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升级   99.43% TA的每日心情 | 擦汗 2016-1-30 03:42 |
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英文原文:<br/>2.1 The essential idea of Autopoiesis<br/>+ D: C4 l% Q0 k3 q. m! O" `
The fundamental question Maturana and Varela set out to answer is: what" q) ]+ m4 g3 ^0 ?' E$ E G
distinguishes entities or systems that we would call living from other
8 f7 a: @. v, O5 jsystems, apparently equally complex, which we would not? How, for
) b8 n# c0 e9 Z) @1 m( H: |7 Aexample, should a Martian distinguish between a horse and a car? This. }2 S0 ]9 E% i; O4 H
is an example that Monod (1974, p. 19) uses in addressing the similar
( m3 u7 |; R2 z; d2 I' lbut not identical question of distinguishing between natural and/ c. X! o3 z2 Q8 `+ q$ B0 C; F1 M4 _4 g
artificial systems.<br/>
! D( O4 p/ L% \7 \This has always been a problem for biologists, who have developed a8 Q' P9 Y) d( X' ~
variety of answers. First came vitalism (Bergson, 1911; Driesch, 1908),% b7 R' n* p" n8 w, L0 `
which held that there is some substance or force or principle, as yet
2 Y4 y' ^' n0 Q4 _unobserved, which must account for the peculiar characteristics of
- R5 _' q4 B8 Y' klife. Then system theory, with the development of concepts such as
$ C$ A0 e# C$ @7 O) zfeedback, homeostasis, and open systems, paved the way for explanations
% X6 c% O' p3 b. }1 F; \of the complex, goal-seeking behavior of organisms in purely8 b: u+ X7 r* ?" H4 B0 _
mechanistic term ( for example, Cannon, 1939; Priban, 1968). While this
6 W( [6 e, e& L3 G1 U' qwas a significant advance, such mechanisms could equally well be built4 B* p, s6 `) ~! R+ v% {
into simple machines that would never qualify as living organisms.<br/>: c& F" G5 x; R# O( w
A third approach, the most common recently, is to specify a list of
; ]" i9 V2 E- s+ s4 k% S' c3 ~5 pnecessary characteristics that any living organism must have – such as
0 q4 h$ e! E, t2 x& ?. Freproductive ability, information-processing capabilities, carbon-based& c/ s9 c0 j# x7 G' j
chemistry, and nucleic acids (see, for example, Miller, 1978; Bunge,9 ?9 b- p7 O( }3 y) l+ _% @
1979). The first difficulty with this approach is that it is entirely* Z& m% f5 u* S7 L
descriptive and not in any real sense explanatory. It works by3 o6 r6 \5 ^9 G% i7 r2 ~; Z
observing systems that are accepted as living and noting some of their) o E) R) L8 o: R: d$ ?/ W: S
common characteristics. However, this tactic assumes precisely that$ Y7 ^( x: I5 i# u
which is in need of explanation – the distinction between the living" A& [) Q* B: R5 `" F
and the nonliving. The approach fails to define the characteristics. p! {* u. A& n H* A& W9 c+ i
particular to living systems alone or to give any explanation as to how
* o/ a' S* j& {such characteristics might generate the observed phenomena. Second,
& E9 W$ I: L* othere is, inevitably, always a lack of agreement about the contents of% T. c% |! X4 m/ g! \
such lists. Any two lists will contain different characteristics, and
! k7 P4 e: J* Cit is difficult to prove that every feature in a list is really$ t/ m$ ]2 k* i& _' n
necessary or that the list is actually complete.<br/>3 X% H2 Z3 x/ I( }: T
Maturana’s and Varela’s work is based on a number of fundamental
f7 ~5 S2 y% f; I) q/ z7 Zobservations about the nature of living systems. They will be
5 z, ^/ Z% u9 ~7 L- Gintroduced briefly here but discussed in more detail in later chapters.<br/>5 G& [& \4 f3 V
1. Somewhat in opposition to current trends that focus on the species# D- C5 n6 O0 `- b$ u% d
or the genes (Dawkins,1978), Maturana and Varela pick out the single,; o+ j9 V/ S/ p* W t2 _ |
biological individual (for instance, a single celled creature such as, u2 i) G) ?2 V
an amoeba) as the central example of a living system. One essential2 K0 o- l$ ]" {. X: f: o4 e
feature of such living entities is their individual autonomy. Although
* j, b1 C) F+ zthey are part of organisms, populations, and species and are affected
9 Q% I- f3 ]0 b4 m; I3 e6 oby their environment, individuals are bounded, self-defined entities.<br/>
, s1 F% X4 f: {* v3 W2. Living systems operate in an essentially mechanistic way. They8 `) W. ~. C1 D
consist of particular components that have various properties and
* |) g( P/ H( q) w6 \interactions. The overall behavior of the whole is generated purely by
* m# ^ q2 c/ u! `$ z: }7 T# Wthese components and their properties through the interactions of
4 Y7 t* F; u6 @' c$ |neighboring elements. Thus any explanation of living systems must be a
* i# l) g6 ?' g, q' f) Ypurely mechanistic one.<br/>* A# h' R! C$ I% l+ f% P6 L6 l
3. All explanations or descriptions are made by observers (i.e.,
. R! y- y \& {* wpeople) who are external to the system. One must not confuse that which0 u: O2 ^2 f6 W
pertains to the observer with that which pertains to the observed.. w/ a- T f$ w) A( o! s
Observers can perceive both an entity and its environment and see how
; M+ G0 i, R+ y9 k% ~9 X+ {" tthe two relate to each other. Components within an entity, however," j. q$ R8 y" W* J5 \! H- E2 l
cannot do this, but act purely in response to other components.<br/>* \$ e( q1 b# l* R
4. The last two lead to the idea that any explanation of living systems) @5 B/ I9 l: ~/ @2 t
should be nonteleological, i.e., it should not have recourse to ideas0 D# ` i' }& F
of function and purpose. The observable phenomena of living systems
* A7 x" f8 V% r, d2 v( `/ lresult purely from the interactions of neighboring internal components.. r0 i: j0 A1 B8 U. C7 d
The observation that certain parts appear to have a function with* o" x5 \" l$ d; i
regard to the whole can be made only by an observer who can interact" u' l3 |4 D) }& S, u7 G+ P$ Y+ {
with both the component and with the whole and describe the relation of$ K' i) p T9 A5 y' O* L- E9 h! A
the two.<br/> ]. z! z- a. S3 h
<br/>7 X- o/ [* A9 l" }
To explain the nature of living systems, Maturana and Varela focus on a3 p5 X- z" H! P( o' O9 t& i: U0 h0 U" ?
single basic example – the individual, living cell. Briefly, a cell* R/ l5 n6 h' h& ]4 ]
consists of cell membrane or boundary enclosing various structures such
) F% C3 T9 g3 p$ ~) Fas nucleus, mitochondria, and lysosomes as well as many (and often
; G- i9 Z/ ?! S. h# E: ucomplex) molecules produced from within. These structures are in
4 @; i8 j& M8 G! {8 B7 Kconstant chemical interplay both with each other and, in the case of( b& ~6 [$ n6 T; @
the membrane, with their external medium. It is a dynamic, integrated0 f4 ]' f/ x5 G1 Z4 I" I" n% }
chemical network of incredible sophistication (see for example Alberts
" Q! W1 n5 s* z$ y& B5 Tet al.,1989; Raven and Johnson,1991).<br/># W [ t' |9 a: V U% n
What is it that characterizes this as an autonomous, dynamic, living
9 v: e; }* I4 h( Bwhole? What distinguishes it from machine such as a chemical factory: U3 f9 M( \3 i: _
which also consists of complex components and interacting processes of P& _# \# [% Z& }! s
production forming an organized whole? It can not be to do with any
4 I& O/ \6 T4 W Y6 gfunctions or purposes that any single cell might fulfill in a larger) ]+ j3 @8 L* {# Q1 [# {( L6 v
multi-cellular organism since there are single-cellular organisms that
4 s e2 y8 O0 Z) Csurvive by themselves. Nor can it explained in a reductionist way3 p8 D2 b# z: z% q% v
through particular structures or components of the cell such as the {3 ~7 H# n+ S! E( P- S; {% d9 N
nucleus or DNA/RNA. The difference must stem from the way of the parts9 s- ~+ z; ?+ ~7 F0 i3 f
are organized as a whole. To understand Maturana and Varela’s answer, d/ J7 \/ }7 ?( P8 u+ v2 q4 d$ [
we need to look at two related questions – what is it that the cell
% v( l5 U6 x1 I# L s7 Wdoes, that is what is it the cell produces? And what is it that
7 {/ Q8 F+ i5 w" ~* ^7 Q% Yproduces the cell? By this I mean the cell itself rather than the
& |% B5 M! k# d/ Cresults of their reproduction.<br/>
$ D! {7 b1 q) i/ E. O! jWhat does a cell do? This will be looked at in detail in Section 2.35 K3 B8 n' [8 }% G, b
but, in essence, it produces many complex and simple substances which! i% v' s2 N' d' L3 D
remain in the cell (become of the cell membrane) and participate in
& r& i+ T7 i* [# Y7 R: Bthose very same production processes. Some molecules are excreted from
5 y5 N [' O" H# a5 i) [/ Cthe cell, through the membrane, as waste. What is it that produces the: S) G$ T* W# u% O4 {
components of the cell? With the help of some basic chemicals imported$ x: ]+ q& S' C' E+ E5 R& k, Z$ D5 d
from its medium, the cell produces its own constituents. So a cell
! v5 |& `* R9 i4 k5 Xproduces its own components, which are therefore what produces it in a1 H" K" @% f& P
circular, ongoing process (Fig. 2.1)<br/>2 J( ^9 y8 Y/ ~. M! X
It produces, and is produced by, nothing other than itself. This simple
' W3 g7 c' g6 J' uidea is all that is meant by autopoiesis. The word means& f0 o& M, O# v! x% k. r% P3 S
“self-producing” and that is what the cell does: it continually/ w, B. c2 _9 ]9 I) n6 }$ f1 z
produces itself. Living systems are autopoietic – they are organized in
7 B4 O; |0 F: `# u3 Osuch a way that their processes produce the very components necessary* {" ]8 U4 r/ R( R: C: G5 o) P
for the continuance of these processes. Systems which do not produce$ J A) z1 M8 ^$ ?$ p" g
themselves are called allopoietic, meaning “other-producing” – for: z5 H: J" M% O# x% E, h
example, a river or a crystal. Maturana and Varela also refer to6 o. h* ^' k3 a
human-created systems as heteropoietic. An exemple is a chemical
& h- v6 P4 G! n* r- m( E! J6 ~) ?# `factory. Superficially, this is similar to cell, but it produces9 @0 c2 _+ }6 Y& B
chemicals that are used elsewhere, and is itself produced or maintained& r; S% w# ]1 w/ M4 ^' l
by other systems. It is not self-producing.<br/>
- N- r$ t0 Q5 [/ I: XAt first sight this may seem an almost trivial idea, yet further contemplation reviews how significance it is. For example:<br/>! b+ J! d* y$ r/ m6 H' h# n3 Z( r. J
1. Imagine try to build autopoietic machine. Save for energy and some
5 q% ~, N5 a( t6 abasic chemicals, everything within it would itself have to be produced
1 s. s; ~/ _$ a# \2 `by the machine itself. So, there would have to be machines to produce
6 p: K8 t$ V+ [7 Pthe various components. Of course, these machines themselves would have" c+ B n0 o# |7 \/ D2 Y0 }9 h3 n
to be produced, maintained, and repaired by yet more machines, and so4 y: a* B- m% ?% O/ p
on, all within the same single entity. The machine would soon encompass' z5 @* b: Y3 i- e' Y% E
the whole economy.<br/>
5 H$ k5 S9 C) T8 v2. Suppose that you succeed. Then surely what you have created would be
6 X; f+ I/ {" g4 \- Kautonomous and independent. It would have the ability to construct and e: ]4 t4 U0 ~9 |+ l
reconstruct itself, and would, in a very real sense, be no longer
* p) U4 l# m0 q6 A ]+ Acontrolled by us, its creators. Would it not seem appropriate to call
! Y3 S# g* {( R; Bit living?<br/>. Z6 ]( z$ ~. Z4 j" S
3. As life on earth originated from a sea of chemicals, a cell in which
Y b u* A. ^/ y3 W$ U) Da set of chemicals interacted such that the cell created and re-created
: f+ `$ Y; G# @4 z% R, Iits own constituents would generate a stable, self-defined entity with" r% Q Y6 v, _
a vastly enhanced chance of future development. This indeed is the9 i; h; { a6 a7 C$ `, V
basis for current research, to be described in section 2.4.1<br/>
. X) x; d% u3 l0 ]6 P4 q4. What of death? If, for some reason, either internal or external, any
: A5 @0 A3 `: }4 upart of the self-production process breaks down, then there is nothing8 z5 U8 C8 C; z: y$ L
else to produce the necessary components and the whole process falls+ b9 j, N5 C# Y& a$ \& z; m
apart. Autopoiesis is all or nothing – all the processes must be
3 t. m7 T$ c/ k' A/ R) m- aworking, or the systems disintegrates.<br/>0 Q' y \2 N0 v! }1 Y
This, then, is the central idea of autopoiesis: a living system is one8 j* T& H3 S' ?! p' G) w
organized in such a way that all its components and processes jointly
& g; _ e3 k3 r: s2 i/ wproduce those self-producing entity. This concept has nearly been( x3 c- h) c, h% F9 O3 s. P# ?/ j
grasped by other biologists, as the quotation from Rose at the start of" g! C% o0 s; o4 E
this chapter shows. But Maturana and Varela were the first to coin a+ Q8 t+ \- L' [' N" H+ z& j
word for this life-generating mechanism, to set out criteria for it
8 M" p. L& t* u2 e2 U" w(Varela et al., 1974), and to explore its consequences in a rigorous9 X) j) |/ L- y
way.<br/>1 o4 ?4 \5 J# O" U9 ~
Considering the derivation of the word itself, Maturana explains that4 q* ~! ~( D S# {$ A
he had the main idea of a circular, self-referring organization without
( Z0 c g7 u- z4 Y" q- {2 |the term autopoiesis. In fact, biology of cognition, the first major4 e; m# A! K2 U1 S+ {3 M6 Q1 D
exposition of the idea, does not use it. Maturana coined the term in ]1 ~( X0 B. I7 C( b6 @
relation to the distinction between praxis (the path of arms, or
7 h j6 I4 s3 g' ^- _action) and poiesis (the path of letters, or creation). However, it is
" f; v5 B0 b) Q6 V. X6 B+ |6 xinteresting to see how closely Maturana’s usage of auto- and
/ H9 Y7 X2 X; c( p% w8 zallopoiesis is actually foreshadowed by the German phenomenological5 m7 V" P- Q, W/ c
philosopher Martin Heidegger. In the quotation at the start of Chapter
; M) h% t. l4 R# Y+ s' a1, Heidegger uses the term poiesis as a bringing-forth and draws the
- L. B! T% X2 l' ~5 mcontrast between the self-production (heautoi) of nature and the' o4 O' I+ L7 f; J9 j9 U
other-production (alloi) that humans do. Heidegger’s relevance to2 c3 v4 F: Q u
Maturana’s work will be considered further in Section 7.5.2<br/>$ {5 F% i/ f1 [* C: \1 R" C
2.2 Formal Specification of Autopoiesis<br/>
2 H7 h4 y3 ]1 j* j* |Now that I have sketched the idea in general terms, this section will) {5 O& q0 q+ w+ ~
describe in more detail Maturana’s and Varela’s specification and
# j- P# S4 L; H7 J3 E [vocabulary.<br/>
/ B6 r. I6 G0 A5 nWe begin from the observation that all descriptions and explanations! Y/ K) l' w: R5 X4 F
are made by observers who distinguish an entity or phenomenon from the
& k) v+ n6 d( S$ o4 lgeneral background. Such descriptions always depend in part on the
3 a: r- a' H. i( mchoices and processes of the observer and may or may not correspond to- p8 I% X- i- X4 v9 A& `
the actual domain of the observed entity. That which is distinguished q, U5 A9 A( N. c. Q
by an observer, Maturana calls a unity, that is, a whole distinguished
1 `1 J4 w4 ^# L6 E Lfrom a background. In making the distinction, the properties which
# t7 ]% N& o" {7 u0 N- r) j) j) Sspecify the unity as a whole are established by the observer. For2 J3 y7 B) j9 p( X' j
example, in calling something “a car,” certain basic attributes or
4 M% Z' g4 M( X4 P/ v0 J! e: Rdefining features (it is mobile, carries people, is steerable) are
* _* }: c/ F1 y( b Nspecified. An observer may go further and analyze a unity into0 Z1 |$ ?1 P) L8 L7 Q. X
components and their relations. There are different, equally valid,
- P/ g: D& q9 w- x; ~6 x$ S! ]4 S7 Yways in which this can be done. The result will be a description of a
: D" s0 |$ G4 V$ M3 kcomposite unity of components and the organization which combines its' y7 V9 l$ d. N% d( p
components together into a whole.<br/>; ~. q8 {; K7 w7 }' C
Maturana and Varela draw an important distinction between the organization of a unity and its structure:<br/>
3 r3 w; e# N- x- F, m[Organization]refers to the relations between components that define
" d& B# [1 W0 }8 C4 X- p/ Yand specify a system as a composite unity of a particular class, and$ w: Q' h" k+ n6 }# i
determine its properties as such a unity … by specifying a domain in, w* k8 O% ]. { [- M/ N0 ]
which it can interact as an unanalyzable whole endowed with
- l# I- V/ ~( |9 r, P; V1 Pconstitutive properties.<br/>' E9 ^# _; M# ]2 G
[Structure] refers to the actual components and the actual relations
* I2 T I' i' s% T. a# {" X. J% x3 Qthat these must satisfy in their participation in the constitution of a
9 b @1 k: o; H8 I; vgiven composite unity [and] determines the space in which it exists as0 p8 [4 Q1 E1 D3 _0 |. [
a composite unity that can be perturbed through the interactions of its; ?; |+ } [( E0 J" Y
components, but the structure does not determine its properties as a
) w- Z6 ?4 R+ C( P3 }3 A* f1 punity.<br/>
* K4 w5 a+ h( Y+ h1 M1 }1 E% GMaturana (1978, p. 32)<br/>
: X; U, A6 x& x" aThe organization consists of the relations among components and the
1 F$ [! R4 M1 k$ D- A- t6 r( J) Knecessary properties of the components that characterize or define the
& ^1 a2 D# s& A/ m) e2 V5 ~/ X6 cunity in general as belonging to a particular type or class. This
, {) w Y4 @/ D" d$ jdetermines its properties as a whole. At its most simple, we can
1 A" {# j7 k0 N; ?" Oillustrate this distinction with the concept of a square. A square is
s3 ?. F0 o s" u, z6 A0 S: Edefined in terms of the (spatial) relations between components – a
: y" U" Q! v0 \1 y! I) xfigure with four equal sides, connected together at right angles. This L7 N e. {9 n& N# S' d
is its organization. Any particular physically existing square is a9 {9 S# E; e3 B6 e! T
particular structure that embodies these relations. Another example is; M; x7 f. K2 h0 q1 m
a an airplane, which may be defined by describing necessary components$ ~. @0 E8 D7 M0 x s" X/ m/ `
such as wings, engines, controls, brakes, seating, and the relations
; d0 r( ?- @% B1 _# g0 d5 m% ~between them allowing it to fly. If a unity has such an organization,2 \0 q1 @/ X& b/ i
then it may be identified as a plane since this particular organizatio0 A. ]& D3 X; i- l2 k" f/ B$ d% w
would produce the properties we expect in a plane as a whole.
* `+ x: F0 g& Y! B1 w5 MStructure, on the other hand, describes the actual components and3 v0 e- z) o* l
actual relations of a particular real example of any such entity, such* E/ P( ?* J0 I1 f6 z8 y3 a
as the Boeing 757 I board at the airport.<br/>+ y+ A6 j( K5 K R$ K! c
This is a rather unusual use of the term structure (Andrew, 1979).
) w% C9 \; W% xGenerally, in the description of a system, structure is contrasted with' q/ m. f# U5 q2 R7 y* h
process to refer to those parts of the system which change only slowly;
0 z o0 k% o, t6 l0 Pstructure and organization would be almost interchangeable. Here,
( F* x6 ^- @( N/ t" v8 @2 F3 v! E; k1 Uhowever, structure refers to both the static and dynamic elements. The: c* K. d7 X5 ] u! c% J
distinction between structure and organization is between the reality/ a0 x2 C. e& C) ?% G! O8 p2 P
of an actual example and the abstract generality lying behind all such
4 f6 z% o7 d8 }+ x" \. nexamples. This is strongly reminiscent of the philosophy of classic& n" ~2 c ]) Q& L( u5 N1 s! ]
structuralism in which an empirical surface “structure” of events is
# Z3 ?/ \; w jrelated to an unobservable deep structure (“organization”) of basic
# G) \4 @/ J) u/ V. erelationships which generate the surface.<br/>
: x: ~* H: F6 {( \/ S. h5 CAn existing, composite unity, therefore, has both a structure and an
( f9 |6 }# F& l" @" J) [8 q0 Korganization. There are many different structures that can realize the& l2 Z% M$ x% M3 G1 T4 x
same organization, and the structure will have many properties and
( e1 C: B! j$ k/ u2 `relations not specified by the organization and essentially irrelevant, F; V; G/ S9 i8 q" Q1 }2 E
to it – for example, the shape, color, size, and material of a R+ M F; w% s+ M0 G% J; |
particular airplane. Moreover, the structure can change or be changed
+ }0 m* d6 n) fwithout necessarily altering the organization. For example, as the
2 W3 N* P! T: I, Iplane ages, has new parts installed, and gets repainted it still' o7 Q& @, g; @0 M' O7 B
maintains its identity as a plane because its underlying organization" l% L0 f; m! S+ m
has not changed. Some changes, however, will not be compatible with the
0 [" ?/ J% z6 C1 U! S: X+ }maintenance of the organization – for example, a crash which converts
x1 y. \2 G( c) a( j' ^/ Othe plane into a wreck.<br/>" w2 `' c |% N# t
The essential distinction between organization and structure is between
8 r. R1 h# |9 o; ]% W+ Aa whole and its parts. Only the plane as a whole can fly – this is its1 V: u) u+ ^6 z: n. ]: K( N
constitutive property as a unity, its organization. Its parts, however,' O3 \- W T" z1 t8 ~* I& Y! X
can interact in their own domains depending on all their properties,- I! }. ?/ s; T* c4 E; M( g4 _
but they do so only as individual components. Sucking in a bird can
* H% y# F2 P' T' ^2 w2 Xstop an engine; a short circuit can damage the controls. These are
1 E l3 P7 y) @5 @6 s- G5 iperturbations of the structure, which may affect the whole and lead to3 V: Y6 ~0 @8 K5 P) h
a loss of organization or which may be compensable, in which can the' P% d: s; b7 {
plane is still able to fly.<br/> J- V) {; ?1 v- i8 \! N/ U, N
With this background, we can consider Maturana’s and Varela’s0 y9 {5 Z: V+ Z8 d: Q7 u
definition of autopoiesis. A unity is characterized by describing the. j' S6 d+ G6 {' [% N. n9 M; L: C
organization that defines the unity as a member of a particular class
: A3 W W. s# h. ?) wthat is, which can be seen to generate the observed behavior of unities# w4 i5 f% A- \, s& ]. g
of that type. Maturana and Varela see living systems as being- {9 V8 \# ^- e g3 C
essentially characterized as dynamic and autonomous and hold that it is' O. D& Q9 K5 ^0 u; o
their self-production which leads to these qualities. Thus the2 P/ ~9 J1 b3 e3 q, }( O; [
organization of living systems is one of self-production – autopoiesis." l7 r. a1 U* o; s
Such an organization can, of course, be realized in infinitely many; l0 `! }7 w& U) w% E- H3 ~3 z
structures.<br/>0 y D5 Q' h- E
A more explicit definition of an autopoietic system is<br/>
3 A* ^& V* ^# E8 q- ^, BA dynamic system that is defined as a composite unity as a network of productions of components that,<br/>7 Q" L- b8 Y6 t- b
a) through their interactions recursively regenerate the network of productions that produced them, and <br/>. H( c; @8 C/ a! a# j
b) realize this network as a unity in the space in which they exist by
8 g" \( a: u8 _; Y. I U- econstituting and specifying its boundaries as surfaces of cleavage from
$ }& g6 [( z0 { A; nthe background through their preferential interactions within the- }: P% @) p& F
network, is an autopoietic system. Maturana (1980b, p. 29)<br/>6 v$ c" f# P* S3 p
The first part of this quotation details the general idea of a system
i- w1 [' U1 \/ ^+ Zof self-production, while the second specifies that the system must be/ ~$ ~6 k- ^* r9 `9 K
actually realized in an entity that produces its own boundaries. This
% X( D) Z* L3 Y1 y( Clatter point, about producing boundaries, is particularly important3 k) \# p" P- h. H7 o
when one attempts to apply autopoiesis to other domains, such as the% t5 }( X ?' M
social world, and is a recurring point of debate. Notice also that the
4 |9 u0 x, P' n) p# f# W, p9 Qdefinition does not specify that the realization must be a physical/ f1 X b7 [0 B ]8 I1 r4 d. C( f
one, although in the case of a cell it clearly is. This leaves open the
! S: t; X1 G$ P( k2 O* ?, widea of some abstract autopoietic systems such as a set of concepts, a' D1 L. K, s8 ]8 A
cellular automaton, or a process of communication. What might the
7 N/ h' [) d6 b1 l4 ^# k4 U) mboundaries of such a system be? And would we really want to call such a' @# R& \6 B+ g( O/ p
system “living”? Again, this is the subject of much debate – See( a* B3 M, U; \, U0 }% w
section 3.3.2<br/>
- e+ Z. F# c' j. i+ \( W. o! t8 bThis somewhat bare concept is further developed by considering the6 O3 S0 t+ T. n4 D: ?
nature of such an organization. In particular, as an organization it' @& C/ V/ o: l/ R( B: w) a
will involve particular relations among components. These relations, in" U0 N& |6 {) M6 C
the case of a physical system, must be of three types according to" m5 {& X0 q3 `: M
Maturana and Varela (1973): constitution, specification, and order.
$ _3 ?" p& w* J- MRelations of constitution concern the physical topology of the system/ R/ V# l. ^: T0 X
(say, a cell) – its three-dimensional geometry. For example, that it
. v. F( @! G3 `3 fhas a cell membrane, that components are particular distances from each
6 N! d. \2 K' o) \- D9 iother, that they are the required sizes and shapes. Relations of
* x$ \ x0 L: p8 I2 [4 wspecification determine that the components produced by the various. O& E* ~0 F: N5 j6 R8 `
production processes are in fact the specific ones necessary for the% V7 Y0 x- t% u. @
continuation of autopoiesis. Finally, relations of order concern the
/ d7 G2 L! E1 `* e: d* u4 Tdynamics of the processes – for example, that the appropriate amounts9 ^; b Y# }# P
of various molecules are produced at the correct rate and at the
& m7 T n: e$ |correct time. Specific examples of these relations will be given later,
. n! t8 Y& P* Y M* \: r9 Mbut it can be seen that these correspond roughly to specifying the
! s3 M7 i6 e- x) L5 g' N“where”,”what”, and “when” of the complex production processes8 q- z9 K* v0 n
occurring in the cell.<br/>
# f' Z% v1 n. L6 M2 ~It might appear that this description of relations “necessary” for8 w, L' h9 D- M6 Z; b4 K# `
autopoiesis has a functionalist, teleological tone. This is not really: [3 P& r0 z5 ^# I) w
the case, as Maturana and Varela strongly object to such explanations.
9 C1 u3 h" x9 U3 eIt is simply that, if such components and relationships do occur, they: O* `) R: X( j4 A
give rise to electrochemical processes that themselves produce further9 W$ m1 o9 r; A
components and processes of the right types and at the right rates to9 X% |# Y; u$ [8 Y( ]. s
generate an autopoietic system. But there is no necessity to this; it0 z. @% F5 J7 K$ [- Z
is simply a combination that does, or does not, occur, just as a plant+ K1 k6 U6 H9 ~0 f" c( S" o
may, or may not, grow depending on the combination of water, light, and2 g- t1 y2 K3 s: q* b' H, R7 \
nutrients.<br/>! ^4 N, v4 S: z5 V
In an early attempt to make this abstract characterization more5 Z" s* {. u, a* z2 r; e2 x
operational, a computer model of an autopoietic cellular automaton was5 b( Q* p% |3 H1 U. }% q
developed together with a six-point key for identifying an autopoitic
7 _5 t5 q$ m0 ~; ^system (Varela et al., 1974). The key is specified as follows:<br/>. D) P2 Z( q$ H7 W- Y: A* ^+ j
i) Determine, through interactions, if the unity has identifiable
# G' }- M; D1 Z; @. I5 J3 O! Oboundaries. If the boundaries can be determined, proceed to 2. If not,) I5 D8 a- l$ E4 @7 |) B
the entity is indescribable and we can say nothing.<br/>9 z9 t& u3 J7 x9 G) |- D
ii) Determine if ther are constitutive elements of the unity, that is,
/ `5 ~ i; O/ f9 {components of the unity. If these components can be described, proceed4 ~* r& e$ p& c; R" Q5 v
to 3. If not, the unity is an unanalyzable whole and therefore not an# S% D/ o4 _8 h* I5 _
autopoietic system.<br/>8 f% }$ e, V# ?# h9 C2 l
iii) Determine if the unity is a mechanistic system, that is, the
3 X* L; h6 c# i. a' Ucomponent properties are capable of satisfying certain relations that O9 J" E& \. Z2 A7 K; S& R! z
determine in the unity the interactions and transformations of these
, U. D" t) w' ]& ^components. If this is the case, proceed to 4. If not, the unity is not
9 H& ^5 u- ?+ ian autopoietic system.<br/>
1 M$ v v7 V0 M8 D& d8 l, M2 yiv) Determine if the components that constitute the boundaries of the( L1 x+ J1 @; J+ g; k& b
unity constitute these boundaries through preferential neighborhood1 b1 f* T$ k0 [0 p
interactions and relations between themselves, as determined by their
8 z {1 C( Z1 c0 |- [ V% V6 \properties in the space of their interactions. If this is not the case,
" I. q! C r) ^6 ]you do not have an autopoietic unity because you are determining its
% r8 c2 f$ h! H# P j: V' bboundaries, not the unity itself. If 4 is the case, however, proceed to
, A6 h# X# J. C2 m! r5 [5.<br/>
( A' w. k; ~" E* Wv) Determine if the components of the boundaries of the unity are
3 I. Z" L/ V) k# Dproduced by the interactions of the components of the unity, either by* u# |4 A$ c6 n% e8 X
transformation of previously produced components, or by transformations
- ]' Z8 v* R2 V' o' w+ L) nand/or coupling of non-component elements that enter the unity trough4 n9 T5 n' k, M( ^$ j, y7 ^
its boundaries. If not, you do not have an autopoietic unity; if yes$ R3 C1 Z( X) x& t
proceed to 6.<br/>" v# ~1 d3 U9 ?% M: X4 Y: ~+ o
vi) If all the other components of the unity are also produced by the. F, b. x! n- |
interactions of its components as in 5, and if those which are not- A! f2 `- j% u: C& H! p; K6 m1 P
produced by the interactions of other components participate as
6 ^* h4 j. \2 H9 ]necessary permanent constitutive components in the production of other2 F' J# W3 `* C" E
components, you have an autopoietic unity in the space in which its8 }8 M1 R8 z, \; X( a5 K
components exist. If this is not the case, and there are components in
% D" R, d( a9 w/ o- zthe unity not produced by components of the unity as in 5, or if there/ Q/ B) G# F& L1 w! {9 a8 l0 T
are components of the unity which do not participate in the production
& f# ^9 o$ ]; S) a' M! @of other components, you do not have an autopoietic unity.<br/>' N0 z9 ?- C4 e/ N0 P& _ D, c
The first three criteria are general, specifying that there is an
1 C" F5 f6 j7 S W1 a) Widentifiable entity with a clear boundary, that it can be analyzed into
/ c3 d3 @. P6 q8 Z3 u' l/ ^* e& ucomponents, and that it operates mechanistically, i.e., its operation! |$ j, q: r4 \5 o m
is determined by the properties and relations of its components. The* I) M# A8 ^+ u* b/ ]3 z
core autopoietic ideas are specified in the last three points. These
: M7 K" g7 e6 j/ D3 vdescribe a dynamic network of interacting processes of production (vi),
) K/ C- V4 a& p5 W7 ]* k7 d1 d0 scontained within and producing a boundary (v) that is maintained by the0 k v2 j9 e9 w7 V
preferential interactions of components. The key notions, especially) e% f/ F3 E4 p7 Q
when considering the extension of autopoiesis to nonphysical systems,
/ z) z# t7 g. ?4 Yare the idea of production of components, and the necessity for a
' f# {! }/ D2 a7 g* dboundary constituted by produced components.<br/>
5 d" E" T( C0 S( P- ?These key criteria will be applied to the cell in the next section.+ L$ d* J* I5 X5 @5 Q! e( ~
This section will describe briefly embodiments of the autopoietic
7 Y4 P G! O% x0 Jrelations outlined above in the chemistry of the cell. Alberts et al.7 B/ N5 h* Y; U* H
or Freifelder are good introductions to molecular biology, as is Raven
9 i" m" c, | q& y! C# xand Johnson to the cell.<br/>6 ~# x( l) |& d" e
2.3 An illustration of Autopoiesis in the Cell<br/>3 p9 L& @$ h. h) f3 c# s
This section will describe briefly embodiments of the autopoietic ]; u& ]9 }8 Y+ W- r
relations outlined above in the chemistry of the cell. Alberts et al.( W0 }) d x; {6 G
are good introductions to molecular biology, as is Raven and Johnson to9 C/ l/ ]5 E7 r1 f
the cell.<br/>
, I+ G9 k0 l7 x# h2.3.1 Applying the Six Criteria<br/>1 ^9 z% ?% o: [: I y( v4 k1 u
Zeleny and Hufford analyze a typical cell with the six key points. A/ X. Y- |1 ~$ b. z( i. D
schematic of two typical cells is shown in Fig 2. One is a eukaryotic- ?7 B' ~% e) A$ x
cell, i.e., one that has a nucleus, and the other is a prokaryotic
9 `* R3 \: K7 L# O4 o9 k! ]. Scell, which does not.<br/>
: U5 y3 _/ V- B3 | [; D, O0 d1. The cell has an identifiable boundary formed by the plasma membrane. Thus, the cell is identifiable.<br/>6 n/ E' W K/ x1 o. z
2.The cell has identifiable components such as the mitochondria, the
1 L0 W# x/ c3 K c( Z1 \nucleus, and the membranous network known as the endoplasmic reticulum./ Q8 ^( z4 W& a( g( S
Thus, the cell is analyzable.<br/>; q. \7 ~) H8 s3 R. V* T
3. The components have electrochemical properties that follow general
: E( ~9 f( i$ I; v+ ~ E; D& rphysical laws determining the transformations and interactions that
$ [# j7 z/ I% T* Aoccur within the cell. Thus, the cell is a mechanistic system.<br/>
5 M0 X8 z7 K. `4.The boundary of the cell is formed by a plasma membrane consisting of
" Z: b% `! p) Iphospholipids molecules and certain proteins (fig 3). The lipid
* P: } b! L* k2 G4 Z! G xmolecules are aligned in a double layer, forming a selectively
v8 @. y9 |$ _, z; N8 Spermeable barrier; the proteins are wedged in this bilayer, mediating; k3 s9 ^, n0 I1 a; L: i
many of the membrane functions. A lipid molecule consists of two parts9 r. T. I1 R3 q1 p
– a polar head, which is attracted to water, and a hydrocarbon (fatty): N2 m& G4 W! ^: f5 I
tail, which is repelled. In solution, the tails join together to form1 o% c7 g4 \ g+ W' l1 i
the two layers with the heads outside. The integral proteins also have% G$ V( j4 Z2 H2 L
areas that seek or avoid water. The boundary is therefore6 z4 a% h% X* t# r! _' M' l- b# H
self-maintained through preferential neighborhood relations.<br/>
/ d: f, \/ Z$ M. R: s M5. The lipid and protein components of the boundary are themselves
' i* v. R* w& e! G$ P- a e/ `3 {produced by the cell. For example, most of the lipid molecules required/ \' i a' e! C: f* C# ?6 C' |' n
for new membrane formation are produced by the endoplasmic reticulum,' n! o1 W+ d9 b4 Z
which is itself a complex, membranous component of the cell. The
e; q0 @* | s# z3 x- u9 Oboundary components are thus self-produced.<br/>
( V6 d. Y/ F7 b( k D$ _6 {6. All of the other components of the cell (e.g., the mitochondria, the6 E$ R$ U2 |' l& x5 S! M
nucleus, the ribosomes, the endoplasimic reticulum) are also produced- G0 Q. u+ e/ e8 v% I4 R% q
by and within the cell. Certain chemicals (such as metal ions) not
7 Z5 V0 o0 U9 B0 ]produced by the cell are imported through the membrane and then become- h& k( @- Q" n! V6 Y' t$ T
part of the operations of the cell. Cell components are thus: o! c2 x3 |% ?& q( U. @$ M! x
self-produced.<br/>
9 E. D+ j$ r+ C5 I2.3.2 Autopoietic Relations of Constitution, Specification, and Order<br/>2 W+ q( s6 \" k
Apart from the six-point key, autopoiesis was also defined by three
* w" h6 k1 c+ R2 d# g4 F9 X0 J$ d$ {necessary types of relations. These can be illustrated as follows for a
/ j; ?7 @7 h; S' Dtypical cell.<br/>2 Y* r5 }+ n* @7 N2 q- n& v
2.3.2.1 Relations of Constitution<br/>
" V* e+ a% i! }1 e# i* aRelations of constitution determine the three-dimensional shape and
$ ]* {1 I. `7 J2 Lstructure of the cell so as to enable the other relations of production# O! Q( b$ h. r& a2 g3 W
to be maintained. This occurs through the production of molecules3 @. R" _' M, R& l! u( v; e
which, through their particular stereochemical properties, enable other
! e2 f: F% C% b4 i5 O2 ~& L1 eprocesses to continue.<br/>
7 v* E7 [1 U9 h+ F/ Z) A2 o! v7 g& yAn obvious example is the construction of membranes or cell boundaries.+ O' e) |$ v* Q+ a& o
In animal cells, the membrane surrounding the mitochondria, like that6 w1 V6 o+ y3 |3 J1 Q/ N
around the cell itself, serves to harbor cell contents and control the: I' m, s" S* u, W
rate of reaction through diffusion. Various reactive molecules are6 X7 \6 p# T( z9 b( r) P' A& Y
distributed along the inner membrane in an appropriate order to allow7 {# L* r4 G! y
energy-producing sequences to proceed efficiently. In plant cells, in
' N3 ?2 a3 K! ]3 v" taddition to the plasma membrane, there is a cell wall, which consists7 k& r& e/ ^1 B, @0 t
of cellulose, a material made up of long, straight chains of glucose
% L3 E- Y, U3 g5 N0 M7 x, eunits packed together to form strong rigid threads. These give plants7 t' l5 H6 o/ k/ B. W. V6 J
their rigidity.<br/>
+ D9 F) i- q! G hA second example is the active sites on enzymatic proteins. These act
- k7 u/ X$ z: J# ^! w/ Was catalysts for most reactions, changing a particular substrate in an& k4 F# R7 A4 |3 S
appropriate way to allow it to react more easily. Generally, the active& C( c* O: Y# ]# X" c$ V
site is found in certain specific parts of the enzyme molecule where
0 p3 K; M, S* F( x$ I" cthe configuration of amino acids is structured to fit the particular6 K0 w: [9 _3 h- m" e4 y
substrate, sometimes with the help of “activators” or co-enzymes. The) u& f2 C* G5 F6 {. g9 ?5 e
substrate molecule interlocks with the active site and in so doing
5 c& \; S* H' r1 kchanges appropriately so that it no longer fits, and thus frees itself.<br/>
3 a v+ {! @) t: [2.3.2.2 Relations of Specification<br/>3 H3 r& F& O. R& `" c/ d8 Z6 _
These determine the identity, in chemical properties, of the components
+ @" C) V; a2 u9 I4 d2 Xof the cell in such a way that through their interactions they% Z' ^; J2 E( a
participate in the production of the cell. There are two main types of
1 x; m! c) k5 O( P$ |4 Sstructural correspondence, that among DNA, RNA, and the proteins they
9 X- U: n$ B- sproduce and that between enzymes and the substrates they catalyze.<br/>; `$ D3 L' m' N# J& e& ]9 X( D
Protein synthesis is particularly complex because each protein is9 |, d) P' @. G7 r
formed by linking up to twenty different amino acids in a specific
( V- H2 I; b( }: \; G% {$ Z$ D9 mcombination, often containing 300 or more units in all. This requires
' \) \; c: V3 W+ ?an RNA template molecule, tailor-made for each protein, containing
2 x2 i4 H: P" O* \- Hspecific spaces for each of the amino acids in order, together with an: M# y" o" ?' \9 b
enzyme and t-RNA for each acid.<br/>
8 N" x% F4 W- G. l) ~As already mentioned, enzymes are necessary to help most of the) g* [$ H( x! q4 r
reactions in the cell, and again, each specific reaction requires an
- V. ?& r# X* n5 K! ? H) Tenzyme specific to the reaction and to the substrate involved. Hundreds
6 c" C8 X7 C' K6 Gof such enzymes are needed, and all must be produced by the cell.<br/>- K1 `' N! ~9 R% y/ Y K
2.3.2.3 Relations of Order<br/>
- b2 H0 _ ?3 A( R" G: W1 Q' iRelations of order concern the dynamics of the cell’s production l& P; ^2 R: ^/ B" b- s
processes. Various chemicals and complex feedback loops ensure that# j0 L4 z$ {. ^9 p2 ?
both the rate and the sequence of the various production processes0 g( V( p+ v- r8 j4 D
continue autopoiesis. For instance, the production of energy through; I! ^9 `# O& @& B0 ?$ ~
oxidation is controlled by the amount of phosphate and ADP (adenosine: o! L$ V7 U T5 ]/ m6 v5 B6 s
diphosphate) in the mitochondria. At the same time, reactions that use; V; k" ?' s1 l7 k1 ^
energy actually produce ADP and phosphate so that, automatically, a
6 J g& \/ [ [/ J! qhigh usage of energy leads to a high production rate of these necessary
^0 A. k, P n3 ~0 j1 c3 C! Osubstances.<br/>4 k) Y' d- @4 I# o: A: J$ I! @
2.3.3 Other Possible Autopoietic Systems<br/>0 o6 Q0 o8 m) H
An interesting question leading from the idea of the cell as an; w6 p9 e: B( W/ L4 _
autopoietic system is whether or not there are other instances of T+ T/ P6 d) H/ s& F: V
autopoietic systems. Are multicellular organisms also autopoietic
0 v- D6 |; S# u6 hsystems? Maturana is equivocal, suggesting that organisms such as) c8 S! x9 F+ z2 ~' ?# x
animals and plants may be second-order autopoietic systems, with the
$ f3 f5 r1 v4 i& H6 M% @components being not the cells themselves but various molecules P0 s9 \$ h* [3 V
produced by the cells. On the other hand, he suggests that some
1 }2 |8 c$ W8 {cellular systems may not actually constitute autopoietic systems, but! X" z: K+ a8 m6 f8 i. y
may be merely colonies. What about a system that appears to have a
' M9 K/ e8 I0 D) oclosed and circular organization but is not generally classified as0 E; e8 ^8 G+ F+ ?: H3 \, h
living, such as the pilot light of a gas boiler? Finally, what about( u' g2 c. }; o, [% ` `0 P; r
nonphysical systems such as the autopoietic automata mentioned in
5 R) f6 q: }6 J, ^' o4 ]2 m" C/ osection 2.2.1 and described more fully in section 4.4, or systems such
# f. \. @% i: ~1 M& P: oas a set of ideas or a society? These possibilities will be discussed$ h1 u* O+ v" G% j' r0 i$ n) J
in more detail in Section 3.3.<br/>0 s0 F2 E) `3 w
2.4.Applications of Autopoiesis in Biology and Chemistry<br/>3 s6 G& ^" B- C3 x3 t! N/ \2 `
One would have expected that, given the importance and nature of its
4 e1 H& n* ~( r5 c3 H3 Wclaims, autopoiesis would have had a major impact on the field of* A$ [+ l: [! r! M, w
biology. In fact, for many years there was a noticeable reluctance to" n4 i9 Q7 E0 N2 T, ?0 C0 @
take the ideas seriously at all. In 1979, I wrote to an eminent British; \9 k3 i( [# E8 _2 a( A
biologist – Professor Steven Rose at the Open University – querying the
$ \7 a' T/ a8 `0 Sstatus of autopoiesis. He replied to the effect that he did not wish to
% Z& M6 t& \% G) G/ U' L5 bcomment on autopoiesis but that Maturana was a reputable biologist. One
. W2 k4 Y2 k! A2 g' f+ Q. vnotable exception is Lynn Margulis, whose own theory, that eukaryotic
. E6 W- r& ]5 @; ]8 p( K. a4 R$ [cells evolved through the symbiosis of simpler units, is itself quite
! K- X. k7 L1 A+ ]; C1 d3 pcontroversial.<br/>" Y" B% Z- T2 r( b- Q& L- Z
However, recently interest has been growing in two areas: research into3 F8 Z5 F" t7 u
the origins of life and the creation of chemical systems that, although
' W7 e; y% V: d4 z( f: d. d- V anot living, display some of the characteristics of autopoietic, D& j! `( d0 Z% H
self-production. Autopoiesis has also been compared with Prigogine’s1 X# P+ T- z! V+ N, G# S9 g* @
dissipative structures. Varela has also pursued work on the nature of
% q2 t, o% e a4 }9 Y2 W% }$ Jthe immune system, viewing it as organizationally closed but not1 g" v4 [. R$ ]: r6 H" A) h
autopoietic. However, as this topic is very technical and not of
3 d" Q5 o; `/ g' N* I4 ]; R; v3 Bprimary relevance, it cannot be pursued here.<br/>2 e! o: V3 w+ a6 b' z- S
2.4.1 Minimal Cells and the Origin of Life<br/>
- G) T# o- `! T- |, ^& @5 x/ SThere are two main lines of approach to theories concerning the origin0 D( w" T! J; k4 Q
of life on Earth. In the first approach, based on study of the enzymes1 Z# l: K4 k. } u( i* g5 {- V
and genes, life is characterized as being molecular and a defining
: Z ?+ f1 p& K+ Lfeature is the structure and function of the genes. In the second
, s9 f: [7 W* M# k8 A6 S2 Aapproach, life is characterized as cellular, and its defining feature
1 v9 U9 G# [' v- o: pis metabolic functioning within the cell. However, neither approach can9 @; J( F0 B) d7 V7 H
really specify a standard or model for life against which important9 d9 Z1 V) C" D# h5 y% d
questions may be answered. In particular, at what point did prebiotic
5 L& x5 A" i: H# z. a2 S3 ]chemical systems become biotic living systems? And how could we# t) T' f0 k; _! D
recognize nonterrestrial living systems. Which might be radically, p! w) `2 H& b3 B, m6 U0 H
different in structure from our own?<br/>
& C* ~* |0 _- |: t+ B# cFleischaker proposes that the concept of autopoiesis, together with
3 H9 h5 Z8 k1 t% rnotions of minimal cell, can provide a sound theoretical framework to7 I7 y& ~# z, y. f8 ~
tackle these questions within the second tradition mentioned above.) q/ J2 D5 d5 B9 m
Autopoiesis clearly does aim to provide a specific and operationally
6 j/ I2 {% {& guseful definition of life, although Fleischaker argues that the concept
6 q# }3 D5 O) v2 _9 sof autopoiesis does need some modification. This modification would: \7 |9 ]+ a& K3 ^2 I3 F
restrict “living” systems to autopoietic system in the physical domain, p3 _; T3 W$ w
rather that allow the possibility of nonphysical living systems, a/ b. P$ q- O) b9 s( U
possibility which ( as mentioned above) is left open by the formal& r5 W1 K; v# f0 R7 p0 U
definition of autopoiesis. This will be discussed in Section 3.3.2<br/>8 H# H5 Q7 P5 h% K
Given autopoiesis (or modified version) as a definition of life, the* J0 P7 D9 |' ?. H
next step in theorizing about the origin of life is to consider how an
x1 j& D; N% [( I1 y4 ielementary autopoietic system might have formed. Note that autopoiesis# R2 j1 e5 N& x4 Q7 z
is all or nothing. A self-producing system either exists and produces9 W/ H& g! m8 D+ ?, }9 K( _( P
itself or it does not – there can be no halfway stage. This leads to: }% ~( E d" Q) g2 X2 v
the idea of a theoretical “minimal” cell which could plausibly emerge,9 R4 n! _% d, E! a3 Z6 e1 ` h
given the early conditions on earth. In fact, Fleischaker considers6 @2 X7 ]7 S9 _0 W. B
three different characterizations of minimal cells: a minimal cell
/ G2 d. k$ \6 \2 Yrepresentative of the evolved life forms that we know today; a minimal1 e3 f! G( \0 ?! ?3 M
cell that would characterize both terrestrial and nonterrestrial life
7 t, s; }* Q. `$ ]! E; ~2 s# Lregardless of its constituents.<br/>8 Y9 m: D3 }6 ]5 l; H' h
About the last, little can be put forward beyond the six-point, F6 p0 T2 M1 m7 C2 ?6 [* g L+ v0 q S
autopoietic characteristics in the physical space; to be more specific8 ^# E1 m% f$ z# ~) W* j* N1 U0 o
would constrain the possibilities unnecessarily. On the other hand, we2 o* v- o" B5 x% G# n9 f
can be quite specific about a modern-day cell. Such a cell could be
6 d3 f3 G: | F7 N9 vdescribed as “a volume of cytoplasmic solvent capable of DNA-cycled,
# e" K5 E2 \3 HATP-driven and enzyme-mediated metabolism enclosed within a
& L* T- ]" x. S+ d; Bphosphor-lipoprotein membrane capable of energy transduction”, This
5 Q9 l. A; K( m" @* z- N# r* t3 Wgeneralized specification can cover both prokaryotes (bacterial) and( r, X3 \ r8 {- X+ z* g d
eukaryotes (algal, fungal, animal, and plant cells) even though there* |$ _" [, z. E6 d" O
are important differences in their operation.<br/>
4 y2 t1 w+ { i K n4 W: HThe most interesting minimal cell scenario concerns the origin of life.
6 n( K! n2 i B8 ^4 c2 {) d( HThe first cell need be only a very basic cell without the later; b6 b2 O6 h4 _2 Z- t) t
elaborations such as enzymes. Fleischaker suggests that such a cell
+ \1 ~) z9 {( V. D9 imust exhibit a number of operations (Fig.2.4):<br/>: R+ r7 s8 L% k: x& b- }' |+ y6 X
1、The cell must demonstrate the formation and maintenance of a boundary* k$ s( K9 E6 n
structure that creates a hospitable inner environment and allows# W/ T+ m- N- @9 a- k
selective permeability for incoming and outgoing molecules and ions.
5 {/ a% H, A" j6 Q( W/ ZThe lipid bilayer found in contemporary cells is a good possibility q+ p; Y1 G- E8 f" j K, R
since the hydropholic nature of lipid molecules leads them to form+ `, @" L3 N0 [+ X
closed spheres in order to avoid contact with water. Lipid bilayers are* K8 _6 x# k! i1 a/ J
also permeable in certain ways – for example, to flows of protons or2 M5 F* ?7 F5 Y2 H
sodium atoms – without the need for the complex enzymes prevalent in
, T2 I( m$ J8 v3 k0 k# N" o1 Mcontemporary cells.<br/>: @! j A7 ^3 j- J% B( W# }
2. The cell must also demonstrate some form of active energy0 C7 \; Q' m- v9 Y, N0 w/ ]
transduction to maintain it away from entropic chemical equilibrium.- X" s" A7 k- B/ Z6 Y
One possibility is an early form of photopigment system driven by
% f) U1 F2 @, ~9 Flight. Pigment molecules would become embedded in the membrane and act- v3 \+ l7 D. u; y9 X
as proton pumps, leading to the concentration of variety of raw0 y0 J+ |- R9 Z' p6 G' Z; ^
material in the cell.<br/>
) M8 G4 ]. z m% w) @! J+ A3. The cell would also need to transport and transform material4 z+ S; |& j3 D2 O W4 ^/ m/ p8 f
elements and use these in the production of the cell’s components and& \0 _7 e) h0 l. u. A+ B
its boundary. A possible start in this direction would be the import of' z3 _- D1 q3 O U) G) L
carbon dioxide and the physio-chemical transformation of its carbon and; x1 m. {; p) F* Y6 W
oxygen through light-driven carbon fixation.<br/>4 d6 f' K7 c1 I4 I
What is important is not the particular mechanisms for any of these
# Y; p) h- |" Egeneral operations but that whichever mechanisms are postulated, all6 e5 |6 s% ?) q/ z' {
operations need to be part of a continuous network to form a dynamic,
$ A$ ?' K+ q5 dself-producing whole.<br/>! h. p4 ?& g6 B& N" x4 h/ w5 ?
2.4.2 Chemical Autopoiesis<br/>
! d/ o3 S+ D C+ z8 ]9 pBeyond theoretical constructs of minimal cells, it is also interesting/ `/ f Z1 u$ a( D* C1 e2 F/ x
to look at attempts to identify or create chemical systems based on( u6 N/ s* a5 Y; i& O6 s# I
autopoietic criteria, and to consider whether or not these are living.
+ x6 @7 M( x+ ~+ DWe shall look at three examples: autocatalytic processes, osmotic7 W/ [9 q2 i G) c/ _8 ?
growth, and self-replicating micelles.<br/># z; N% c) d4 M$ [ g. S
2.4.2.1. Autocatalytic Reactions<br/>
: [, n6 V( T7 W5 ]' [+ BA catalyst is a molecular substance whose presence is necessary for the3 O& L; P+ b" e9 u- D% M
occurrence of a particular chemical reaction, or which speeds the
2 R3 ^ @, H) W* t! }reaction up, but which is not changed by the reaction. The complex3 Z) c+ H/ G9 w( i! O" {
productions of contemporary cells (as opposed to cells that may have1 c+ \( ~+ V7 o9 {9 n& i. i
existed at the origin of life) require many catalysts, and this is one
E- i( q K0 J' G: J& b& g5 Uof the main functions of the enzymes. An autocatalytic process is one' n: D( I+ [9 J6 \2 \
in which the specific catalysts required are themselves produced as
* x' z* Y; C2 T4 @6 kby-products of the reactions. The process thus self-catalyzes. An
4 g; M; K5 |5 }1 ]+ e" n; texample is RNA itself which, in certain circumstances, can form a
2 y3 h! h+ S: d* zcomplex surface that acts like an enzyme in reaction with other RNA |: Z- p9 Q5 d* L/ J* K* P
molecules (Alberts et al.) Kauffman has a detailed discussion within
8 F9 _! v0 _6 y9 L7 b% y) othe context of complexity theory.<br/>1 Y( [( u' a# ^
Although this process can be described as a self-referring interaction,
4 Z+ h0 N5 m+ fthe system does not qualify as autopoietic because it does not produce
, `7 m$ A" A+ T% ^% ?) \its own boundary components and thus cannot establish itself as an
" V0 ~' S+ X, n; M- hautonomous operational entity (Maturana and Varela). Complex,$ U% Z# H4 @5 L, Z
interdependent chemical processes abound in nature, but they are not0 a8 k- ]: Y) o
autopoietic unless they form self-bounded unities that embody the
' S {4 }/ G) Y& c* K( yautopoietic organization.<br/>" p/ o+ }. c( ~$ Z
2.4.2.2 Osmotic Growth<br/>
& s4 h. D* W- s* j# W1 O! p6 kZeleny and Hufford have suggested that a particular form of osmotic7 Z$ Q. D8 ?7 t
growth, studied by Leduc, can be seen as autopoietic. The growth is0 H) P) u/ ^& ?; Y# k
precipitation of inorganic salt that expands and forms a permeable+ ]* N% M7 N7 L: w5 G. E: C7 K
osmotic boundary. This can be demonstrated by putting calcium chloride! p# J9 d3 W0 d9 V+ `+ Y: E9 [$ Q
into a saturated solution of sodium phosphate. Interaction of the7 P' R$ z. ~, y. l* Q4 c) w9 `6 s
calcium and phosphate ions leads to the precipitation of calcium1 m1 t+ o% B- H* @- Z
phosphate in a thin boundary layer. This layer then separates the" n- T1 j6 b4 n$ [0 f
phosphate from the calcium, water enters through the boundary by& L5 }' w. F/ c
osmosis, and the increased internal pressure breaks the precipitated
/ u: ~' q6 h0 \5 t: S& Ecalcium phosphate. This break allows further contact between the
- J5 l5 j8 h3 Iinternal calcium and the external phosphate, leading to further
' k! m, |/ `- A3 z( z: mprecipitation. Thus the precipitated layer grows.<br/>
6 {9 `: _( G# }5 B8 D9 Q: pZeleny and Hufford argue that this system fulfills the six autopoietic criteria:<br/>
) b+ O3 |: T7 R8 Z* P% G' i) ]1. It is distinguishable entity because of its precipitate boundary.<br/>- l0 L8 Y: b0 ?, t `
2. It is analyzable into components such as the calcium phosphate boundary and the calcium chloride.<br/>
4 a, h* {6 b3 S2 j; W2 D4 K! z3. It follows mechanistic laws.<br/>
' `8 @: x4 t h0 a0 r& `, Q, n& w4. The boundary components (calcium phosphate) aggregate because of their preferred neighborhood relations.<br/>7 @$ ~0 l! n/ ^! d7 w1 U
5. The boundary components are formed by the interaction of internal
: M1 W0 u7 ?! gand external components following osmosis through the membrane.<br/>
( Z1 m) ]- p9 o6. The components (calcium chloride) are not produced by the cell but4 p! X& q+ g w) Z: w
are permanent constituent components in the production of other1 c! O8 q9 P. C) m
components (the precipitate)<br/>
$ m& f8 C4 b# X4 W6 M) sThis hypothesis does cause problems, as Leduc’s system is clearly
0 |( V3 z2 M0 n* R- s. M* Yinorganic and not what would be called living. If it is accepted that
) D, u7 m* F1 u( O {1 b4 L1 j6 wthe system does properly fulfill the criteria of autopoiesis, i.e.,) t! P6 H$ @4 f& ^( Y5 G) t
that it is an autopoietic system as currently defined, then either we# Z2 C: ?; \, |, n0 O. F
must expand our concept of living or accept that autopoiesis is in need
) o! N! C1 B# Z1 ^- b. Cof redefinition to exclude such examples. In fact, it is debatable+ V9 ?" L2 A, S* ]; t7 n# i1 G2 _
whether or not this osmotic growth does correctly fulfill the six: r8 g. ~/ u( n/ A8 I- _1 y3 k. S
criteria. It certainly meets the first three, but it is not clear that
; s. P0 J& R2 V% b2 xit is a dynamic network of processes of production.<br/>, |) }/ s; F2 v: B( C
As for the fourth criterion, the precipitate that forms the boundary is! T y" M7 T" z8 s. ?5 b
unlike a cell membrane. It is static and inactive, more like a stone
* @* q; P \; A8 @5 x ~wall than an active membrane. It is not formed through “preferential8 }% q3 A- K- c! t& `5 I! h" ~
neighborhood interactions”; in fact, once formed, it does not interact+ ?( O1 j7 x, B! r4 w g9 ?
at all. Considering the fifth criterion, the boundary components are
. B W4 y8 Z, h, i( ]2 z$ znot continuously produced by the internal processes of production.( o! p6 O# v. U! W" G9 r, w5 P
Rather, a split or rupture occurs and more boundary is precipitated at+ I @- {1 O/ d
the split through the interaction of internal and external chemicals.
0 T& @' _7 B4 UIt is only because of, and at, the rupture that new boundary is
# V. `9 ]. L6 Wproduced. Finally, chloride, which is introduced artificially at the* S6 x I. c, ~) c$ i1 I
beginning, is not produced by the system, and eventually runs out.<br/>, X2 R: G$ B! R2 P
2.4.2.3 Self-replicating Micelles<br/>
8 d3 A' X2 c/ d7 P* NAn approach with more potential, currently being researched by Bachmann! r( M8 w5 X% e
and colleagues, was first proposed by Luisi. It has been discussed by% ]3 E$ k5 b i6 c2 I( y" I
Maddox and Hadlington. A micelle is a small droplet of an organic
& Y, `3 m' u6 h" k9 gchemical such as alcohol stabilized in an aqueous solution by a- o7 G& Q9 z0 A( _) O4 F Q
boundary or “surfactant” A reverse micelle is a droplet of water% G0 q) w+ Y. W: K
similarly stabilized in an organic solvent. Chemical reactions occur/ P# r. M( R* Y/ X; {1 S7 a1 @
within the micelle, producing more of the boundary surfactant.
) W$ ~9 r$ k# S! |Eventually, this leads to the splitting of the micelle and the
* \# j+ G0 M- \" j. P% w( ?generation of a new one, a process of self-replication. Experiments
9 K7 |3 }. o: H% \( ^have been carried out with both ordinary and reverse micelles and with
2 Z( ]4 Z& o ~0 W: dan enzymatically driven system.<br/>$ W. F: w3 B$ Y+ k# k- d! b$ w
In the reverse micelle experiments, the water droplets contain
( z+ q3 r1 Q6 W) m( ^& |dissolved lithium hydroxide, one of the surfactants is sodium
: I+ x$ L' m) h: M9 ^( k; eoctanoate, and the other is 1-octanol, which is also a solvent. The; q2 `( p+ S4 W
other solvent is isooctane. The main reaction is one in which the3 [) s M9 G. Q. W& D/ D' X$ |: O. v7 ~
components of the boundary are themselves produced at the boundary.5 x X8 q5 n/ D. u6 u
Octyl octanoate is hydrolyzed using the lithium as a catalyst. This# @) ]4 B+ G; {; I" E
produces both the surfactants (sodium octanoate and 1-octanol). Since$ p7 n- H. o' `! E
the lithium hydroxide is insoluble in the organic solvent, it remains3 h# l" ?. j$ |2 {
within the water micelle, thus confining the reaction to the boundary' X% a, ~% F) M
layer. Once the system is initiated, large numbers of new micelles are
4 o s7 x `1 k, Hproduced, although the average size of the micelles decreases.<br/>
& V# j& \4 G( |6 \- ?# q! LIt is not clear that these systems could yet be called autopoietic.
% ^% s1 W. [ Y# D! s! s% _, BFirst, the raw materials(the water-lithium mixture or the enzyme
2 @; B9 E2 N4 k& ucatalyst) are not produced within the system. This limits the amount of
?. |; f1 I3 X: jreplication which can occur; the system eventually stops. Even if these, r0 _5 i5 a) y4 v3 s7 @
materials could be added on a regular basis, the system would still not
- `5 b& M1 [, a D8 wbe self-producing. Second, the single-layer surfactant does not allow# x" X" K' q5 O
transport of raw materials into the micelle. For this to happen, a6 Z" `! S4 z) }$ _) y: q5 W
double-layer boundary would be necessary, as exists in actual cell0 e: g5 z: w. X
membranes. Moreover, the researchers themselves, and seem most+ u- x- w& O2 c( r' K4 }$ s- g
interested in the fact that the micelles reproduce themselves, and seem
. y8 s/ D$ d+ G: Vto identify this as autopoietic. However, reproduction of the whole is
, e4 ^/ M9 E0 \- S R1 t6 b) u& `quite secondary to the autopoietic process of self-production of
- D5 Y; a- w9 Y& c8 [: T9 G2 w. y/ dcomponents. Nevertheless, this does represent an interesting step1 ^, ^+ ]( M+ X+ o- n/ y
toward generating real autopoietic systems. |
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