ON THE MAKING OF A SYSTEM THEORY OF LIFE: PAUL A WEISS AND LUDWIG VON BERTALANFFVS GONCEPTUAL GONNEGTION

7/25/2019 ON THE MAKING OF A SYSTEM THEORY OF LIFE: PAUL A WEISS AND LUDWIG VON BERTALANFFVS GONCEPTUAL GO

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VOLUME 82 No. 4 THE QUARTERLY REVIEW

OFBIOLOGY DECEMBER

2007

ON THE MAKING OF A SYSTEM THEORY OF LIFE:

PAUL

A

WEISS AND LUDW IG VON BERTALANFFVS

GONCEPTUAL GONNEGTION

M A N F R E D D R A C K

Departffum l

of

Theoretical BiologyandDepartnumtof Neuroffiology

and CognitionResearch,

University

of

Vienna

1090 Vienna, Austria

E MAIL MAN H i E D .D R A C K U N I

VIE.

AC .A T

WiLFRIED A P F A L T E K

Deparlvumt ofPhilosophyand Department of Nnirobiology

ami Cognition

Research,

University

of

Vienna

1090 Vienna, Austria

E-MAIL: [email protected]

D A V I D P O U V R E A U

Schoolof Advanced Studies

in

Social

Sciences,

Koyre Center

75231 Paris, France

E M A I L

D A V D . t ' O U V R E A U @ W A N A D O O . F R

KEYWOR DS

orgaiiismic biology theoretical biology Biologische Versuchsanstalt

systems biology

A B S T R A C T

In this ariide, me revitw howlino enmient Viennese system Ikinkeix Paul A Wens and Ludxidg

von Berfnlanffji, begun to develop their ownperspectives Cowardiisystem, tfirary of life in the 1920s.

Their luoi-k

is

especially rooted

in

pxperimental biology as performed

at

the Biologische Versuchsamtatt,

as tvell

as in

philosophy,

and

they converge

in

basic imicepts. H>underline the concefHual connections

of their thinking, among them

the

organism

as an

organized system, hietarchical organization,

and

primary activity. W ith their system thinking, both biologists shared

a

.strong desire

to

nverrome what

they vieu>ed

as a

mnhanistir approach

in

biology. Their interpretations are relevant tot.heren.aissance

of system thinking

in

biology systems biology. Unless otherwise

noted all

transUuions are

our

oum.


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35 0

THE QUARTERLY REVIEW OF BIOLOGY

VoLUMi: 82

I N T R O D U C T I O N

N TERMS OF publications, systems biol-

ogy is curre ntly an explod ing field (ste ,

e.g.. Ch ung and Ray 2002; Gra nt 2003). No te,

however, that sy.steni thinking is not at all new

in biology. In the .sciences of life, it can be

traced back at least to the Ge rma n teleo-me-

chanicist tradition inatigurated by Imm an-

uel Kant's

Criticsofjndgernmt

of which Joh an-

nes Muller and Karl von Baer were the main

pro pon ents (Kant 1789: Lenoir 1982), More

generally, integra tive (or holistic ) thin king

is no t a discovery of the 20th cen tury . It has

a rather long history going back to ancient

Greek thinke rs: Aristotle already claimed that

tlie whole is of necessity prior to the pan

(Aristotle 1966:I253a20). Between the I4th

and 16th cen turies , it was a major intellectual

trend, which strongly reemerged at the turn

of the 19th ceniury in German Romanticism

and post-Kantian idealism. Among others,

Nicholas of Gusa, Gottfried W ilhelm Leibniz,

and Joha nn WoIIgang von Go ethe are consid-

ered precurs ors of m od ern systems think-

ing (B app 1921: Bertalanffy 1926a; Harr ing -

ton 1996). One of the leading figures, who

recurrently referred to those thinkers, is the

philosopher and theoretical biologist Ludwig

von Bertalanfiy (1901-1972), mostly known

for his project on a general system theo ry

(GST). Unsurprisingly, he is still regularly

quotediUtliough mostly laconicallyby ad-

vocates of systems biology in the early 21st

centur y (e.g., Chon g and Ray 2002).

Pursuing the historical roots of biological

systems tlieory inevitably leads to eaily 20th

century Vienna. Ludwig von Bertaianffy lived

there and was infltienced by biologists Hans

Leo Przibram (1874-1944), the founder of a

laboratory for experimental biology, and his

student Paul Alfred Weiss (1898-1989), both

of whom were also ba.sed in Vienna. We will

more closely examine the convergences of

the experim ental and theoretical works of

Weisswho allegedly claim ed tha t he was re-

sponsible for Bertalanlfy

initial appreciation

ogy. The early writings of Weiss and Berta-

lanffy- can be fruitful for modern research;

they already developed many concepts that

can shed light on certain contemporary de-

bates and perhaps only need to be adapted lo

current intentions.

The puipose of this article is therefore two-

fold. We first provide a historical overview of

the works of these system thinkers, and then

refiect the early system conceptions on recent

developments in biology.

C O N T E X T O F E A RL Y V I E N N E S E

SYSTEM THKORY

The fin de siecle and the following years

were characterized by innovations in many

fields, among them science, arts, and poli-

tics.

In particular, the developments in phys-

ics and psychology, together with problems

in biology; provided an impetus that led to a

sophisticated system conception of organ-

isms.

Au important revolution took place in

physics, which many scholars from other dis-

ciplines liked to cite. A statement by Max

Planck, inspired by the new qtiantum me-

clianics and referring to the false dictum that

all physical processes caji be displayed as a

stringing together [Aneinanderidhungl of lo-

cal processes, summarizes the relevance for

system concep tion s: The physical world is

not simply a sum of spatial and temporal sin-

gle worlds running one be.sides die others,

and tnany phenomena escape

[entzi^ hni .sirli

the understanding when one does not con-

sider a physical object

[Gebilde]

as a whole

(Planck 1929:17).

At about tbe same time, a conceptual shift

occurred in another field, namely psychology,

which was also exten ded to the realm of phys-

ics. Based on the conceptions of tbe Austrian

philosopher Christian von Ebrenfels, iestnlt

theory was developed (Ash 1995); it empha-

sized the claim that an entity's total charac-

teristics cannot be discovered by simply sum-

ming up iis partial characteristics. Wolfgang


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WEISS ANDBERTA IANFFY SGONCEPTUAL CON NECTION

351

m ore than the sum of the parts; and trans-

positio n, e.g., a melody can be played in a

different pitch, but stays the same) as con-

stitutive for

Gestalt

(Kohler 1920). Similarly,

Bertalanfly (1928:69-70) characterized a

(iesUilt

as compri.sing properties that cannot

be found by simple addition of the compo-

tienLs' properties and that furthermore dis-

appear when the

Gestalt

is destroyed; he used

the term Gestalt synonym otisly to systemic

state [Systemzustank]

(Bertalanfiy 1929c:89).

BertalaiiHy ^vitnessed a controversy among

biologi.sis at the turn of the century between

mechanicisin and \itHlism (for co n te m po

rary analysis of this controversy see, in partic-

ular, Haldane 1884; Loeb 1906; Schaxel 1919;

Ne edh am 1928; W oodger 1929; Bertalanfiy

1932; M Ha itinann 1925; and M H attm ann

1937; for a recent discussion see Pouvreau

2005b). Vitalism can have two mean ings , a

metaphysical or a methodological one. Meta-

physically, it a.s,serts tha t biologica l ph en om en a

cannot be explained withotit tlie action of a

nonspatial principle harmonizing tlie matter

and energies involved in living phenomena.

Metliodologically. it is not antinatui alistic and

only as.serLs that, at least provisionally, biology

should have its own categories, methods, and

laws.

As for the term me chanicism, it was

sometimes loosely referred at that time to

the opposed view that should be termed

physicalism in or de r to avoid confu sion;

that is, the idea that the sole con cepts, m eth-

ods, and laws of physics and chemistry en-

able biological phenomena to be grasped.

But this

w s

not the only mea ning . More gen-

erally and precisely, me chanicism seems

(and seemed) to refer to a complex group of

mo re or less coherently related positions. The

m ore fundam ental p osition is the analytico-

summ ative approa ch to biological phen om -

ena: its basis is tlie (methodological or meta-

phy.sical) postulate that any entity can be

analyzed in parts whose properties can be

studied ill isolation from others without in-

convenience (tbe relationships among the

parts being extern al, not constitutive ).

Throtigh decomposition into independ ent

causal chains and their linear com position,

repres ented in a biology that com bines the

analytico-summ ative app roac h with on e or

several of the following positions: physical-

ism, determ inism (each state is imivocally de-

rivable from previous state s), and reacUvism

(the changes in the behavior of an entity are

ascribable to the sole action of its environ-

ment ) .

An extreme example isjacques Loeb's

theory of tropism, which combines all of

these items.

A paradigmatic variant of m echanicism is a

machine-like model applied to the organ-

ism. This m ode l can be seen as a successor of

Descartes's bete m ach ine, which explains

life m erely by m eans of physics and chem istry.

Bertalanffy (1932:55) saw this Car tesian meta-

phor as one of the most unfortunate concep-

tions in the hi.story of science and philosophy.

Such a mach ine the ory was oppo sed by many

scientists. Some biologists in the late 19th

cen tury were con sidering a t>pe oi living mol-

ecule e.g., Darw in's g em inulae, Pfliiger's

living pro teins, or Haec kel's Plastidulen

(Penzlin 2000:434)although most physiol-

ogists rejecte d vitalism. Two findings w ere im-

portant for rejectbig vitalism (see Lenoir

1982;

Penzlin 2000): the first law of thermo-

dynamics and .successful physical


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352 THE QU RTERLYRESTEW OF BIOLOGY

VOLUMK 82

INSTITUTIONS OF EXPKRIMENTAI , BIOLOGY

Another debate in biology at that time was

on inetliodolog)'. Since the IS.^Os, a lively dis-

cussion arose between proponents of two sci-

entific approaches in biology: should mere

observation or experimentation be employed

(Nyharl 1995; Querner 2000)? Argtiing for

observation, traditional niorphologists stich

as Ernst Haeckel saw no sense in artificial, vi-

olent, experimental studies of development

because they considered it to be methodolog-

ically sufficient to thorotigh ly draw conc lu-

sions from observing nature's own diversity

and genuine transformations. In their view,

an experiment alters tlie events that normally

occur during developtnental processes and,

thus,

provides no causal explanation. A rguing

for experimentation, scientists such as Her-

mann von Helmholtz, Sigmttnd Exner, and

Wilhelm Roux favored well-constructed ana-

lytical experiments from which they hoped to

derive causal explanations.

Apart from older institutions stuh as the

Stazionc Zoologica in Naples {founded in

1872) or the Marine Biological Laboratory in

Woods Hole. Massachusetts (founded in

1888), a novel, avant-garde Viennese fiti de

siecie institution became famous for its ex-

perimental research on developmental and

evolutionary processes. This Biologische Ver-

suchsanstalt (Institute for Experimental Bi-

ology),

or popularly Prater Vivarium (as

there were other vivaria in Vienna, too), was

situated in a Neo-Renaissimce building at the

so-called Prater, Vienna's poptilar recreation

and en tertain m ent aiea (including parks, for-

est land, iuid fields) at the beginning of the

Pratei- Hatiptallee (Prater main avenue; Fig-

tire 1), Equipped with automatically regtt-

lated installations for controlling humidity

and temperature, the Biologische Versuch-

satistalt was an innovative research facility

both metho dologically and technologically. It

vras privately e stablished as a new type of bio-

logical researcb instittition in 1902. and

op en ed in 190.S, devo ted exclusively to exp er-

tackle all big question s of biology (Pr /ibrain

1908-1909:234) by transdisciplinarily apply-

ing similar experiments to both plants and

animals (e.g., with transplantation, breeding,

processes of selection, orientation, and polar-

ity in development). To perform those bio-

lf)gical experiments, methods from pbysics

and chemistry as well as physiology were ap-

plied to living organisms. Gonipared to phys-

iological rese arch , the Vivarium biologists

needed long-term experiments to deal with

questions of embryogenesis, regeneration,

and evolution. This called for methods that

allowed investigations over a series of gener-

ations.

Th ere w ere several de par tm ents at the Biol-

ogische Versuchsanstalt: one each for zool-

ogy, botany, and physiology of plants. For

some time, there were also phy.sioiogy and

physics/chemistry departments, Wolfgang

Pauli Sr., a colloid chemist and close friend

of Ernst Mach, headed the latter. ,AIthotigh

Przibram, Paul Kammerer, and Figdoi had al-

ready received their

alnUtation

(grand tmi-

versity teaching licen.se) at the University of

Vienna, the Prater Vivarium researchers

in many cases, liberal or left-oriented, open-

minded individtials with a [ewish backgroinid

(Re iter 1999; Goen 2006) were subject to ef-

ficacious disadvaruages orch estrate d by estitb-

lished, and often anti-Semitic, academics

from the university. However, effective Janu-

ary 1, 1914, tihe successively enlarged and

highly productive Biologische Versuchsan-

statt formally became a part of the Kiiiserliche

Akade mie der Wissenschaften (Imp erial

Academy of Sciences) in Vienna. At the sam e

time,

the three founders were appointed a.s

unsalaried heads of their departments, and

Paul Kammerer was appointed

djunkl

(civil

serv ant) , which, at that time, was the only po-

sition with a .salary.

There were also strotig international activ-

ities and lively network ing. Abou t one -third of

the positions were reserved for scholars from

abroad. Those gtiests included people such as

Karl von Frisch (Germany), Erich von Hoist


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D E ( ; F . M B K K

2007 WEISS ND BERTAEAMY S CONCEFWAL CONNECTION 353

Fu . t IRK

I.

T H E

B lOLOGISCH E VF.RSUCHSANSTAt.T

T he Prater Vivarium is depictedin theleft fo reground as seen from theRiesenrad. Vienna's giant ferris

wheel.Osterreichische N ationalbibliothek/Wien L 39.274


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3 54

THE QUARTERI.Y REVIEW OF BIOLOGY

VOLUME

82

botli tried

to

find

a

physical, chemical,

and

physiological basis

lor

ontogenesis.

In

pa i tic-

ular, Roux conceived Darw in's princ iple ofse-

lection

as

also implying

internal

selection.

When nicntioniiig

a

struggle

of

part.s wiihin

the organism (Roux 1881).

he

especially

highlighted

the

struggle

for

nutrients

and

space, Bertalanfty, although

in

deep disagree-

meiu with the Darwinian selectionist scheme,

later

(the

first time

in

Bertalanff)' I940b:56)

refers

to

this struggle of th e parts,

in

which

he sees

an

illustration

of ihe

metaphysical

(Heraclilean

and

Cusean) principle

of har-

mony through contradiction, when develop-

ing

his

theory

of

allometi

ic

growtli. Arguing

from Roux's point

of

view

and

considering

living beings

as

highly active entities inti-

mately related

to

their environm ents,

the bi-

ologists

of the

Biologische Versuchsanstalt

systematically questioned external selection

as

the

crucial drive directing evolution

(Gliboff 2005). This approach vaulted

the

Prater Vivarium staff directly into highly

controversial central issues

of

developmen-

tal

and

evolutionary biology. C onte m por ary

evolutionary developmental biology (EDB),

or evo devo , exam ines this role

of

devel-

opmen l

in

evolution

and in the

evolution

of

developmental mechanisms (Hall

et

al.

2003;

MO Ilerand Newman 2003). Wagner and Laub-

ichlcr (2004) recognize the Vivarium program

as

a

precursor

of

m ode rn eco^vo-devo.

Describing

the

laborato ry style

at

the Biologis-

che Versuch.sanstall. Przibram programmati-

cally declared: No specialization, [ bu tra ihe r]

a generalization

of

obtained experienc e is our

goal (Przibram 1908-1909:234 ). WiUi exper-

imental research,

not

only

on one

specific

group

of

organisms,

but

rather

on

\-arious

plants

and

animals,

he

aimed

at

finding

gen-

eral answers

to the

big questions

in

biology.

Pi-zibram proposed

to

ascribe

the

Gestalt,

developing u nde r

the

influence oft he inner

forces,

to a

.state

of

equilibrium

[Gleichge-

mchLszustand],

so

that

by

viewing organic

form

as a

state

of

equilibrium

. . . a

compen-

sation

of

disturbance

[ torungsausgleirh]

will

become accessible

to

ma them atical treat-

m ent (Przibram 1923:22-23).

In

this

con-

FiGURK a. HA NS LKO PR/.IBRAM (1^74-11)4,1)

Oster re ichischc Nal ionalbibl io ihck/Wiei i

NB

51U.429-B.

1888 w rote:

An

organism

is a

.system, which

is able

to

.su.stain

its

co nstitution

[Ikschajfcn

heit] (chemical iherm al state) against influ-

ences from outside,

and

which shows

a dy-

namic state

of

equilibrium

of

considerable

stability (Przibram 1906:222).

The

synony-

mous usage ofthe terms

Ofstalt

and form

is obv ious

in

Przibram.

He

refers

to the

influ-

ence

of

the whole

on the

parts

and how

this

can

be

dealt with mathematically. This

no-

tionas discussed below became im porian i

in Weiss's concepts, although

he was

less

in-

volved with mathematical details. Przibram

com mittedly discusses

the

possibility of apply-

ing mathematics

to

biology

in

general

and

t


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DECEMBER

2007

WEISS AND BERTALANFFY S CONCEPTUALCONNECIION

355

portio ns) should be described, also in relation

to physical measures such as surface tension.

Th e investigation shou ld reveal constants

that explain similar forms in various species

(Przibram 1923:23-25). Przibram also di^

cusses Fre deiic Houssay's project of a mor-

phologie dynam ique (Przibram 1923:9-11);

the fundamental idea here is to tmite mor-

phology and physiology, an improvement

compai ed to D'Arcy

Thompson's approach

of mathematical morphology. Thompson's

theory, or ratlier inductive method, which has

little predictive valite, is likewise criticized

by Przibra m (1923:13 -14) an d Bertalanffy

(1937).

Przibram 's oud ook toward a matli-

ematical mo rpho logy is very similar to the

notion on which Bertalanffy (1932:105) later

builds his con cep t of higher orde r statistics

(i.e.,

describing the average behavior of

high er units of cells or organism s, regardless

of physicochemical single events). This is

similarly already expressed by Przibram. His

example is the first formulation regarding

the course of organ ic global growth (precisely

the field where Bertaiantly' later brings bis

main contribution to mathematical biology,

see Pouvreau 2005b; Pouvreau and Drack

2007): It is, however, no t always necessary to

know tlie contribution or the nature of the

involved components of morphogenesis

[Formbildung] to arrive at mathematically for-

mu lated and theoretically usable laws (Przi-

bram 1923:25). With regard to the statistical

view on nature since Ludw'ig Boltzmann, Przi-

bram is influenced by his contemporary,

physicist Franz Exner (Coen 2006). Przibram

envisions and already initiates a m athe m ati-

cal theo ry of the org anic on the basis of quan -

titative inquiry, of tying tip witb geometry,

pbysics, cbemistry, and of sharp definitions of

biological term s. He expects this mathe-

matical biology to serve as a safeguard of

humankind against the forces of natttre and

namely as a bulwark against prejudice, super-

stition and untena ble speculation (Przibram

1923:64).

Furthermore, Przibram also opposes the

organisms are not mere playthings of envi-

ronmental forces.

P A U L W E I S S ' S S Y ST E M C O N C E P T I O N S

In 1918, Weiss took up mechanical engi-

neering, physics, and mathematics at Vi-

enna's Technische Hochschule, now called

Vienna University of Technology. After one

year of training in engineer ing, he decided to

study biology at the University of Vienna (for

Weiss's background and professional career,

see Brauckmann 2003). By 1920, he had al-

ready managed to receive one of the highly

coveted pe rm an en t positions at tbe Vivar-

ium laboratories, where he later con duc ted

work for his doctoral thesis (Weiss 1922;

Hofcr 2000:149). After Paul Kammerer re-

tired a nd left V ienna, W eiss, as his im me diate

successor, was appointed

Adjunkt

in early

1924.But he left tlie Vivarium in 1927 wh en

he applied for dispense of

his

working duties

(letter of March 25, 1928 and ptotocol no. 44

of Jan uary 31 , 1929, Vivarium pap ers. K2) in

Vienna in order to conduct research in Mon-

aco,

Paris, an d Berlin. He definitively resigned

from his salaried Adjunkl position in 1929. In

1931,

he was appo inte d Sterling Fellow at Yale

Universit)'. He went to the University of Chi-

cago in 1933 and became an American citizen

in 1939 (Haraway 1976; Brauckmann 2003).

Tb e Prater Vivarium becam e especially

important for Weiss and his later system

thinking ; so muc h so tbat at the Alpbach sym-

positmi iti 1968, Bertalanffy, when discussing

Weiss's talk, stated that, to bis knowledge,

Weiss was the first to intro du ce (he term sys-

tem in biology in 1924 (Koestler an d Smy-

thies 1969). As a matte r of fact, Bertalanffy

(e.g., 1932:87) knew very well that the philos-

op he r Nicolai Har tma nn (1912). with whom

he was very much in line, had already held

system as the fun dam ental category of bi-

ology as well as of otber fields (Pouvreau

2005a). Th e concep t of tbe harm onic equi-

pote ntial system as applied to the biological

organism was already the core of Driesch's

thinking since the end of the 19th century


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35 6

TH

QU RTERLY REVIEW

Of

BIOLOGY

VOLUME 82

F1GURK5.

P A U I ,

A1.PRHD

W EI S S

( 1 8 9 8 - 1 9 8 9 )

Cou rtesy of ihe Rockefeller U niversity A rchives.

W E I S S S EA R LY B U T rF .R F LY E X P E R I M E N T S

W eiss received his doctoral de gre e from ihe

University of Vien na, snpervised by P rzibram

and Hatschek, for an experimental study on

the normal positions of butterflies in re-

sponse to light and gravity, which greatly in-

fluenced his system conceptions. His experi-

me nts were designed to determ ine if and how

m uch the norm al position of b iiiterflies is re-

lated to the direction luid intensity of light

and gravity. He therefore used one or two

light sources and changed the angle ofa wall

on which the animals rested from vertical to

overhanging horizontal. This experimental

app roa ch t


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D EC I K M B ER 2007

\\JSS

AND BERTALANFFi S CONCEPTUAL CONNECTION

7

final position (Weiss 1922:S;

1925:223).

When

discussing the sj'mmetrical position in rela-

tion to two light sotirces, Weiss claimed that,

after disturbance, an equilibrium state is

again 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 lo a state of uniqtie definileness" (Weiss

1922:20, our emphasis).

NE W

APPROACHES

IN DF.VEt.OPMENTAt. BIOLOGY

With his early work in developmental biol-

ogy, Weiss focused on reg ene ratio n and trans-

plantation in vertebrates. Those studies

proved certain prevailing concepts and dis-

cussions to be inaccurate (e.g., the debate on

preformism versus epigenesis). Weiss also

tried to overcome the debate on mosaic ver-

sus regulative d evelop m ent, which was also in-

fltienced by Hans Spemann's "organizer ef-

fect." He argtied that one cannot distinguish

between them based on a .strict timeline (stich

as the eight cell stage); rather, every devel-

oping orgatiism at some time incltides cells

that are determined and cannot develop in

other ways, Weiss's studies in developmental

biology also tise the system concept. He

clearly states that ontogenesis cannot be un-

derstood by examining single isolated aspects

only. Again, be refers to

kstalt

tlieory and

uses it to describe the occtirrences found in

the organism (Weiss 1926).

The developmental biologist Weiss notes

that, in the organism, spatial heterogeneity is

accompanied by substantial heterogeneity;

thus,

spatial form must not he viewed as sepa-

rated from the variations in the involved ma-

terials or sub.stances. With the mo rphological

structure, certain functions are determined

and everything that app ears as distinct differ-

entiation was, in a form er phase , a latent abil-

ity. Biological form is defined as "tlie typical

localization, i.e,, arrangement and allocation

of different sub-processes within the respec-

tive material systems." Accordingly, already

the fact "tliat at this location tliese and only

bors the concept of inseparable systems with

an argument derived from the dynamic pro-

cesses involved:

If in the developmental process

[Gestalt

ungsprozess]

each part would simply be

connected to its neigbboring parts with-

out taking a specific subordination

within the whole system, theti the small-

est de\iation of a partial process at the

beginning would at the end lead to mas-

sive discrepancies. This contradicts the

observation (Weiss 1926;7),

Weiss's criticism of misleading terms re-

flects his oppo sition to the co ncep ts on w hich

tliey are based. Terms like "eqtiipotent" or

"omnipotent" seem improper to him because

the "material parts " of the early germ have no

ability or potential to build up the whole in

an organizational perspective. Rather, the

parts should be called "nulUpotent" because

the specificity of their development is deter-

mined by their position, and differentiation

is dom inate d by the factors prevailing at a cer-

tain position (Weiss 1926:10-11.

V^).

ln this

context, he derived a field theory for devel-

opmental biology comprising five aspects

(Weiss 1926; Weiss 1928;1570; listed as "field

laws"

in Bettalanffy- 1932:328):

1.

Every field has axial structure, which

leads to hetero-polarity

[Heteropolaritdl]

in at least one axis.

2.

W hen m aterial is sepa rated from a field-

bear ing system, tlie field is con taine d in

the remainder in its typical structure.

3.

Wlien 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 two

fields are bro ught together they can add

up or result in a mixed field depending

on their orientation.

5.

A field has the tendency to take up and

incorporate eqtiivalent fields from its

surrotmdings.


10/26

358

THE QU RTEIUJ REVIEW OF BIOLOGY

VOLUME 82

to the use of this term in physics. A whole

responds to influences from the outside as a

whole by inner reactions of the components

in such a way that the former state i.s re-

gained. The special cases in developing or-

ganisms are mo rphogenetic organization

fi,elds. T he field conc ept is considei ed to be

completely in line wilh Gestallpsychology

(Weiss 1926:23). During the developmental

processes in an organism, various subHelds

can come into existence that govern certain

areas of the material parts withiti the whole,

subsequently leading to an autonom isation

of those enuties (Weiss 1926:31).

Note that Weiss was not the first to talk

abou t fields in developm cnuil biology. Boveri

(1910) already postulated a concept of

morphogetietic fields (see Gilbert ct al.

1990:359), and in the same year, Gurwitsch

started 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 inuodticed tlie organizer

con cep t, also used tlie term field. However.

Weiss is considered to be the originator of

field the ory in developm entiti biology (Har-

away 1976:177)he provided a theoretical

basis for the co ncep t (Gilbert etal.1996:359).

FAR-REACHING CONCLUSIONS

Weiss's results from his butterfly experi-

ments and further theoretical conclusions

from his thesis were first published in Ger-

man. The itrticle was already written in 1922

(Weiss 1925:248), tliat is, before he cam e into

con tact with BertalanfTythey very likely met

each other in 1924 for the first time. At any

rate,

Bertalanfly regarded this work as an im-

portant contribution. This is reflected in the

fact that an English translation was published

in the

General

Syslefris

Yearbook

the book ser ies

coedi ted by Bertalanfi)' (Weiss 1959). Weiss

explained:

That 8()-pagc article stated tlie basic

premises and principles of a holistic sys-

a singled-out fraction of na tur e can right-

fully be acc orded the designation of sys-

tem (Weiss 1977:18).

Weiss conc ludes that his results proved to-

tally incompatible with the mechanistic doc-

trinesor rather, rationalizationsof animal

behavior prevailing at that time (Weiss

1977:17). With such doctrines, he refers es-

pecially to Loeb's tropism theory, by which

the behavior of an animal is reduced to a

mere physicochemical reaction triggered by

factors exclusively coming from outside the

organism, and where the mechanisms thai

govern orientation towaid light and gravity

are tlie same in plants and animals. Weiss

(1925:171, 177-178) rejects this fatal mis-

take,

which consists of identifying similar

phenomena

[Erscheinungsformev ]

as the resu

of similar mechanisms. According to Weiss,

the reduction of biological problems to mere

physical and chemical aspects fails to touch

the core of the problem of life.

Besides animal behavior, a field in which he

pe rfo rm ed only a few studies. Weiss is also

similarly critical of developmental research,

where his main interest lies, and to which

Loeb also made conuibutlons (Weiss 1922).

A CONCEPTUAL APPARATUS

Weiss points out the importance of theo-

retical considerations, but he uncritically

holds that the description of facts does not

change, whereas theories, which are formu-

lated to integrate the findings into a larger

context, may change (Weiss 1925:168-169).

In conOiLst to th e reductio nist attit tide ,

Weiss states that it is more promising and ec-

onomical to first determine the laws in a cer-

tain research field such as biology, before

.separating it into physics and cliemi.stry like

Loeb d id. Accordingly, it is impo rtan t th at bi-

ology develops its own scientific conceptual

apparatus [BegriffsbiMurig] (Weiss 1922:2, 8

The basic terms of a conceptual apparatus

from Weiss's perspective are listed below.

Even his early work contains the basis of a


11/26

Dt-XEMBER 2007

WEISS AND BERTALANFFY S CONCEPTUAL CONNECTION

359

ical reaction, a single mtiscle fiber contrac-

tion, a con

ti

action ofthe muscle, a movem ent

of the organ, or a move by the whole organ-

ism), new features appear that cannot be ex-

plained simply by constituents of lower levels

alone, By splitting the whole into lower-level

elements through quaniitative analysis, the

fealiires of the whole may become lost even

methodologically. This would also be tru e for

inorga nic entities such as mo lecules or atoms.

Although a quantitative approach should be

applied, it can never lead to a complete ac-

count of

a

com plex entity, because when such

an entity is fragmented, some characteristics

are lost. Highe r com plexes can be viewed as

units for which certain laws can be found

without reducing them to their components

(Weiss 1925:173-174).

Another possible term in a conceptual ap-

par atus is the system. In his chap ter on laws

of systems, Wei.ss prop oses an early system

def inition : As a .system we want to de fine

each complex tliat, when parts of it are mod-

ified, displays an effort to stay constant with

regard to its ou tsid e (Weiss 1925:181, 183),

In other words, the state of a system should

be stable within and distinct from tlie outside

con ditions . Th erefo re, Weiss defines a system

by its behavioral feature of reac tion. Th is no-

tion wa.s no t new and can be fo und in Her-

ing, as mentioned previously; in Gustav

Fech ners's principle of tendency towards

stability, acc ord ing to which every natu ral

system, 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 tlie parameters

leads the others to modify' in a direction that

compensates this change and enables the sys-

tem to ret urn to an equ ilibrium siate. As

Weiss interprets it, a constiint system state in

the whole, when parts are altered, could only

be 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-

Weiss distinguishes thr ee different system

reactio ns. [n the first on e, an outside factor

applied to the system results in a reaction of

a single pa rt, such as in sim ple reflexes. In the

second, more than one part of the system is

affected by an outside factor. And in the final

reaction, an outside factor does not trigger a

reaction of single parts of the sysiem, but of

the whole system, where the reaction of the

whole is not a result of the reactions of the

paris: it is the other way around, and tlie re-

actions of the parts are a result of the overall

reaction (Weiss 1925:175-181). T he third re-

actionwhich he illustrates by examples

from physics and Kohler's

fstalt

theoryis

important for his argument that an organism

is not merely a conglomerate of cells, mole-

cules,

or atoms. This notion also allows laws

to be found without knowing all ofthe details

of the involved elementary processes, al-

though the latter

a

e im por tant as well. Weiss

emphasizes that the third case has no rigid

mechani.sni. Rather, th e parts show som e plas-

ticity, while the reaction ofthe whole is lawful.

The physicist's descriptions of processes

based on parts and their relatio ns is very

much in line with that argument (Weiss

1925:186-187).

With regard to dynamic processes and Ber-

talanfl\''s concep t of open systems in flux

equilibrium

[Fliefiglnchgeiincht],

which

was

in -

fluenced by several authors, it is interesting

that Weisswhen discussing system reac-

tionspicks up Hering 's schema of metab-

olism. Equal rates of assimilation and dissim-

ilation yield a metabolic equilibrium. The

same idea is emphasized in 1920 by August

Piitter, another organismic thinker, in hisvol-

imie on growth equations, which had a sig-

nificant influence on Bertal an fly's own tlie-

ory of gi'owth; the end of growth corresponds

to this equilibrium (see Pouvreau 2005b). For

example, if dissimilation is accelerated by an

outs ide ch an ge , then following the laws of

chem ical kinetics which Weiss calls .system

laws of chemistry assimilation is also accel-

erated (Weiss 1925:193-194). He argues that

if this concept is viewed beyond the chemical


12/26

6


VOLUME 82

the one hand and chance on the other, but

he is not satisfied with either concept and

claims that something such as an arch con-

si.sting of single bricks can never come into

being. A mere mechanistic "catisal analysis,"

however, cannot solve such a problem either.

The solution of rhe contradicting perspec-

tives is seen in tlie historical development of

variotis processes wher e, from the very begin-

ning, the p arts are subord inated to the whole

(Weiss 1925:190-191). The problem of rec-

onciling the system perspective and the his-

torical perspective was also mentioned by

Jean Piaget when referring to the central

problem of every structuralism; "have tlie to-

talities through composition always been

composed, but how or by whom, or have they

first been (and are they still?) on the way of

com position ?" (Piaget 1968:10). T he im por-

tance of this problem is also emphasized by

Leo Apostel (1970:167): Ac cording to h im,

structuralism is so domina ted by the notion

of equilibrium that it does not succeed to in-

tegrate gene.sis and history; the problem is

central both to structuralism and system the-

ory.

EPISTF.MOI.OGTCAI. CONSIDERATIONS

Weiss is deeply influenced by Mach; for

example, he refers to the antireductionist

program of the positivist thinker (Hofer

2000:151). Although Weiss contradicts tbe

idea that biological activities can be explained

by pbysics and chemistry alone, without fur-

ther requirements, he leaves open the ques-

tion whethei or not physical or chemical

terms could replace biological ones In the fu-

ture.

He does claim that the laws of complex

is.sues will never be fully rtfplaccd (Weiss

1925:170).

With regard to prevailing positions of re-

ducing biolog)', Weiss suggests tbat, notwith-

standing progress within biophysics and bio

cbemistry. the latter are only con cern ed w ith

parts.

The laws concerning the relationships

am on g al parts are still the realm of biologi-

cal research. But they must not remain mere

that Weiss (1926:43) provides. The funda-

men tal distinction between "rules" and "laws"

bad already been emp hasized by Roux a nd

Prizbram in biology, but also by physicists

such as W alter Nern st (Przibram

1923;

Nernst

1922);

and it was a recurrent theme in sub-

sequent reflections on theoretical biology,

particularly by Max Hartmann (1925) and

Bertalanfiy (1928). It

s

an oth er expression of

the Kantian distinction between "natural sci-

ence"

as "a whole of knowledge ordered by

princ iples" a prio ri, of which "the certiiinty is

apod ictic," and "natural history" as a "descrip-

tion of nature," of which the certainty is only

empirical, and that is knowledge only in an

inappropriate sense (Kant 1786).

We agree with Khittel when he states tliat

Weiss's epistemological background is rather

.simple, and that he uncritically combines die-

ory and ex per im ent (Khittel 2000:160). Weiss

is satisfied with posing questions to nature

that are answered by the experiment and

does not go deeper into cpistemology.

The experimental biologist Weiss never de-

veloped a "system theory." Rather, from the

insight he obtained by his experiments and

observations, he described what such a theory

must comprise and where the problems lie

with the "analytico-stimmative" approach, as

well as its abstraction and shortcomings. This

sheds light, on Weiss's statement that he only

had "marginal ccmtacts with the field [of

general systems theory]" in his later life

(Weiss 1977:18). Weiss was only minimally

concerned wiih systems beyond tbe single

organism, btit he does consider them occa-

sionallyfor example, at the Alpbach sym-

posium (Koesder an d Sm yihies 1969); he also

refers to communities and societies when

talking about systems (Weiss 1970b:9).Oneof

his last descriptions of systems broadens the

scope even more: "a system is an evipimnlen-

lily which in ou rexperiencehm sufficiently durable

identity to be defined on a m aaoscak as a

whob>

although from ourknou U dgeof the parts alone into

which we can physically or mentally fractionate tfie

entity on the miaoscale,

xve

could nei'er

retrieve

the


13/26

DECEMBER

2007

WEISS

A ND

BERTAIANfTY S GONGEFIV L CONNECTION

361

the following quotation demonstrates, Weiss

advocated, in hislate writings, system think-

ing onabroad and deep basis:

It is an u rge nt task for the future to raise

man's sights, his thinking and his acting,

from hispreoccup ation with segregated

things, phenomena, and processes, to

greater familiarity and conce rn with

their natural connectedne ss, to the total

context. To endow the epistemological

foundations forsuch a turn ofoutlook

with the crede ntials ol validation

by

mod-

ern scientific experience, is tliusamajor

step toward that goal (Weiss 1977:19).

LuDWiG

VON

BERTALANFFY'S

ORGANISMK: CONCEPTS

Ludwig von Bertalanffy, borntoa family of

lower nobility, was (from

an

early age) only

interes ted in the works ofJean-Bap tiste de La-

marck, Charles Darwin, Karl Marx,

and Os-

wald Spengle r (Ho fer 1996; PouvTeau 2006).

In his youtli, Berialanffy lived on his mother's

estate outside ofV ienna next to Paul Kam-

merer, oneofthe most prom inent Prater Vi-

varium expe rime nters. Being a neighbor

and friend of the family, Kam mere r often

talkedto the young Ludwig about biological

matters. Thtis, Bertalajifly, even before his

stud ent years, was probably well awareofthe

specific style in which resea rch

was

conducted

at the Biologische Versuchsanstalt laborato-

ries.

At the end of World WarI,the Bertalanffy

family temporarily left Vienna and movedto

Zell am See (Salzbtirg, Austria) because of

fi-

nancial losses. Ludwig started histiniver.sity

studies, focusing on history of ait and philos-

ophy (especially metaphysics, logics, and phi-

losophy of religion), in Inns bruc k (Tyrol,

Austria). To a lesser extent, he was also

trained therein botany by tlie experimental

morphologist Emi Heinricher and plant

physiologists Adolf Speriich

and

Adolf Wag-

nerthe latter heing oneof the main advo-

FIGURE

4. LuDWtGVON BERTALANFFY' (igoi-1972)

Osierreichische Nalionalbiblioihek/Wien

NB

553.771-B.

(in 1925)can belabeledas metaphysicsof

life''anddealt especially with Henri Bergson's

thinking. Indeed, the sttident Bertatanfty'was

essentially what one might call a well-in-

formed autodidactforbiological matters.He

ea rne d his PhD in 1926 with a dissertaticm on

Gustav Fechner's metaphysics of organization

levels,

which already stressed

and

discussed

some contemporary ontological controver-

sies

in

biology. Between 1926

and

1932,

he

devoted most

of

his studies

to

philosophy of

biology, justifying and developing

a

project of

theoretical biology. Hereceivedhis Habili-

tationin1934, which wasin theoretical biol-

ogy. He livedinVienna until 1937, develop-

ing both his theore tical perspectivesand

some experimental applicationsin thefield

of animal organic growth.

A

Rockefeller

scholarship was granted tohimforon e year.


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6

THE QlJ RTERiy

REVIEW OF IOLOGY

VOLUME 82

in March 1938, Bertalanffy- tried to find a way

to stay in ttie United States. But conditions

were unfavorable in that regard, with priority

being given to persecuted scientists: he was

forced to return to Austria in October 1938.

The dark period of Bertalanffy's life then

started. Despite liis deep disagreements with

the essential parts of Nazi ideological com-

m itme nts, he becam e a m em ber of" the Nazi

Party in N ovem ber 1938, It was a m ean s to

facilitate his promotion as a profe.ssor at the

University of Vienna, which he was awarded

in September 1940. During the war, in par-

allel to the elaboratjon of his tlieory of or-

ganic growth, he closely linked his "organis-

mic"philosophy of biology* to the totalitarian

ideology in general and the

Fiihrerprinzip

in

particular. Bertalanff>- lived in Vienna until

1948,when he decided to leave pe rmanently.

H e was imp atient regarding his career oppor-

tiuiities in post-1945 Austria becatise of un-

favorable outcomes of his personal "denazi-

fication" procedure and because of the

enmities induc ed by his opportunistic behav-

ior during the war (Hofer 1996; Pouvreati

2006),

He then went throu gh Sw itzerland an d

England to Canada,

Bertalanfly was influenced by many peo ple

(see Potivreati and Drack 2007). From the

philosopher's side, it was by no means only

the Viennese Circle but, among others, also

incltided neo-Kantian thinkers such as Hans

Vaihinger and Ernst Cassirer. His organismic

biology was largely influen ced by JuHtis

Schaxel, Emi Ungerer, Nicolai Hartmann,

Hans Spemann, and Friedrich Alverdes, but

to some extent also by the "Prater Vivarium"

and Paul Weiss,

BERTALANFFY MEETS WEISS

When still a student, Bertalanffy met the

recently graduated Weiss and tbey discussed

system issues in biology, Weiss with an exper-

imental and Bertalanffy with a philosophical

background. Five years after BertalanfTy's

death, the 79-year old Weiss remembers:

soon found that his thinking and mine

moved on the same wave-lengthhis

coming from philosophical speculation,

mine from logical evaluation of practical

experience. And so it remained for half

a century, each of

us

hewing his separate

patli according to his predilection. That

is,

kept on as the empirical experimen-

tal explorer, interpreter, and integrator,

for whom the "system" concept re-

mained simply a silent intellecttial ginde

and helper in the conceptual ordering of

experience, while he, more given to ex-

trapolations and broad generalizations,

and bent on encompassing the cosmos

of litiman knowledge, made the theory

[genera l system theory ] itself and the ap-

plicability of it to many areas of human

afiairs his prime concern. Does not this

confluence here, once again, prove the

"hybrid vigor" of the merger of ideas

that, coming from a common source,

have converged tipon common ground,

alheit by .separate routesthe one offer-

ing a distillate of a life of study of living

systems, the other the extensive elabo-

ration of an intuitive philosophical ide-

ology, tested in its pertinence to human

evolution? (Weiss 1977:18-19),

During the last three months of Berta-

lanffy's initial

U.S.

visit (July dir ou gh Septem-

ber 1938), he worked with Weiss at Woods

Hole (Hammond

2003;

Pouvieau 2006), This

was dieir last meeting until 1952, In autumn

of that year, they met again at the University

of Austin (Texas), where Weiss was working.

At that time,Bertalanff\ was on a tour, mak-

ing abou t 15 conference presentations

Ln

dif-

ferent parts of the cou ntry; he cam e to Austin

at the invitation of the biochemist Chauncey

D Leake. Weiss probably played a role in this

invitation (Pouvreau 2006:76). Years after go-

ing abroad, Bertalanfi)' and Weiss met again

in Austiia at the small alpine village of Alp-

bach , Tyrol, on the occasion of a special sym-

posium devoted to ,systems tiiinking, orga-


15/26

DECEMBER

2007

WEISS AM) B ERTALANFFY'S GONCEPTUAL CONNECTION

Bertalanffy after 1945. As for th e per iod b e-

fore 1945, no letter s to or from Bertalanffy

are cons erved in this estate, because his hom e

was bu rne d at the en d of World War II.

EARLY WORKS OF BERTALANFFY

In the course of his studies, Bertalanffy fo-

cused on philosophy, having Robert Reinin-

gerthe leader of neo-Kantianism in Austria

and president of the Viennese Philosophical

Society (1922-1938)and Moritz Schlick

among his teachers. Bertalanfly recognized

his student days in Vienna as a time of high

intellectual intensity, triggering many devel-

opments that later became important for him

(BertalanfTy 1967). Concerning the positivist

ideas of the Vienna Circle aro un d Schlick,

Bertalanffy logically rega rds emp iricistn as an

erroneotis doctrine. He holds that empirical

data have to be orde red by laws, because one

cannot base and develop any science solely

on exp erien ce and ind uctio n (BertalanfEy

1927a:660). Referring to Kant (1787:A51/

B75),

he liked to say that, thoug h theory widi-

out experience is mere intellectual play, ex-

perie nce w ithout theo ry is blind (Bertalanffy

2003:101). Fu rth er m ore , when Bertalanffy

outlines the difference between science and

metaphysics, mythical thinking, and mysti-

cism, he argues that a dialogue between them

is neces.sary. For exam ple, h e writes: T h e

most noble task of science is that mysticism

does not bring us back to the night of igno-

rance and faith, but that it becomes itself a

means for the deepest understanding of na-

ture

(Bertalanffy 1927b:264). Convergences

with positirism can nonetheless also be

found: especially a taste for logical analysis of

concepts, the rejection of substantialism (see

Mach 1906 and 1933), and the will to save the

dignity of science in tbe face of the rise of

irrationalisma characteristic trend in the

interwar German and Atistrian contexL

Bertalanffy's doctoral thesis (Bertalanffy

1926a) was concerned with intrigtiing ques-

tions:

Can tmits of higher orde r (integrating,

concept of organization, as opposed to spec-

ulative metaphysics? His dissertation already

covers tlie stratified levels of biology, psychol-

ogy, and sociology, and he emphasizes the

perpe tual r ecu rre nce of the same in all levels

of in tegra t ion [hUegralionsstuJeri] (Bertar

IanfTy- 1926a:49). To illus trate this, he employs

the model of the atom as a small planetary

system, reviving the ancient doctrine of the

homology of macrocosmos and microcosmos.

No te, however, that this mo del was already re-

garded as erroneous at that time in physics,

after having been understood as a useful fic-

tion.

After giadtiating, he wrote many articles in

the field of theoretical biology. In 1928, he

published his critical theory of morph ogen -

esis

(Bertaianffy 1928), which acq uain ted

him wilh the Berlin Society for Scientific

Philosophy and Hans Reichenbacha coun-

terpart to Lhe Vienna Circlean d op en ed

contacts with

Gestall

psychologists (Ho fer

2000 :154). In that work, Bertalanffy regards

tlie development of an organism out of a

germ as a paradigm

\Paradigma]

illustrating

the nee d for an organism ic a nd theoreti-

cal

biology. He therefore analyzes the vari-

ous theories of development

[FormbilduJig],

among them the approaches of Przibram and

Weiss (Bertalanffy 1928). Prz ibram 's analogy

of a germ and a growing crystal is criticized

becatise the first one consists of inhomoge-

neous parts, while the second is homoge-

neous,

althou gh b oth have a vectorial pote n-

tial

and tend to regenera te if disturbed. Also

criticized is the lack of differentiation be-

tween life and the inorganic realm (Berta-

lanffy 1928:1 66-175). The field the ory in

Weiss's developmental biology is categorized

by Bertalanffy (1928:18 9-190) as an organ is-

mic theory (.see below). Bertalanffy con-

cludes Irom the analyzed theories of devel-

opm ent that wholeness

[Garizhdl],

G estalt, is

the prim ary attrib ute of life (BertalanOy'

1928:225). He uses who leness

[ anzkeil]

and

estalt dn

synonyms (Bertalanfly' I929c:89).

In May 1934, Ludwig von Bertalanff)' was


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6

THE QUARTERLY REVIEW OF BIOLOGY VOLUME 82

been published two years earlier

1932).

THEORETICAL BIOLOGY AND

EPISTEMOLOGV

At a young age, Bertalanffy was probably

introduced to the ongoing controversies in

biolog). Th e neo-Lamarckian Kam merer has

already lieen mentioned above. Kammerer's

impact is clear in Bertalanffy's early criticism

of selectionist or mutationist theories of evo-

lutio n (Bertalanff 1929a). But very early on ,

he distanced himself fiom the different kinds

of neo-Lamarckism that were flourishing in

the first two decades of the 20th century as a

conse quen ce of the inner w eaknesses of Dar-

winism (related in particular to its apparent

contradictions with the Mendelian theory of

heredity). He therefore criticized Kammerer

(Bertalanffy 1928:95) no t only lor his La-

inarckist spec itlations, but also for his al-

leged theory free emp iricism. As a conse-

quence of his early contact with the

(psycho-)\ntalist botanist Adolf Wagner, Ber-

talanffy very likely had firsthand knowledge

regarding the criticism of the Darwinian

school and abo ut attacks on biological me-

chanicism (Reiter 1999; Hofer 2000; Pou-

vreau 200(i). Like Weiss, he was also very

much influenced by Kohler, whom he re-

gard ed as a precu rsor of his general sys-

tem olog \ (Bertalanffy- 1945:5; 1949b:115).

Kohlcr, who wanted to extend Gestalt theory

toward biological regulation, coined the term

Systemlehre

(Kohler 1927), which BertaianfJly

also used an d that we transla te as systemol-

ogy (Pouvreau and Drack 2007). Kohler's

aim was to show that self-regxilation by means

of dynamic interactions is a common ability

in both physical and organic systems.

An impo rtant figure regarding the need for

a theoretical basis for biology was Schaxel,

who in 1919 started a book series Abhandlun-

gen zur

thearetischen

Biolo^e;

Treaties

in Theoreti-

ca l

Biology

dedicated to implementing theti-

retical biology. It is not by chance that works

the term organi.smisch (orga nism ic), which

was coined by the entomologist Ludwig

Rh um bler in 1906 (Schaxel 1919:203). This

term 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 g enesis of

his ide;is and his associated vocabulary.

Spem ann's works (1924) on organizers of

deve lopm ent also influenced Bertalanff)'.

Th e depe nden cy on th e positions of the parts

can only be tinderstood when considering the

organism as a whole, including processes of

differentiation and hierarchization.

Bertalanffy's first book, in which he em-

phasized

the

need for a theoretical biolog)',

appeared in 1928. His motivadon for going

into biolog)' is much related to the fact tliat

biological categories (especially wholene.ss

and organization) were very often and un-

critically used in areas beyond biology, and

often served as a basis for dogmatic ideolo-

gies.

The science of life has nowadays to a cer-

tain extent become a crossroad, in which

the contemporary intellectual develop-

me nts converge. Th e biological theories

have acquired a tiemendous ideological

[ioeltanschauliche\,

yes even p ubl ic an d

political significance. . . . The condition

of biology, problem atic in m any respects,

has led to the situation that the philos-

oph ies (if life were until now by no

means satisfactory from the scientific as

mu ch as the practical po int of view; we

see all the more clearly the importance

ofthe theoretical clarification of biology

(Bertalanffy 1930:4-5).

Ano ther consideration de ma nding a sound

epistemologicat basis in biology was its mere

descriptive or experimental character, which,

acc ordin g to Bertalanffy, ha d to be overco m e.

He argues in the same line as Weiss in so far

as both think that biologv' has to be autono-

mous,

and that life cannot be reduced to the

realm of physics and ch emistry alon e, as lne-


17/26

DECEMBER

2007

WEISS AND BERTALANFFY S CONCEP TUAL CONNEC TION 365

broad sense) or organismic, teieological, and

historical (e.g., Bertalanffy 1928:88). The an-

aiytico-stimm ative app roac h expressed by

the physical and chemical investigation of

iso-

lated processes leaves aside organization,

which is a core problem of biology. Second,

Bertalanffy regarded tbe conflict between

mechan icism an d 'Vitalism as an essentially

metaphysical one, which cannot be solved

empiricallythis can also be seen as a criti-

cism ot empiricism. A new approach was

ther efo re necessary. Bertalantly' called it or-

ganism ic biology (Bertalanfiy' 1932).

In starting his theoretical program, he was

conv inced that biology's right to exist as a sci-

ence requires a theoretical background that

had not been established yet, altliough many

so-called theorie s were aro un d. For him, bi-

ology of tbis time was little more than some

kind of natural history in a precritical ( pre-

Cop ernican note the allusion to Kant)

stage (Bertalanffy 19 28 ;l- 3, 51-5 6, 90- 100 ;

1932:1-9, 20-35). Empirical data have to be

ordered by laws, and BertalanfTy distin-

guished between empirical rules and laws,

a distinction that empiricists usually do not

makeas m ention ed, Weiss did so but no t in

an elaborated wayand that consequently

leads 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 by

opposite facts. On the contrary, the law

is a logical relation between conceptual

constructions; it is therefore deductible

from upper

[libergeordnete]

laws and en-

ables the derivation of lower laws; it has

as such a logical necessity in concor-

dance with its upper premises; it is not a

mere statement of probability, but bas a

compelling, apodictic logical \'alue once

its prcmi.ses are accepted (Bertalanffy

1928:91).

Another confusion involves mixing de-

scription, which is the me re repo rting of

The role of the experiment is overesti-

ma ted by empiricists and, from a logical po int

of view, notjtistified. The law does not belong

to the empirical realm. Furthermore, it can-

not be induced because, in such a proce dur e,

the disturbances in reality have to be known

before the law is derived from em pirical d ata,

which is logically impossible (Bertalantfy

1928:93). Laws therefore have to be derived

hypothetico-deductively, with exp erim ents

being an incentive and instance of proof

(Bertalanffy 1932 :27-28 ). A final mistake of

empiricists is the illusion tha t a hypothesis

free

science could possibly exist. All the ac-

tivities of science con tain a certain theoretical

momentum. Scientists who are unaware of

that instance might easily become victims of

metaphysical or ideological bias.

According to Bertalanffy, biology must also

become ati exact nomotbetic science (Berta-

lanfiy 1928:55). On the on e h and , a theory of

knowledge and methodology of biology is re-

quired. On tbe otber, general principles and

laws of life for deriving par ticula r

l ws

are also

part o fa theore tical biology; within wbicb th e

different fields of biology should also be syn-

thesized and unified in a coherent system.

Tbeoretical biology has to plan experiments

and strive for the conceptual mastery of the

phenomena. Furthermore, the unstructiu^ed

accumulation of empirical data and the spe-

cialization into various fields, which obscu res

the links among the problems, must be over-

com e (Bertalanffy 1932:32-34). Nevertheless,

establishing exact laws does no t m ean that

biology has to be reduc ed, by causal analy-

sis,

to physics and cbemisUy. The essential

and stifBcient ch ara cte r of a law is to link

univocally a definite initial state with a defi-

nite final on e, thereby it is not indispens -

able

for such a det erm ina tion that all inter-

me diary processes are know n (Bertalanffy

1930-1931:400). This is similar to Prizbram's

position but, of course, also to Boltzmann's

statistical approach in thermodynamics to

which Bertalanffy repeatedly refers, or to the

contemporary approaches in quantum me-

chanics.


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TI-IE QUAR TERiy REVIEW OF BIOLOGY

VOLUME 82

ism. His organismic pro gram aims at de-

termining the corresponding laws that are

complementary (in Niels Bohr's sense) to the

analytico-summative approach. Ftirthermore,

tlie concept of teleology is rehabilitated: not

in a \italistic way (in the metaphysical sen se),

but as a logical form of the maintenance of

the whole. The basic assumption that the

world is stratified (N H ar tm an n 1912) is used

in order to connect causality and finality in

connection with the concept of dynamic equi-

librium: Wliat in the whole deno tes a causal

equilibrium process, appears for tlie part as a

teleological event (Bertalanfiy 1929b:390;

1929c:102). His de-anthrop oinorphization

of teleologyit might be less confusing to

employ the word teleonom y instead, which

was introduce d by Pittendrigh (1958)and

use of

.systemic catego ry is in line with o th er

authors such as Ungerer, Schaxel, Weiss, and

Nicolai Hartmann.

Just like the concept of energy is die

form of expression for the causal law rtil-

ing the inorganic world, the concept of

organism is the form ol expression for

the finalist

[finale]

perspective, which we

have to apply to the organ ism besides the

causal one. The characteristic of the or-

ganism is first that it is more than the

sum of

its

parts, and second tliat tlie single

processes are ordered for the ma intenance

of the whole (Bertalanffy 1928:74).

Important in this context is the idea that

an equin nal state of ihe whole may be

reached independently of the initial states of

the parts and in the way they reach it. This

conception is crucial in developmental biol-

ogy and clearly contra dicts a mec hanicist at-

titude.

The epistemolog )' of organism ic bioUjgy

is based on the conception of the organi.sm

as a system, and can thus be called a system

t h e o ry o f

life,

T h e t e rmSystemthe.one[ %\ &iem

theory ] first appears in Bertalanfty's writings

in 1930-1931 (p 391), although he does not

explicitly define what is meant by system be-

statistics, wh ere the causalities driving the

single pa rt are left aside, exact laws are nev-

ertheless derived without redtiction to the

physicist and chemist realm. Clearly, Berta-

lanfly^'s thinkin g and the a ttem pts of Przib ram

converge. With his growth equations, Berta-

lanffy (1934 , 1941) late r .shows how a mathe -

matical approach in that direction can be im-

plemented in the field of the dynamics of

morphogenesis (Pouvreau 2005a.b).

It should also be mentioned that Bertalanff)'

rejccLs emergcntism and holism in their origi-

nal sense (Lloyd M organ 1923; Smuts 1926).

He considers them as belonging to metaphys-

ics and as sterile in sciencehoth implying an

ontological irreducibility of the whole to its

parts and relationships, even when they

i e

ex-

haustively known (radical qualitative

novelty .

He al.so easily resists critiques such as those of

Phillips (1976). ,\lthotigh that latter critique

toward a simple-minded Hegelian attitude oi

holistic thinkersfocuses on the late works of

botli Weiss and Bertalaniiy; it should be dis-

ctxssed here becau.se it also provides insight

into th eir early works. Bertalanffy's position

might be interpr eted as echoin g Georg W F

Hege l's doctrine of intern al relations (ac-

cording to which the whole relational struc-

ture is cons titutive of each part ) as well as

holism, which is not the case. In contrast to

Phillips's critique , B ertalanfly's statem ents a re

ptuely epistemological. Weiss was probably

too pragmatic and loo much an experimen-

talist to incorporate any Hegelian attitude.

But the chief argument brought forward

against Hegelian thinking natnely, against

tbe idea that the parts gain certain properties

when combined to a wholecannot be sus-

tained for the two biologists. Bertalanffy un-

ders tood the whole as the sum of its part.s

andtheir relations, and thou ght tliat this is all

that can be tackled by science. T he only prob -

lem lay in the epistemological difficulties

posed by organized complexity as W arren

Weaver (1948) calls itwhich sets into prac-

tice the impossibility to satisfy the required

condition of knowing all the parts and their

relations in order to know the whole:


19/26

DECEMBER

2007

WEISS AND BERTALANFFY S CONCEPTUAL CONmCTTON

7

the components brought together and

all the relations existing

between

them,

t h e n

the higher levels are derivable from

their components (Bertalanfiy 1932:99;

1949a:140),

Th e system laws thus dea l with the relation-

ships among the parts: the existence of such

laws relies on the idea that the relational

struc ture may cha ng e, while the parts stay in-

trinsically the same (i,e,, as studied in isola-

tion from the whole), so that the whole has

properties of its own. Only in that sense is

em ergen ce tangible by science, and not in the

mentioned metaphysical sense (Bertalanffy

1932;99). A later statement by Weiss is inter-

esting in this context, which is very much in

the same line. In the phr ase the whole is

m ore than the sum of its parts , Weiss inter-

prets more not as an algebraic term , but re-

fers to the m ore inform ation tliat is ne ed ed

to describe the wbole compared to the parts.

With this conception, Weiss bridges the gap

between reductionism and holism (Weiss

1970a:20).

Bertalanffy is also very much in line with

Weiss (see above) wben he states that bio-

logical system laws are not once and forever

irreducible to physics, although Weiss refers

to biological terms that in the futtire might

be replaced, but fie holds that the laws of

complex issues will never be ftilly replaced by

physics and chemistry (Weiss 1925:170).

To the question whether the concepts of

physics are at present sufficient for sci-

entific biology, the answer is no. . . , To

the question whether the [purely biolog-

ically applied] concepts can anyway be

replaced by physical ones, the answer is:

wait and see (Bertalanffy 1932:113-114

following Woodger).

If the system laws of the organ ism can not

today be formulated in a pbysico-chemi-

cal way, Uien a biological version of those

laws sjtistified. T he question w hethe r an

ultimate redtiction of the biological sys-

tem laws is possible , , , is of secondary

(Bertalanffy 1932;331), which shou ld m ore

appropriately be regarded as conceptual

models of biological organization. These

principles had already been previously

mentioned by others, but Bertalanff)' unifies

them and extends their scope from the single

organism to biological organizatiotis in gen-

eral (from cell to biocoenosis), thereby open-

ing the way from organicism to systemism.

Th e first one is the principle of the orga-

nized system as an open system in flux

equilibritmi

[ liepgUnchgewicht]

(Bertalanff)

1929c:87; 1932:8.V84, 116, 197; I940a:521;

1940b:43). This equilibrium cannot be com-

pared to chemical eqtiilibria becatise the latter

are characterized by a minimtmi of free en-

ergy. In contrast, the organism is an open sys-

tem that maintains itself throtigh a continuotis

flux of matter and enei g\'. by assimilation and

dissimilation, distant from true equilibrium,

and is able to .stipply work. Metabolism is thus

an essential property of the organism.

Th e second principle, also first enunci-

ated in 1929, is the striving of the organ ic

Geslalt

for a maximum of formne.ss

[Geslaltel

//^i] (Bertalanff)'1929c:104), This

s

thought

to play a role in ontogenesis as well as in phy-

logenesis, and later becom es the principle of

hiera rchiz ation or principle of progressive

organization (or individua lization) (Berta-

lanff)'

1932:269-274, 300-320). The dynamic

interactions in the system give rise to order.

The principle is very mtich 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 sort

of centralization , with leading parts that

control the development of other stibsystems,

can also occur. Characteristic is the inherent

trend toward an ever-increasing complexity

a trend that he later Bertalanf^- 1949a) calls

anam orpho sis, after Richard Woltereck had

coined the term in 1940.

Besides the principles oi'w hole nes s and

dynamics, a third o ne , namely that of primary


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368


VOLUME 82

also has ancient roots. It must be viewed in

opposition to a mere passive concept of or-

ganisms that merely react to the outside

world. The concepts are not new, bta were

already forwarded by researchers of Lebens-

philosophie ( philosophy of life ), which was

an intellectual fashion in the first two decade s

of the 20th century (Potivreau and Drack

2007). Weiss and Przibram also argue along

the lines of prim ary activity, but Bertalanffy

does not refer to tliem in tliis regard. Berta-

lanfTy largely cites Weiss when discussing the

theory of the morphogenetic fields (e.g., Ber-

talanfly 1928), although he at least once

clearly writes about system theory with regard

to animal behavior without mentioning Weiss

(Bertalanffy 1932:57-5 8). Th er e, he refers to

the

rstarrung

(congealment) into a machine

as a secondary process and the

Fdhighrii zum

Ganztn

(ability toward being a whole) as pri-

mary to the plasticity and regulation abilities

of the most rigid, machine-like reflexes and

instincts of the organisms. But Weiss's theory

of animal behavior is never explicitly men-

tioned. Bertalanfty (1937:136-142) provides

detailed critici.sm of Loeb's theory, but only

Friedrich ,\lverdes is mentioned, not Weiss.

In the 1920s and 1930s, Alverdes

was

the m ain

German advocate of an organismic approach

to animal behavior (individual, but especially

social);

his entire body of work is dedicated

to it.

BEYOND W F : ISS ' . S C O N C E P T I O N S

Note that Bertalanffy's concerns went be-

yond the co nverge nces with Weiss's conce pts.

Th e prog ram of his organismic biology

comprises fotir dimensions. It is (1) a scien-

tific program for opening the

way

toward c on-

structing exact theories of biological systems

of different levels. It also is (2) a na tura l ph i-

losophy of the biological world, where the two

principles m en tion ed above probably have

in the organic realm a similar basic impor-

tance as the physical fundamental laws for in-

orga nic na tur e (Bertalanfly 1932:282). Es-

sendal for this philosophy is the stratified

basis for developing an inductive metaphysics

has to be mentioned. Metaphysics can, ac-

cor din g to Bertalanff) (1932 :283), find a solid

fundam ent in organism ic biology for re-

newing answers to old problems. Finally,

there is (4) an ideological dimension, insofar

as organism ic biology is sup pos ed to lay the

foundations of an alternative worldview be-

yond mechanicism a found ation in which

the

kmzheit

[wholeness] plays a central role

(Pouvreau and Drack 2007).

The fact that Bertiilanffy's organi.smic biol-

ogy was not an em pt)' program

is

exemplified

by his theory of organic growth, which even

today remains a central reference in that field.

In this theory, global growth and relative (al-

iometric) growth are tackled not only by find-

ing growth constants, but also by linking

growth witli anabolism and catabolism. This

step had not been made before, and combines

the open system concept with equlHnality (see

Pouvreau 2(K)5a.b; Pouvreau and Drack 20 07).

As already described above, this is in line with

laws

of higher ord er and mathematical mor-

phology as m ention ed by Przibram.

C O N C E P T U A L C O N N E C T I O N

The fields of interest of Bertalanffy and the

Vivarium biologists ove rlapp ed co iisidei-

ably. The Vivarium tackled some t>f the ba-

sic questions in biology experimentally and

Bertalanffy strived to set up a theo retic al basis

of biology. Unsurprisingly, they argue in simi-

lar fields, such as developmental biology;

which comprises fundamental biological

questions: indeed, observations from embry-

ology also inf luenced Bertalanffy's sy.stem

the ory of life. The research style at the Vi-

varium clearly offered on e, althou gh no t the

only one, introductory approach to issues of

system theory. W hen Weiss and Bertalanfly

met in 1920s Vienna, their discussions very

certainly centered around concepts that later

converg ed in Bertalanfly's system the or y

Bertalanffy, however, was also well a cquain ted

early on with the works of Przibram, and first

mentioned Przibram in his first paper in the


21/26

DECEMBER

2007

WFJSS AND BERTALANFFY S CONCEPTUAL CONNECTION 369

wise,

BertalanfJy would not have mentioned

him at the ,\lpbach symposium as a key figtire

regarditig tlie system approach in biology.

Weiss was one of the first researchers who not

only described but effectively grasped the or-

ganism as a system. The experiments that he

performed support tlie system approach ofthe

organism and , thereby, also Bertalanfly's or-

ganismic biology. Like Bertalanfly, Weiss ad-

dresses system theory later on in his career.

However, the development of both thinkers

was more parallel, and the influence of one

on the other was only minor.

The concepttial connections between

Weiss and BertalanfH- can be summarized as

follows: Botli think dynamically and in hier-

archical term s, an d both consider whole-

ness

as a key conception for understanding

living STStems or organisms as systems, as op-

posed to a mechanicist appr oach . In this re-

gard, tlie principle of primary activity is cru-

cial for bo th. Th e idea of conserva tion is

also fundamenuil to their system concepts.

Weiss's tentative definition of system and for-

tnulation of system laws also has a certa in

generality, which is paralleled in Bertalanffy's

later effort toward a general system theory .

Both also argue for biolog)' as an aut ono m ous

discipline with its own conceptual apparatus,

which cannot be reduced to physics and

chemistry; and for biologists to find laws of

higher order. Nevertheless, Bertalanfly was

much more interested in establishing a theo-

retical biology than Weiss and, therefore, his

epistemological attempt was more sophisti-

cated.

POTE NTIA L CON TR IBUT ION TO FU TURF.

SYSTEM APPROACHES IN BIOLOGY

As the conceptual connection between

Weiss and Bertalanff>' arises from both exper-

imentiU and philosophical considerations,

these can serve as a useful framework for fur-

ther re search reg ard ing biological systems. In

rec en t years, systems biology (see Kitano

2002) and bioconiplexity research (see

Michener et al. 2001) have appeared as new

biology can be described as the study of the

interactions among the components of bio-

logical systems and the cons equ ent function

and behavior of the system as a .system (i.e.,

how a biological whole behaves over time).

Th is already shows tha t systems biology goes

beyond a me re analytico-summative ap-

proach. Still, quite often concepts only from

eng ineer ing or cybernetics are used to model

biological systems; for example, when feed-

back loops are introduced or when robust-

ness is investigated. These concepts are often

derived from physics and do not comprise

central characteristics of life. Woese (2004)

criticizes this engineering approach, which

he already observes in the Human Genome

Projec t, a pivotal kickoff impulse for systems

biology. Som e scientists aim to expla in life

within systems biology on the basis of mo -

lecular interactions with emphasis on genes

and proteins. Furthermore, the question of

how the phenotyp e is gene rated from the ge-

notype is im por tant w ithin systems biology,

as is the correspondence between molecules

and physiology at a larger scale.

Kirschner (2005) notes that the perfor-

mance of

a

protein is not only dete rm ined by

a gene, but that it depends on the context in

which it occurs, and this context is history de-

pendent The historical dimension of all liv-

ing systems can be detected in the work of

Weiss and Bertalanffy as well.

The concept of hierarchy was already cen-

tral to Weiss and Bertalanffy. Thus, they were

far removed from what Mesarovic and Srecn-

ath (2006) criticized as a flat ear th pe rspe c-

tive in systems biology.

In addition to the concept of control, as

stemming from engineering feedback sys-

tems,

the importance of coordination in liv-

ing systems has recently been pointed out by

Mesarovic and Sree nath (2006:34): Coordi-

nation . . . provides motivation/incentives to

the parts to function in such ways that they

contribute to the overall system's goal while

at the same time they pursue their local ob-

jectives.

This parallels the concepts of


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7

THE QUARTERI.Y REVIEW

OF

BIOLOGY

VOLUME 82

prevail in "systems biology." Here, the con-

cepts of Weiss and Bertalanfly- might help

avoid the risk of "mechanicism." As shown,

some of these c onc epts have iUready been in-

corporated or encouraged to be integrated in

"systems biology." Prim ary actixit); however, is

not an issue in "systems biolog\" today, but

wa.s acknowledged as important by Weiss and

Bertalanffy.

Although skeptical about earlier system at-

tempts in biolog), O'Malley and Dupre

(2005) emphasize that furt

Documents

ON THE MAKING OF A SYSTEM THEORY OF LIFE: PAUL A WEISS AND LUDWIG VON BERTALANFFVS GONCEPTUAL GONNEGTION