UKO ZYLSTRA

LIVING THINGS AS
HIERARCHICALLY ORGANIZED STRUCTURES

ABSTRACT. Hierarchical organization is an essential characteristic of living things.

Although most biologists affirm the concept of living things as hierarchically organized
structures, there are widespread differences of interpretation in the meaning of hierarchy
and of how the concept of hierarchy applies to living things. One such basic difference
involves the distinction between the concept of control hierarchy and classification hierar-
chy. It is suggested that control hierarchies are distinguished from classification hierarchies
in that while the former involve authority relationships between levels, the latter do not.
This is illustrated in an analysis of proposed hierarchies of replicators and interactors.
The analysis of levels of hierarchies and their relationships also brings up the part-
whole problem. An authority relationship between levels implies that the whole has a
determining influence on the parts that make up the whole, and that parts have no
independent, meaningful existence apart from the whole. The concept of an authority
relationship in a part-whole relationship introduces the question of the independence or
sovereignty of the components of the subordinate levels in a hierarchically organized
living thing. This problem is discussed along with an analysis of the rather novel theory
of enkapsis proposed by H. Dooyeweerd, in which he distinguishes part-whole relation-
ships from enkaptic relationships.

J. H. W o o d g e r (1929) p o i n t e d to a f u n d a m e n t a l characteristic o f living

things by his rhetorical question, "Is not organization the very back-
b o n e of the c o n c e p t o r g a n i s m ? " . A r e m a r k a b l e feature of a living
organism is that its identity is maintained during the d y n a m i c flux of
its constituent parts, b o t h in its d a y - t o - d a y existence and in its g r o w t h
and d e v e l o p m e n t . H o w this is accomplished is a p r o b l e m which has
long interested biologists.
T h e resolution of this p r o b l e m requires u n d e r s t a n d i n g the structure -
indeed, the unity, wholeness and integrated nature - of living things.
This p r o b l e m comes into focus in the often used expression 'living
m a t t e r ' . But as W o o d g e r correctly observed, " t h e r e is no such thing to
be f o u n d in nature as 'living m a t t e r ' " ( W o o d g e r , 1929, pp. 308-09).
W e always find whole living organisms. T h e expression 'living m a t t e r '
is really an o x y m o r o n ; m a t t e r as such is not alive. Molecules do not
b e c o m e 'alive' w h e n they enter a cell or w h e n they are i n c o r p o r a t e d
into cells or tissue structures. T o recognize a cell or any living thing as
being alive, does not m e a n that the molecules which c o m p o s e such
living things are alive. E v e n c o m p l e x protein and D N A molecules are

Synthese 91: 111-133, 1992.

© 1992 Kluwer Academic Publishers. Printed in the Netherlands.
112 uI~O ZYLSTRA

simply physico-chemical things subject to physico-chemical ordering

principles.
But if matter is not alive, what then is the basis for the integrated
unity of living things? What is there about the wholeness of living things
which accounts for their being alive? The concept of organization has
become one avenue for understanding and resolving this problem. Or-
ganization is seen as fundamental to the integrated unity of living things.
The emphasis is not only on the recognition of the phenomenon of
organization but also on the fact that it is present due to, and structured
by, ordering principles which determine the organization and conse-
quently the basic character of life. Furthermore, these ordering prin-
ciples are considered by many proponents of organization theory to go
beyond those which hold for chemical and physical phenomena. Biotic
ordering principles are also involved in life phenomena.
The concept of organization as a basic feature of living things began
to receive prominent status in biology with the rise of organismic bi-
ology in the 1920s. Key figures in this development were J. H. Woodger,
L. von Bertalanffy and P. Weiss. They not only stressed the importance
of organization as a key characteristic of living beings, but they also
emphasized the hierarchical nature of the organization. A basic theme
of organismic biology came to be that parts of a whole are affected or
influenced in some way by the whole, or, as commonly stated, the
whole is greater than the sum of the parts. But the simplicity of this
statement doesn't fully capture the significance of the nature of parts
and wholes and the relationships between them. One of the purposes
of this paper is to analyze the nature of 'parts' and 'wholes' in the
context of hierarchy theory by introducing a theory of enkapsis (enclos-
ure).
The theme of hierarchical organization is now commonly accepted
in biology. Pattee even argues that "hierarchical control is the essential
and distinguishing characteristic of life" (Pattee, 1970, p. 120). Most
general biology textbooks affirm the concept of living things as hierarch-
ically organized structures in which the biotic realm is described as
consisting of levels arranged in such a way that a particular level serves
as a sublevel for the next higher level. However, though the concept of
hierarchy has become widely accepted in biology, there are considerable
differences not only in hierarchy theories, but also in applications of
the concept of hierarchy to living things. These differences can lead to
 H I E R A R C H I C A L L Y O R G A N I Z E D STRUCTURES 113

Table 1. Terminological distinctions in hierarchy theory

One type may be a form of another type as indicated in the sub-listing.

TERM REFERENCE
Control hierarchy Grene (1969, 1987, 1988)
Taxonomic hierarchy Grene (1969, 1987, 1988)
-Linnean type hierarchy
Command hierarchy Allen and Starr (1982)
Constitutive hierarchy Mayr (1982)
Aggregational hierarchy Mayr (1982)
-Taxonomic hierarchy
Inclusive hierarchy Mayr (1982)
-Nested type hierarchy
-Taxonomic hierarchy
Exclusive hierarchy Mayr (1982)

considerable confusion in the understanding of hierarchy theory and

consequently in our understanding of the nature of living things.
A fundamental issue in dealing with hierarchy theory is whether the
perceived hierarchy has an ontological basis or whether it is chiefly of
epistemological significance: What does the concept of hierarchy do for
us in understanding the nature of life? In other words, is hierarchy used
as a way to account for the integrated unity of living things - a distinc-
tive mode of being - or is it simply a way to organize and rank the
diverse structures and types of organisms? Are the various biological
disciplines organized in a hierarchical manner for conceptual con-
venience, or does such organization reflect the actual structure of living
things? I suggest that the principle of hierarchy is mainly an ontological,
not an epistemological matter, and that the possibility of epistemologi-
cal hierarchies is based on the reality of ontological hierarchies.

C R I T E R I A FOR H I E R A R C H I E S

Due to the various types of hierarchy theories, it becomes necessary to

distinguish among the hierarchies by establishing criteria for analyzing
them. Table 1 provides a short list of terminological distinctions which
are applied to hierarchies. An important element in such distinctions
concerns the nature of the relationships between the levels of the
114 UKO ZYLSTRA

hierarchy. Etymologically, the term 'Hierarchy' implies some form of

rule or authority of a higher level over a lower level. Authority or rule
is still an important criterion in analyzing hierarchies, although the
meaning of this authority relationship or rule needs clarification. In
addressing the problem of the variety of meanings of hierarchy, Grene
(1969, 1987, 1988) proposes a basic distinction among hierarchies be-
tween control hierarchies and taxonomic hierarchies.
Control hierarchies are characterized by some type of authority re-
lation of a higher level upon the elements of a lower level (Pattee,
1970, 1973a). Polanyi (1968) develops the concept of dual control be-
tween levels of organization and argues that the lower level provides
the initial conditions for the higher level, whereas the higher level sets
boundary conditions for the lower level. An example of this dual control
is the cell, in which the molecules provide the substratum (initial con-
ditions) for cell activity, and the cell provides the context (boundary
conditions) for molecular activity.
Taxonomic hierarchies are those formed by grouping entities in some
ranking order. Each level is a rank in the ordering of such entities.
Whether or not there exists some authority relation between the ranks
depends largely on the nature of the organization. In a taxonomic
hierarchy of, for example, species, genera, families, orders, etc., an
authority relation is absent.
Mayr (1982) introduces two other criteria for distinguishing hierarch-
ies: a distinction between constitutive and aggregational hierarchies;
and a distinction between inclusive and exclusive hierarchies. In consti-
tutive hierarchies the members of a lower level are combined in forming
(constituting) the next higher level. This higher level becomes expressed
through the interaction of the constituents which make up the next
lower level. An example of a constitutive hierarchy is the series: mol-
ecules, cells, tissues, organs, organ systems, and organism. An aggre-
gational hierarchy results from a collecting or grouping of entities into
a larger unit. The larger unit does not necessarily involve interactions
among the entities; it is simply an aggregate of the units which are
grouped by the definition of their properties. The larger unit does not
possess any novel properties apart from the common collective proper-
ties of the units. A pile of sand is a simple example of such a relation-
ship. Aggregational hierarchies would include the taxonomic type hier-
archies in which entities are assigned to a particular rank, such as in a
Linnean hierarchy of taxonomic characters.
 HIERARCHICALLY ORGANIZED STRUCTURES 115

Mayr introduces the designations 'inclusive' and 'exclusive' to distin-

guish between arrangements in which lower ranks are subdivisions or
Subgroups of the next higher level, thus inclusive, and those in which
the lower rank is not a subdivision of a higher rank, thus exclusive.
An example of the former would be the subgroups of proteins and
carbohydrates included in the class of organic molecules; an example
of the latter would be the ranks in an army where sergeant is not a
subdivision of lieutenant. Allen and Starr (1982) and Grene (1988)
appropriately designate this latter type of hierarchy as a command
hierarchy (as distinct from control hierarchy). With this criterion, all
inclusive classifications would be hierarchical. If we distinguish taxo-
nomic hierarchies from command hierarchies, then all taxonomic hier-
archies would be inclusive, with the higher groups containing larger
and larger numbers of individuals or entities. Such hierarchies are
often also described as nested type hierarchies where one category of
individuals is simply contained within another category of individuals.
There is little hint of a control element in such nested hierarchies.
From Mayr's discussion it is not clear, however, whether the distinc-
tion between inclusive and exclusive hierarchies applies equally to both
aggregational and constitutive hierarchies or only to aggregational hier-
archies as indicated in the above examples. In a constitutive hierarchy
series such as molecule, cell, tissue, organ, etc., molecules are not
simply subgroups of cells, nor are cells simply subgroups of tissues, nor
are tissues subgroups of organs, etc. Subgroups are not equivalent to
parts or components. A subgroup is characterized by the group to
which it belongs. This implies that inclusive hierarchies which are
characterized by subgroups rather than parts are distinct from constitu-
tive hierarchies. Grene (1988), in recognizing alternative meanings of
inclusion in 'Chinese box' hierarchies, makes the distinction between
'arbitrary serial enclosures' and 'constitutive serial enclosures'. I suggest
that the term 'inclusive' be restricted to the arbitrary type of serial
enclosures where we are confronted with subgroups that contain mem-
bers or aggregates as distinct from parts. In a constitutive hierarchy we
are confronted with distinct types of integrated structures as parts at
each level, with each type of structure subject to the peculiar set of
ordering principles which holds for that level. In the example given,
the ordering principles for cell structure and function are distinct from
the ordering principles for molecular structure and function.
Another criterion which then must be considered concerns the con-
116 UKO ZYLSTRA

cept of the part/whole relationship as distinct from a member/class or

member/aggregate relationship. This distinction has to do with the
nature of the relationship between the levels. The concept 'whole'
suggests organization; it is not simply a collection or aggregate. This
organization is such that the parts that make up the whole display
mutual relations "so that the properties of a part are different when it
is in its place in the organic hierarchy from what they are when it is
removed from it" (Woodger, 1929, p. 310). A part/whole relationship
is thus an integral, functional, as well as structural, relationship such
that the loss, change, or addition of a part affects the entire whole with
a consequent loss or change of properties of the whole. If one were to
cut a tree into two-foot sections, those two-foot sections are not be
considered as true parts of the tree. By contrast, in a member/class or
member/aggregate relationship the members have an integrity of their
own which is not affected or altered by being a member of a class.
Classes or aggregates are simple conglomerates. They not only lack
organization above or at the level of the aggregate (Woodger, 1929),
but they also lack organization in the level below the aggregate, viz.,
the level of the members. As Weiss affirms, "an empirical dichotomy
arises between simple conglomerates and the type of ordered complexes
which we designate as systems" (Weiss, 1969, pp. 9-10).
If hierarchy theories are to have significant explanatory value, then
they must go beyond a simple ranking or grouping of entities according
to some gross morphological featt~res and/or some general function or
behavior. Although hierarchical rankings of objects and individuals may
be useful for classificatory purposes, they convey information about
biological structures only when they reflect the integral character of the
individuals or structures being ranked. Ultimately, hierarchy theory
should be founded upon an integrated ontology of organisms. Attempts
to develop such an ontological framework for hierarchy are usually
present in the proposals which argue for some form of constitutive
hierarchy. An analysis of hierarchy theories on the basis of these criteria
may serve to judge the usefulness of various hierarchy theories that
have been proposed in the past few decades.

CONSTITUTIVE HIERARCHIES

One of the classic examples of constitutive hierarchies is the one pro-

posed by H. Simon (1962). Simon defines hierarchy as a "system that
 HIERARCHICALLY ORGANIZED STRUCTURES 117

is composed of interrelated subsystems, each of the latter being, in turn,

hierarchic in structure until we reach some lowest level of elementary
subsystem" (Simon, 1962, p. 468). Although Simon recognizes a part/
whole dimension in these complex systems, he does not develop the
concept of control, whether it be single level or dual level control, for
such hierarchies. Since constraints (boundary conditions) of a level
upon its sublevels are absent, this type of hierarchy is designated a
structural hierarchy in order to distinguish it from a constitutive hierar-
chy in which the element of control is present.

CONTROL HIERARCHIES

Control hierarchies are characterized as those which display a dual type

of control between the levels of the hierarchy. The systems hierarchy
theories developed by P. Weiss and L. von Bertalanffy are good ex-
amples of such control hierarchies. Weiss (1969, 1971, 1973) viewed
living organisms as a series of subordinate sub-subsystems. Each level
serves to control the parts of the subordinate level. The control is seen
as restraining the degrees of freedom of the parts of the subordinate
level. At the same time, however, the constraints allow for a higher
form of freedom in the dynamic flux of the constituent parts (see also
Pattee, 1973a; Allen and Starr, 1982). This paradox of greater freedom
through constraints not only is an important characteristic of control
hierarchies, it is also basic to the phenomenon of life. The very possibil-
ity of organization of living systems comes about through constraints
which function as boundary conditions of one level upon its sublevel.
This type of control is also a basis for the distinction between living
systems and machines. As stated by Weiss, "in a system, the structure
of the whole coordinates the play of the parts; in the machine, the
operation of the parts determines the outcome" (Weiss, 1969, p. 13).
Although Weiss makes his systems hierarchy analogous to a set of
Chinese boxes, I think the analogy is inappropriate. In a set of Chinese
boxes, one box is simply contained within another, larger box, there is
minimal interaction between the boxes. Weiss, however, stresses the
dynamic interactions between adjacent levels and across levels. He
actually describes multiple interactions of a particular level with all the
other levels, both above and below it. Such a view goes much beyond
the simple view which is illustrated by that of sets of Chinese boxes.
L. von Bertalanffy introduces a similar concept of a systems hierarchy
118 UKO ZYLSTRA

in his development of a general systems theory (1952, 1968). He empha-

sizes that "the problem of life is that of organization" (Bertalanffy,
1952, p. 12). This organization is characterized by a hierarchy of systems
levels. Moreover, a system isn't defined simply by its parts or as a sum
of its parts but, rather, as a complex of parts or elements which stand
in interaction. It is the ordered arrangement of parts and processes
which define a living system. Bertalanffy (1952) also affirms that a living
system is not to be identified as some living substance; there is no such
living substance. Although Bertalanffy seeks to maintain an integral
ontology of living beings in his holistic organismic conception of systems
hierarchy, I think the integrity of his conception breaks down when he
proposes that an organism displays a morphological hierarchy of parts,
as well as a physiological hierarchy of processes. The very concept of
systems implies that parts are defined by their relations and interactions,
viz., the processes that characterize the system. Splitting parts from
processes into separate hierarchies undermines the holistic ontology for
which he argues. Physiological processes do not have independent status
apart from the structures which display such functions. Inversely, struc-
tures are integrally related to function; a dead cell is not the same
integrated and unitary structure as a living cell. Structures and functions
are thus not separate entities but are integrally related, and any onto-
logical hierarchy must reflect such integral wholes.
Nevertheless, it is quite clear that the systems hierarchies proposed
by Weiss and Bertalanffy are constitutive, not aggregational, hierarch-
ies. No level can be considered as simply a sum or aggregate of the
parts. Organization does not arise from aggregation of parts or elements
but, rather, as Weiss succinctly points out, "omnis organisatio ex organ-
isatione" (Weiss, 1973, p. 94). The configuration or organization of the
constituent elements in a system makes possible novel properties of the
system which go beyond the properties of the subsystem. Grobstein
(1962, 1973) develops a good case for the phenomena of novel proper-
ties that appear in the higher levels of hierarchical order. He emphasizes
that such novel properties stem from the particular relationships be-
tween components of a particular level, as well as from the properties
of the components themselves. The phenomena of neogenesis, or the
emergence of properties of a structure which appears at a higher level,
depend on the context in which the structure is found. As the context
is changed or modified, new properties may emerge. This context is
multMevel. For Grobstein, an analysis of hierarchical levels requires a
 HIERARCHICALLY ORGANIZED STRUCTURES 119

description not only of a particular level itself, but also of the compo-
nents (subset) that occur within that level, as well as of the higher level
(superset) of which it is a component.

GENEALOGICAL AND ECOLOGICAL HIERARCHIES

Recently there has been a new interest in hierarchy theory with the
intent of providing a basis for establishing an evolutionary hierarchy
theory (Eldredge and Salthe, 1984; Vrba and Eldredge, 1984; Eldredge,
1985; Salthe, 1985). A basic concern in this recent development is to
establish a hierarchy theory which is 'ontologically based'. Whether this
claim for ontological status is justified depends upon a critical evaluation
of the proposed hierarchy theories in the light of our criteria for analyz-
ing and evaluating hierarchy theories. The two chief hierarchy theories
proposed by Eldredge and Salthe are referred to as the genealogical
hierarchy and the ecological hierarchy. The core of each of these hier-
archies is related to the distinction which Hull (1980) makes between
replicators and interactors.
The proposed genealogical hierarchy is a hierarchy of replicators,
viz., entities which produce new entities of like kind at the same level
(Eldredge and Salthe, 1984). It is a hierarchy of information transfer.
The 'glue' that unites the subparts of the individuals at each level is
'more-making' of the subparts of an entity (Eldredge, 1985). The speci-
fic levels of replicators are designated as codons, genes, organisms,
demes, species, and monophyletic taxa. The relationship between the
levels of the genealogical hierarchy is such that replication (more-
making) of entities at one level is required for the next level. In fact,
such replicated entities constitute the entities of the next level. Thus
organisms come about through replication of cells; demes come about
through replication of organisms, etc. Each level is thus considered to
be a class of entities (individuals) which themselves exist due to the
'more-making' of like kinds by (some of) the parts which compose
the sublevel. What we have here is simply a series of nested entities
characterized by the property of more-making.
The proposed ecological hierarchy is a hierarchy of interactors, viz.,
entities which are involved in "matter-energy transfer" (Eldredge, 1985,
p. 167). The 'glue' which lends cohesion to each level arises from the
energy exchanges in the sublevels. The specific levels in this hierarchy of
interactors are designated as enzymes (or molecules), cells, organisms,
120 UKO ZYLSTRA

populations, local ecosystems, biotic regions, and biosphere (Eldredge

and Salthe, 1984). Again, each of the units is considered to be a class
of individuals which is nested in the structure of the hierarchy.
Both genealogical and ecological hierarchies are process hierarchies.
They are characterized by intrinsic dynamic processes evident, respec-
tively, as information transfer through more-making and as matter-
energy transfer. Miller (1978) makes a similar distinction between sub-
systems which process matter-energy and those which process infor-
mation. For Miller, however, the subsystems belong to a single hierar-
chy of living systems, not to separate hierarchies. Furthermore,
Eldredge (1985) argues, it is through the interaction of these two pro-
cess hierarchies that evolution (selection) occurs, giving rise to the
evolutionary hierarchies of taxonomy and homology. This relationship
to evolutionary hierarchies becomes the key significance of the geneal-
ogical and ecological hierarchies. Because the genealogical and ecologi-
cal hierarchies are perceived to be ontologically based, evolutionary
hierarchies, which presumably result from their interactions, are also
assumed to have an ontological basis.
The presentation of these two hierarchy theories provides an illus-
tration of the difficulties which can arise when using ambiguous criteria
in developing a hierarchy theory. First of all, it is not entirely clear
what 'ontologically based' means. Does it mean that at each level the
hierarchy consists of actual, unitary, integral, wholes - individual be-
ings - or, does it mean that the entities of the higher levels of the
hierarchies are integrated wholes, constituted out of (rather than mere
aggregates of) lower level entities? The claim that these hierarchies are
ontologically based appears to be linked to the argument for integrated
and holistic character of the entities composing the hierarchies. If this
is correct, the compositional (and perhaps control?) character of these
hierarchies is of great biological importance. In practice, however, the
criterion used for individuality is whether the entities can be defined as
spatio-temporal entities. This is not a sufficient basis on which to sup-
port the claim that the entities at the upper levels are integrated compo-
sitional wholes.
A basic flaw in the genealogical and ecological hierarchies proposed
is that they are not clearly defined as aggregative or as constitutive.
This ambiguity is partly due to the confusion of inclusive hierarchies
with constitutive hierarchies. Eldredge (1985) gives an illustration of
 HIERARCHICALLY ORGANIZED STRUCTURES 121

inclusive hierarchies as organisms which are composed of organs, which

are composed of tissues, which are composed of cells, and so on. But,
as indicated above, this illustrates a constitutive hierarch~, not a merely
inclusive one. His emphasis on the individuality of the component
entities that make up the hierarchy suggests that these are aggregative
hierarchies. Although Eldredge states that we are "not to think of the
higher levels merely as aggregates of lower-level individuals" (Eld-
redge, 1985, p. 141), he proceeds to describe the individuals in each
level or rank as 'aggregates' or 'classes' of the entities or individuals of
the sublevels (see also Eldredge and Salthe, 1984).
To the extent that these hierarchies are aggregative, their significance
as hierarchies is undermined. Aggregative hierarchies contribute very
little beyond a basic ranking of entities. Ranking hierarchies trivialize
the concept of hierarchy (Pattee, 1973b); they are "just rather boring
ways of arranging things" (Grene, 1988, p. 9). The absence of control
or authority relationships between levels in aggregative hierarchies min-
imizes the ontological significance of such hierarchies. Perhaps aggre-
gative hierarchies should be designated as serial rankings rather than
hierarchies insofar as the element of rule (arch,) is non-essential. Doing
so would rid us of much of the confusion surrounding hierarchy theory.
Consequently, in view of the minimal importance of aggregative hier-
archies, the role proposed for the genealogical and ecological hierarch-
ies in establishing an evolutionary hierarchy is also diminished.
The alleged constitutive nature of these hierarchies raises a more
fundamental problem concerning their ontological basis. Constitutive
hierarchies are hierarchies of embedment in which the components of
each level interact in such a way that novel properties are expressed
at higher levels. The problem with the genealogical and ecological
hierarchies is that the individuals as defined for each level do not
actually interact as constituents at that level. They are seen as members
of nested levels (one level contained within another level) rather than
as structural components of levels. Consider the relationship of codons
and genes in the genealogical hierarchy. The more-making of codons
is presumably the 'glue' that holds together the genes of the next
level. But is that really so? Genes are formed or held together by the
association of various codons, not by codons making more of them-
selves. Likewise, chromosomes are held together by the association of
various genes, not by genes making more of themselves. This concept
122 UKO ZYLSTRA

of integration at the higher level affecting the interactions of entities

at the lower levels is precisely what is missing in the nested type of
genealogical hierarchy that Eldredge and Salthe propose.
The shortcomings of the genealogical and ecological hierarchies are
basically due to the question of individuality of the entities composing
the levels of the hierarchy. The issue is not so much the definition
given of individuals as spatio-temporal bounded entities but, rather, the
criteria for delimiting individuals in the respective hierarchies. The
criteria of more-making and matter-energy transfer are abstractions,
and they only describe a single aspect of the entities. More-making and
energy-transfer are undoubtedly functions of certain spatio-temporally
bounded things, but such functions do not exhaust the nature of the
entities which possess such functions. As a consequence, we are dealing
with a reduced ontology because the entities which are to provide the
ontological basis for the hierarchies have been stripped of full, integral
relationships. The criteria of more-making or energy-transfer are thus
not sufficient for establishing individuality. Another problem which
,arises from using a single abstract criterion for entities in a hierarchy
is that many other interactive components of the sublevels of individuals
are excluded from the hierarchy. The strength of a constitutive hierar-
chy lies in the very interaction of all of the components in each level.
A fundamental maxim in biology is that all components of a system
play a role in the system, and if a component is added, altered, or
missing, the system as an ontological whole will be affected. This notion
is absent in the proposed genealogical and ecological hierarchies; only
selected entities form the levels of the hierarchy. But entities always
exist in a context, and it would seem that an ontologically based hier-
archy would certainly incorporate the context in attempting to define
the entities, as well as the levels of the hierarchy.
If we evaluate these hierarchies in terms of control hierarchy theory,
we encounter further problems. In a control hierarchy, the upper level
provides boundary conditions for the sublevels. These boundary con-
ditions or constraints act through a form of "downward causation"
(Campbell, 1974, p. 180). But control or downward causation in hier-
archical structures is generally viewed as operating within the particular
hierarchical structure in question, viz., between levels in a hierarchy
(Pattee, 1973a). Eldredge and Salthe (1984; Eldredge, 1985; see also
Vrba and Eldredge, 1984), in contrast, seek to apply the concept of
downward causation as also extending from one hierarchy to another,
 HIERARCHICALLY ORGANIZED STRUCTURES 123

not just within a particular hierarchy. They argue that the genealogical
hierarchy guides the processes of the ecological hierarchy. This destroys
the very essence of a control hierarchy and it runs counter to the
concept of hierarchy in which authority relations extend from level to
sublevel within the hierarchy. If, indeed, we are able to recognize
phenomena of downward causation of entities in one hierarchy upon
entities within a different hierarchy, this would suggest that they are
all of a single, ontological hierarchy rather than of separate hierarchies.
The question of individuality also introduces a problem concerning
the meaning of levels in a hierarchy. How are levels defined? Tradition-
ally, organismic biologists have defined hierarchies in terms of levels
of organization or levels of systems (Bertalanffy, 1952, 1968; Weiss,
1969, 1973; Grobstein, 1962, 1969, 1973; Miller, 1978). This view is
given a twist by Hull when he attempts to correct a faulty assumption
of "our traditional way of organizing phenomena into a hierarchy of
genes, cells, organisms, kinship groups, populations, species, and eco-
systems or communities" (Hull, 1980, pl 311). This particular series,
however, does not completely resemble the traditional organizational
hierarchies. The level below the cells is usually designated as the mol-
ecular level, not the genetic (gene) level. Genes perhaps may be con-
sidered to be components of the cellular level, but they certainly do
not exhaust the level or system below the cell; cells contain components
besides genes. Although the levels are often designated by the noun
forms 'molecules', 'cells', 'organisms', etc., they are generally interpre-
ted in the adjectival forms, 'molecular', 'cellular', 'organismal', etc.
But Hull seeks to emphasize the individuality of the levels for the
reason that only individuals can be selected. In doing so, Hull shifts
the meaning of hierarchy to deal, not with levels of organization, but
with levels of individuals. When Hull then proceeds to define such
individuals as replicators or as interactors, he is limiting n o t only the
range of individuals which are included in the hierarchy but also the
grounds for inclusion of individuals included to a single aspect of their
functioning. Hull argues that "selection can only act on spatio-tempor-
ally localized entities" (viz., individuals), but "these entities must be
cohesive wholes" (Hull, 1980, p. 314). However, in his definition of
entities as replicators or as interactors Hull destroys their wholeness by
defining such entities in terms of a single function. This limits the
conceptual recognition of such entities as cohesive structural or func-
tional wholes.
124 UKO ZYLSTRA

A similar problem is present in the discussion by L. Buss (1987)

concerning the evolution of hierarchical organization. Buss argues that
"life is hierarchically organized, with species composed of populations,
populations of individuals, individuals of cells, cells of organelles, or-
ganelles of chromosomes, and chromosomes of genes" (Buss, 1987, p.
183). This hierarchy is also one of levels of individuals rather than
levels of organization. This becomes obvious when we recognize that
cells are not composed of only organelles, nor are organelles composed
of genomes. Even the eukaryotic chromosomes contain both structural
and regulatory proteins in addition to DNA (genes). This type of
hierarchy has limitations because it is designed to establish a hierarchy
of units of selection rather than of levels of organization.
Genealogical and ecological hierarchies thus appear to have a weak
ontological basis. It becomes questionable whether theories that deal
with such hierarchies can adequately establish an evolutionary hierar-
chy. In fact, one wonders whether the desire to establish an ontologi-
cally based evolutionary hierarchy has become the driving force in
making these proposals for the genealogical and ecological hierarchies.
It often appears that these two hierarchies are constructed so as to fit
the conditions of a preconceived evolutionary hierarchy rather than to
reflect the actual nature of living things.

PARTS AND WHOLE

So far, our discussion of hierarchy theory has highlighted the need to

establish hierarchies upon a sound ontological basis. This implies that
the entities that compose the levels in the hierarchy are unitary beings,
that they are real, integrated entities. Furthermore, to avoid triviality,
hierarchy theories should articulate a well-developed theory of the
relations between the levels in a hierarchy. Are such relationships
to be seen as part/whole relationships, or, are there other forms of
relationships involved as well? Of particular concern is the definition
of both parts and wholes and the individuality of parts. The terms 'parts'
and 'wholes' demand careful definition if they are to be serviceable in
the context of hierarchy theory.
A basic theme of organismic biology, 'the whole is greater than the
sum of the parts', implies inherent distinctions between the concept of
parts and that of pieces, portions, or fragments. If a plant or animal
body were to be cut into numerous equal portions by weight, such
 HIERARCHICALLY ORGANIZED STRUCTURES 125

portions cannot really be considered as parts in the sense that the whole
is greater than the sum of the parts. As Hart (1984) argues, a genuine
whole is an integral totality of parts, not of fragments or pieces. Leaves
or branches may be considered as legitimate parts of a tree, and legs
or ears may be considered as legitimate parts of an animal body. A
part has significance only in terms of its relation to other parts and to
the whole of which it is a part. A true part is not independent from
the whole but, rather, it is defined by the whole of which it is a part
(Hart, 1984). This relation is entirely absent, for example, in a heap
of bricks. There is no meaningful interaction or relation between the
bricks which make up the pile. The pile is simply a sum of the individual
bricks and is more appropriately designated as an aggregate rather than
a whole (Laszlo, 1972). But if parts lack independence and are only to
be defined in terms of the whole, can they then possess individuality?
It would seem not. Individuality implies a measure of structural and
functional independence in relation to other structures; true parts lack
that independence. This poses a dilemma with regard to a theory of
levels of organization. Can a whole, as a structure of individuality, in
turn be itself a part of another whole of a higher level of Organization?
If a whole is a genuine part of another whole, then the latter would
define the nature and character of the whole as a part. The whole as a
part would then lose its sovereignty and internal integrity as a whole.
An analogy may be useful in our attempt to clarify the distinction
between part and whole. A home, as a whole, is composed of a number
of parts, such as the walls, ceilings, rooms, etc., which constitute the
structure of the home. A home also contains furniture which contributes
to the individuality of the structure designated as home. It is obvious
that a wall and a table are not to be considered as parts in the same
sense. The relation of the wall to the home is quite different from the
relation of a table or bed to the home. Furniture pieces are themselves
integral wholes consisting of parts (e.g., the legs, seat and back of a
chair). A wall, in contrast, is not really a genuine whole; it has no
meaningful existence apart from the whole to which it is related. It is
the concept of the home as a whole which defines the meaning and
character of walls or even of rooms which exist as parts of the home.
But that is not so for the pieces of furniture in the home; they possess
meaning (functional and structural meaning) as entities apart from the
home. We can recognize two fundamental relations in this analogy:
that of part to whole (wall to home) and that of whole to whole
 \

126 VKO ZYLSTRA

(furniture to home). Such fundamental differences in types of relations

are also present with living things. Hierarchy theories which claim to
be ontologically based should seek to incorporate these distinctions.

THEORY OF ENKAPSIS

H. Dooyeweerd 1 (1957) proposes a novel theory to account for these

distinct relationships in his theory of enkaptic structural wholes as it is
applied to living things. Dooyeweerd borrows the term 'enkapsis' from
the anatomist Heidenhain, who used the term to describe the relation
between the separate organs (such as the liver and heart) and the
organism as a whole. Dooyeweerd rejects this application of the term
enkapsis by Heidenhain, though he borrows the idea behind the term.
The term 'encapsulating' (derived from the same root word) has also
been used by Weiss (1973) and Koestler (1969) to denote the relation-
ship of structures in a hierarchical organization.
Dooyeweerd distinguishes between part/whole relationships and en-
kaptic relationships. He wishes to limit the concept of part/whole re-
lationships to refer to those relations in which the parts are qualified
by the same function as that which qualifies the whole. A part is
conceived as being qualified by the structure of the whole and is entirely
dependent on the whole. The possibility of organs being transplanted
does not negate this dependency. Parts of wholes do not have their
own internal destination (teleonomy), and for this reason the whole
cannot be grasped in terms of its parts. Thus, for Dooyeweerd, organs
such as kidneys, liver, and heart are to be considered as parts of the
whole animal. Although such organs may possess a relative degree of
autonomy, their nature is dependent upon the total structure of which
they are a part. Wings, legs and other organs can only live as 'parts'
of the ensemble (Mayr, 1982). Their functional significance is defined
and determined by the totality structure (whole) of which they are a
part. As parts they lack sovereignty as an independent structure of
individuality.
Dooyeweerd defines an enkaptic relation as one which involves a
relation between different types of individuality structures (wholes) in
the formation of an enkaptic structural whole. We are dealing here
with a whole/whole relationship rather than a part/whole relationship.
A cell is a good example of such an enkaptic structural whole. The
molecules in a cell are enkaptically related to the living cell. The
 HIERARCHICALLY ORGANIZED STRUCTURES 127

molecules are thing-structures with an internal teleonomy independent

of that of the cell. The cellular molecules are not to be conceived as
parts of the cell since they are not living components of the cell.
Even within the confines of the cell, molecules continue to function
subjectively as physico-chemical entities, not as biotic entities. They
remain subject to physico-chemical ordering principles, not to biotic
ordering principles. Their sovereignty as physico-chemical entities is
not affected by being enclosed (encapsulated) in the cell. The only
structures to be considered as parts of the cell are the cell organelles,
such as the nucleus, endoplasmic reticulum, plasma membrane, mito-
chondria, chloroplasts, and Golgi complex, which are biotically quali-
fied; they are subject to biotic ordering principles. As living components
of the cell, they do not have an independent teleonomy apart from that
of the whole cell. Such distinctions nearly always involve borderline
cases. The eukaryotic chromosome is a case in point. As mentioned
earlier, the DNA molecule itself is not a living structure. Whether
chromosomes are to be considered as living structures is dependent on
the degree of sovereignty which they have in the nucleus.
The relationship between the cell and the cellular molecules is one
in which the cell embraces or encapsulates the cellular molecules to
form the enkaptic structural whole which we perceive as the living cell.
This embracing of cellular molecules within the living cell does not
attribute subjective2 life functions (viz., functions which are subject to
biotic ordering principles) to the molecules; nor does the enkaptic
relation obliterate the physico-chemical subject functions of the mol-
ecules. In fact the sovereignty of the molecules as physico-chemical
structures is maintained in the enkaptic relationship. The cellular mol-
ecules in the enkaptic relationship serve the function of the living cell;
they do not determine the nature or teleonomy of the cell itself. The
physico-chemical functions of the molecular structures of the cell are
opened up under the guidance of the subjective biotic functions of the
cell in such a way that molecular interactions occur which do not
normally occur outside of such an enkaptic relationship with the cell
(see also the discussion on the unity of the cell by Weiss (1963)). The
enkaptic relationship can be conceived as an expression of boundary
conditions which provide constraints for the enclosed structures. Such
constraints make possible particular patterns of molecular functions
that are integrated with the biotic functions of the cell as a structural
whole. This may be illustrated by considering the example of protein
128 UKO ZYLSTRA

structure and function. The significance of cellular proteins is only

realized within the context of the living cell. A particular protein, which
may be a structural component of a cellular membrane, has a specific
function in the cell by virtue of its being a component in a particular
membrane. Its physico-chemical function is under the guidance of the
biotic function of the organelle which has opened up the potential
physico-chemical functions of the protein in the service of the organelle
and thus of the living organism. This may be reflected in the position
and orientation of the protein in the membrane, as well as in the
interaction between the protein and the neighboring molecules. In
general, cellular proteins function in their specific way only within the
ordered environment of the cell.
A similar enkaptic relationship is evident in an analysis of the struc-
ture and function of the DNA molecules in the cell. As a molecule,
DNA is subject to physico-chemical ordering principles, not to biotic
ordering principles. The physico-chemical ordering principles determine
the DNA's structure, physical properties, and even the various ways in
which it can interact with other molecules or cell structures, but they
do not determine the specific way in which the DNA molecule is
integrated in the cell as a structural whole. As a molecular structural
whole, DNA is bound enkaptically to the cell which is a living structural
whole. That enkaptic relationship is critical for understanding the sig-
nificance of DNA as a molecule which contains genetic information.
As Polanyi (1968) argues, the particular information content in the
sequence of bases of the DNA molecules is not specified by physico-
chemical laws. This notion is affirmed by Pattee in his statement that
"molecules have no inherent message, but that message behavior em-
erges only in the new context of a larger set of constraints" (Pattee,
1973b, p. 134).
The theory of enkapsis suggests that the base sequence of DNA
molecules possesses potential information only as a consequence of
being enkaptically bound to the living cell. In the living cell the func-
tioning of the DNA is ordered in such a way that it can possess infor-
mation as a template for protein structure. The DNA itself does not
direct the protein synthesis activity of the cell. Nor does DNA direct
the structural and functional activity of the proteins. Such activity is
under the direction of the cell as a whole; DNA functions primarily as
an indirect template for protein synthesis activity. Apart from the en-
 HIERARCHICALLY ORGANIZED STRUCTURES 129

kaptic relation in which the cell organism embraces both the DNA and
regulatory and enzymatic protein molecules, the DNA would not serve
any meaningful biotic function. Although processes of protein synthesis
or other cellular activities are physico-chemical functions of the living
cell, they are subject to the biotic guiding function of the cell. They
are not determined by the physico-chemical properties of the molecular
composition of the cell since such functions lie outside of the internal
structure (teleonomy) of the molecules. Thus the enkaptic relation is
not determined by the ordering principles for the embraced structures
such as DNA but, rather, by the ordering principles for the thing (the
living cell) in which the embraced thing (DNA or proteins) functions
enkaptically.
Dooyeweerd also introduces the term 'form-totality' to designate an
enkaptic structural whole in which two or more individuality structures
(wholes) are interwoven or united in a thing of our experience. A
molecule is an example of such a form-totality. A molecule is an individ-
uality structure which embraces two or more atoms in such a way that
the atoms retain their own individuality and sovereignty. In a glucose
molecule, for example, the carbon, oxygen, and hydrogen atoms retain
their individuality structures as carbon, oxygen, and hydrogen atoms.
In this enkaptic relation, the atoms become serviceable to the glucose
molecule as a form-totality which expresses novel functions only real-
ized by the enkaptic structural whole. The carbon, oxygen, and hyd-
rogen atoms do not become parts of glucose as in a part/whole relation-
ship.
The multicellular bodies of plants and animals are also form-totalities
in our experience. They are constituted of both cell bodies and non-
cellular molecular structures of individuality which are interwoven into
a unified whole, the enkaptic form-totality. The enkaptic relationship
between the body and its component thing-structures serves to make
possible functions of the component structures which could not be
realized apart from the enkaptic relationship. Examples of the non-
cellular molecular structures in animals are collagen, blood plasma, and
extracellular ground substance. A snail, for example, is a form-totality
which embraces both the shell as a physico-chemically qualified individ-
uality structure, the cells as biotically qualified individuality structures,
and the other physico-chemically qualified extracellular structures.
Plants are also enkaptic structural wholes. The cell walls are physico-
130 UKO ZYLSTRA

chemically qualified form products of the cells. The cell walls are in-
terwoven with the cell bodies in an enkaptic relationship with the plant
as a form-totality.
We can perhaps now discuss the relation of organs to the whole
body. Is it an enkaptic relation, as Heidenhain suggests, or a part-
whole relationship? The key to such a distinction involves the question
of an independent teleonomy. There does appear to be a distinction
between cells and organs in that cells display a certain degree of sover-
eignty as a fundamental unit of living things. Cells can be removed,
isolated, and grown in culture with varying degrees of integration
among cells. In some cases they can even develop into a new individual,
multicellular structure. Organs, though they may be preserved for some
length of time outside of the body, are not able to function as indepen-
dent structures apart from the body as a whole. They do not appear to
possess an independent teleonomy, but seem to be entirely dependent
on the whole body which determines the structure and functions of the
organ parts. On this basis we conclude that the organ-body relation is
a part/whole relation, not an enkaptic relation.
The theory of enkapsis thus provides us with an alternative insight
into the relation between the levels of organization of living things.
The definition of the levels is dependent upon whether the levels are
constituted of integral structural wholes or of parts. If we seek an
ontologically based hierarchy, then the levels of the hierarchy should
consist of integrated wholes which have at least the degree of indepen-
dent teleonomy possessed by cells. I suggest that such a hierarchy
theory be designated as a hierarchy of enkaptic structural wholes as
opposed to a hierarchy of part/whole relationships. A preliminary
sketch of such a hierarchy could contain the following levels:
biosphere
ecosystems
populations (demes)
individual organisms
cell and cellular form products
molecules
atoms
The levels above that of individual organisms are tentative. They re-
quire a more thorough analysis with regard to the enkaptic structural
wholes at each level and the nature of the enkaptic relations that may
 HIERARCHICALLY ORGANIZED STRUCTURES 131

exist between the levels. Such an analysis is beyond the scope of this
paper.
This paper is an attempt to provide greater clarity to our under-
standing of the hierarchical nature of living things. Two fundamental
problems arise in the application of hierarchy theory to living things.
One problem concerns the meaning of the concept of hierarchy as used
in a variety of applications. This requires a typology of hierarchies as
argued for and initially developed by Grene (1987). But, as suggested
in this paper, it may also require limiting the use of the term hierarchy
to relationships or systems in which some form of rule or authority
relation is present. A second problem concerns the meaning of parts
and wholes and the relationship between parts and wholes. The present
introduction of the theory of enkapsis as distinct from a theory of
part/whole relationships may serve to elucidate this problem. At the
very least, it is hoped that it will stimulate and guide further interest
and discussion of hierarchy theory and the nature of living things.

NOTES

1 H. Dooyeweerd was a prominent Dutch legal philosopher who also made a major
contribution, through a large number of publications, in the area of systematic philosophy.
2 The term 'subjective' is used here in the sense of be!ng the subject of the function as
opposed to the object of the function. Thus, in this case, molecules can only serve as
objects of biotic functions not as subjects of biotic functions, whereas cells function as
subjects of both biotic and physico-chemical functions.

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Dept. of Biology
Calvin College
3201 Burton St., S.E.
Grand Rapids, MI 49546
U.S.A.