance of es-tablishing a theoretical biology was also influenced byBertalanffy. Later on, Weiss, a fatherly friend as Riedlliked to say, also exerted an influence on him. We fur-thermore wish to thank Ludwig Huber and Gerd BMuller for their support.

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