Thinking of Biology

Biological organization A new look at

an old problem
T he cell theory, or cell doc- ria of self-control and then viewing wrong one-free-cell formation-be-
trine, which states that all or- them as existing at different levels of cause he failed to consider all avail-
ganisms are composed of simi- biological organization. Levels of able theoretical ideas. The correct

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lar units of organization, called cells, control is the subject of hierarchical interpretation of cell formation by
was first enunciated in 1839 and has theory; hence, organisms can be bet- division was finally promoted by
remained one of the foundations of ter described by a hierarchical than a others and formally enunciated in
modern biology. This idea predates mainly structural perspective. In this RudolphVirchow'spowerfuldictum
other great paradigms in biology, article, I explain the relationship Omnis cellula e cellula ("All cells
such as Darwin's theory of evolution between the cell theory and hierar- only arise from pre-existing cells";
(1859), the rediscovery of Mendel's chical theory. I begin by developing Wilson 1896).
laws of inheritance (1900), and the both theories and then try to recon- The cell doctrine reached its
establishment of comparative bio- cile them. present-day eminence in 1896 with
chemistry (1940). Although ultra- the publication of E. B. Wilson's The
structure research and molecular bi- The cell theory-organismal Cell in Development and Heredity,
ology have added much to the cell which was an accumulation of what
theory controversy was known about the roles of cells in
theory, and it has retained its emi-
nent status in biology, the cell theory In 1838, Theodor Schwann and Mat- embryology and chromosomal be-
faces two ongoing problems. thias Schleiden were enjoying after- havior.Nevertheless,some biologists
First, the cell theory began as, and dinner coffee and talking about their held stubbornlyto the idea that nerve
to a great extent remains, a struc- studies on cells. According to his organizationremainedan important
tural idea. This structural view, biographer (Fredericq 1884), when exception to the cell theory because
which is found in most textbooks, Schwann heard Schleiden describe some fibers appeared to come from
describes the components of a cell plant cells with nuclei, he was struck nonliving substances. But even this
and their fate in cell reproduction. by the similarity of plant cells to cells final resistance to the cell theory
Today, however, biology focuses on he had found in animal tissues. The succumbed with Ross Harrison's
DNA and its informational features, two scientists went immediately to convincingdemonstration(Harrison
and the description of a cell needs to Schwann's lab to look at his slides. 1907) that neurons in culture arise
catch up with this contemporary view Schwann published his book on ani- only from other nerve cells.
of life. The cell is, and needs to be mal and plant cells (Schwann 1839) The cell is usually investigated by
described as, a unit of self-control. the next year, a treatise devoid of three approaches: characterizing
That is, the description of a cell needs acknowledgments of anyone else's structural components, identifying
to incorporate ideas about how in- contribution, including that of the biochemical activity of the com-
formation is converted to structure. Schleiden (1838). He did, however, ponents, and noting how the compo-
The second problem with the cell summarize his findings into three nents multiply and segregate during
theory is historical: It first gained famous conclusions about cells: cell division. In 1952, a fourth ap-
prominence along with the organismal proach to the study of the cell began
theory, which is the idea that the * The cell is the unit of structure, when the mathematician John von
organism has its own structural fea- physiology, and organization. Neumann developed a theory of self-
tures apart from those of cells. The * The cell retains a dual existence as reproducing automata, in which he
ensuing debate over which of the a distinct entity and a building block viewed cells as self-regulating ma-
two structural theories better ex- in the construction of organisms. chines. Although others had previ-
plains biological organization, espe- * Cells form by free-cell formation, ously described cells as machines, it
cially in protists, has never been re- similar to the formation of crystals. was von Neumann who emphasized
solved sufficiently by structural the importance of internal control
criteria. The first two statements are still ac- and who provided the formalism
A resolution to both problems can ceptable, although the third is clearly behindthis feature.He proposedthat
be found by restating the cell and wrong. Before Schwann's publica- such a machine would have three
organismal theories in terms of crite- tion, there were two rival theories on components: an assembler, to make
how cells form-one claiming free- another automaton; an information
cell formation and the other claim- tape, to direct the activity of the
by Robert W. Korn ing cell division. Schwann picked the assembler; and a tape recorder, to

January 1999 51

University of California Press

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make a new copy of the tape. based on errors in observations. Sec- activity such as self-reproduction
Because the cell is the smallest ond, many adult animal tissues ap- might come from a set of similar
unit of reproduction, the question is peared to have no cell boundaries, a units working as a "team." von
then: Where are these three compo- conclusion that also reflected poor Neumann took up this idea and redi-
nents in a cell? Certain cell organelles observation. Third, plant cells are rected his energy to what he called
can be recognized as von Neumann's connected by plasmodesmata and "cellular automata." He defined a
components if they are viewed from animal cells have gap junctions; these cellular automaton as an entity that
a generic perspective. The cytosol, seemed to create a continuum of pro- can assume various states as speci-
with its batteries of enzymes for syn- toplasm instead of separate cells, an fied by its own encoded information.
thesis and breakdown of many error in interpretation because cells Although the cellular automata of a
biomolecules, functions as the as- can still be individuals even though group can have copies of the same
sembler; the chromosomes are, col- they are connected. Fourth, some program, they assume different
lectively, the information tape; and animal parts cannot be regenerated, states, depending on their status in
the set of DNA polymerases consti- as Wilhelm Roux found when the the group. Integrated group behav-

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tutes the tape recorder. To complete destruction of one cell of a two- ior comes from the close interaction
these correspondences, the scrapyard celled frog embryo led to a partial between individual automata, and
in which the automaton lives be- embryo (Roux 1888). This result because individual actions change,
comes the nutrient-rich external en- suggested to some embryologists that new and interesting behavior pat-
vironment of the cell, which includes parts, rather than cells, constitute an terns arise, including life cycles.
at least inorganic materials. von organism-an erroneous assumption Around 1970, John Conway's game
Neumann had a problem working because regulative and mosaic pat- Life (Gardner 1970), which dealt
out the logic behind the system, par- terns of development were just be- with a special brand of cellular au-
ticularly with the type of material ginning to be recognized. Moreover, tomata, became the rage at com-
needed in the scrapyard, and so he in 1866, Ernst Haeckel, a dominant puter centers throughout the world.
abandoned the problem. Only later figure in science, claimed that a plant This little game, carried out by com-
were his writings salvaged and ed- begins as a cell that simply enlarges puter or by hand, provided a visual
ited by Arthur Burks (von Neumann and forms compartments by plate representation of how complex group
1966). formation. Adding to the difficulties behavior could emerge from a simple
Clearly, von Neumann was aware for the organismal theory was its set of rules and a few individual
that the biological cell was the only association with vitalism, the phi- automata.
example of self-reproduction known, losophy that immaterial forces gov- A multicellular organism is simi-
and so it probably served to guide ern biological activities, which was lar to a group of cellular automata in
him in constructing his theory. As a espoused by such eminent thinkers three ways. First, each automaton
proliferating system, a cell must in- as General Jan Smuts and Alfred has the same program, just as an
clude at least some central DNA (as North Whitehead, among others organism is composed of cells with
in prokaryotes) or a nucleus (as in (Beck 1957). the same genotype. Second, an au-
eukaryotes), and some cytoplasm To most biologists of the late nine- tomaton can assume any one of a
with assorted enzymes and mono- teenth century, the organismal theory number of different states encoded
meric building blocks. offered little for understanding how in the program, just as a cell can
In contrast to the smooth devel- life is organized. There were, how- differentiate into any one of many
opment of the cell theory over the ever, some positive contributions cell types that are encoded in the
past 150 years, the history of the from the organismal theory, such as genotype. Third, specific ensembles
organismal theory has been fraught the construction of a fate map for the of automata can pass through cyclic
with inaccuracies and misunder- chicken embryo by Wilhelm His. In patterns of structure and behavior
standings. The organismal theory this approach (His 1874), various that are parallel to an organism's life
proposed that a living thing, or or- cytoplasmic regions of the egg were cycle. Although von Neumann's idea
ganism, has a complex organization viewed as destined to become spe- of self-reproduction supported the
and is often, but not necessarily, cific structures of the adult, indepen- cell theory, his concept of cellular
composed of cells. When Carl von dent of any cellular organization. automata gave new meaning to the
Siebolt found, in 1845, that protozo- Similarly, Kaplan and Hagemann organismal doctrine in that it showed
ans were similar to cells of multicel- (1991) cited a number of cases in how a small group of cells with a
lular organisms, they were called which plant form is independent of simple common program could gen-
unicellular organisms by some (e.g., cell size, shape, and orientation; they erate complex behavior.
von Siebolt himself and Max Ver- concluded that at least part of an Just as a biological cell is but one
worn) and noncellular organisms by organism's morphology must come example of the self-reproducing phe-
others (e.g., Thomas Henry Huxley, from something other than the build- nomenon, so the multicellular or-
Charles Otis Whitman, Clifford ing blocks of cells. ganism is but one type of cellular
Dobell). Even metazoans were, for After contributing significantly to automata collective. A cell is charac-
several reasons, considered noncel- the cell theory, von Neumann pro- terized by a set of nucleo-cytoplas-
lular (Singer 1959). First, some early posed a new idea about organisms. mic relationships (involving many
chordate embryos appeared to have His friend, the mathematician Stanis- different components, such as chro-
a free-nucleate stage-a "feature" law Ulam, suggested that complex mosomes and enzymes), whereas a

52 BioScience Vol. 49 No. 1

multicellular organism is composed
of many similar components-cells. fertilized egg
The distinction between the cell and
organismal theories is that, accord-
ing to the cell theory, cells organize organism organism
into an all-prevailing pattern of the
individual organism-that is, cells
contribute actively to making an or-
ganism. According to the organismal
theory, by contrast, cells passively
acquire parts of the prevailing pat-
tern from the embryo and hence have
only a custodial role in organismal h
pattern (Figure 1).

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Hierarchicaltheory Cell doctrine Organismal doctrine
The term "hierarchy" was coined by
Pseudo-Dionysius in the sixth cen- Figure1. The differencebetweenthe cell and organismaldoctrines.The cell doctrine
tury to formalize the relationships considersa cell as an identifiablesystemwith processesfor supportingitself as well as
between levels of Christian orders, processesthat contributeto an intercellularpattern.The organismaldoctrineconsiders
or sacred rulers (Osborn 1967). Al- cells as unitscontributingdifferentfeaturesto an overallorganismalpatternby which
though the idea of a hierarchy began cells are sustained.
as a description of a sacred chain of
authority, it was eventually extended scheme of a nested classification of inclusive view of hierarchies is the
to other human organizations, such cells, tissues, and organs (e.g., Table emphasis on the more passive, mem-
as government and armies. It also 1) is now commonly included not bership aspects of organizations.
came to mean a type of classification only in textbooks (e.g., Raven and I have modified Pattee's idea by
in which different groups are included Johnson 1996) but also in most schol- emphasizing that in hierarchies, con-
under a common heading. This in- arly descriptions of biological sys- straints are nonreciprocal (Korn
clusive approach has appeared spo- tems (Weiss 1971, Guttman 1976, 1994). For example, the constraint
radically in biology. August Comte MacMahon et al. 1978, Koestler of gravitational pull is shared equally
in 1854 saw a "hierarchy of sci- 1979, Lieberman and Vrba 1995). by the sun and the earth, but the pull
ences," Virchow in 1858 noted a It was Howard Pattee, a physicist, influences the small mass of the earth
"hierarchy of pathological condi- who pointed out that hierarchies can more than the large mass of the sun.
tions," and Hughling Jackson in 1881 be either physical (meaning, in fact, Therefore, the earth revolves around
described a "hierarchy of nervous physical, chemical, or biological) or the sun. Such nonreciprocal con-
centres in man" (Whyte 1969). descriptive (cognitive) and that it is straints can be easily identified in vari-
Joseph Woodger first described the physical hierarchies that are ous aspects of plant organization (Korn
biological organization as "hierar- formed by members at a higher level 1986, 1994, in press). Table 1 shows
chical," using a taxonomic approach constraining members at a lower level more levels of organization than are
(Woodger 1929). First, he classified (Pattee 1969). Because energy is re- traditionally cited, and the emergence
the known biological levels of cells, quired to forge and, often, to main- of these levels during evolution can
tissues, and organs; he then exam- tain a physical constraint, physical be understood by the insertion of
ined how these levels are structurally hierarchies are dynamic systems. new features accompanied by new
related. As part of an organism, a Even Pseudo-Dionysius's original constraints over these new features.
cell has many relationships to other idea that an organization is based on This dynamic view of a hierarchy is
cells. These relationships no longer authority implies constraint, because illustrated in Figure 2.
exist when a cell is isolated from its authority dictates constrained behav- Only on rare occasions does the
normal neighbors; thus, it will be- ior. The problem with the traditional, idea of nonreciprocal constraint ap-
have differently. To understand cells,
Table 1. Traditional classificational view of the hierarchyof higher organisms.a
Woodger insisted, they must be stud-
ied either in their normal environ- Level Plant Animal
ment or in artificially reconstructed
conditions. Because Roux's half- Organism Dandelion Mouse
Organ system Shoot Integumentary
embryo of one cell has a neighbor Stem Skin
Organ
missing, it will not make exactly half Tissue complexb Vein Dermis
an adult. This view of the biological Tissue Phloem Epidermis
hierarchy of cells, tissues, organs, Cell complexb Loading system Hair follicle
and organisms has been passed on Cell Companion cell Keratinocyte
through biology textbooks, starting aLower levels (from atom to organelle) are not included.
with Simpson and Beck (1967). The bThese levels are not typically included in the traditional classificational view.

January 1999 53
 ent forms when a substance diffuses
organism:dandeliond from its site of production to sink
organs m: constraint by gradient (control) sites; consequently, the substance is
organ system: shoot found at high concentrations at the
>i~~~ constraint by support source and at progressively lower
organ: stem concentrations farther from the
source. Responses to the substance
organ: leaf constraint by support
by cell or region vary with the con-
organ: leaf centration, such that one type of cell
constraint by enclosure may arise near the source, another
tissue complex: vei T type at the distal end, and other
tissue:phloem
segmeconstraint
by enclosure types in between. This gradient is
tissue: phloem segment _ not a cellular feature because it can
extend either from one end of a cell
cell cmplex:
constraint by bonding (cluster)

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to another or extracellularly, from
cell complex: phloem one pole in a group of cells to an-
loading unit other pole (Wolpert 1969).
In Fucus, the organismal gradient
cloadin el n constraint by bonding (cluster) constrains the fertilized egg as a cell
cell: sieve element d
in dictating, first, how it will divide
Figure2. A dynamicview of a hierarchyby physicalconstraints.A constraintdoes not to form two dissimilar daughter cells
act on a levelper se but on a specificstructureat a particularlevel. Someconstraintsare and, later, that the lower daughter
of the traditionalenclosureor nestedtype (e.g., a leaf containsa vein or a vein contains cell will no longer divide. Therefore,
a set of phloem cells), whereas other constraintsare nontraditional.Nontraditional the answer to the question of whether
constraints include bonding between similar members, as exemplified by phloem a fertilized egg is a cell or an organ-
complexesforminga tissue, supportby a stemof a leaf, and controlthrougha gradient ism is that it is both, making the cell
of a regulatorysubstance. analogous to the situation of the
one-room schoolteacher in the Old
pear to be contradicted. Konrad many old ideas. For example, the West, who was principal, teacher,
Lorenz included in his lectures (Eigen fertilized egg of the brown alga Fu- and custodian, all in one person.
and Winkler 1981) the example of a cus is both a cell and an organism. Constraints need not be of Woodger's
pecking order in which goose A domi- Nucleo-cytoplasmic interrelation- inclusiveness, as in a set of Russian
nates goose B, which dominates goose ships make it a cell, and a persistent dolls, but rather may act on indi-
C, which dominates goose A (a polarity of activity generated from a viduals of like makeup with different
"tangled hierarchy" ).This apparent gradient during early development status, for example, soldiers of dif-
hierarchical contradiction is resolved make it an organism (Figure 3; Jaffe ferent ranks in the same army. If
by noting that the hierarchy exists in 1969). hierarchical theory does no more than
two alternating forms, depending on Indeed, gradients such as that in explain how a fertilized egg can be
the type of fighting strategies used. Fucus can play a critical role in the both a cell and an organism, it has
The idea of a nonreciprocal con- development and maintenance of an earned a legitimate place in biology.
straint presents a new way to view individual cell or organism. A gradi-
Structural hierarchies
in biology
An organismal gradient, as is found
in the protist Fucus, can also be iden-
tified in many plants and animals.
Fern sporlings appear to have a gradi-
ent that gives an apical cell-rhizoidal
dipole configuration (Korn 1993).
Hydra has a mouth-foot double gra-
dient of chemically known substances
(reviewed by Kemmer 1984). The
zygote of flowering plants divides
unequally to form a bottom cell that
later forms the radicle (embryonic
(*) cell feature organismal feature root) and an upper cell that is fated
to form the shoot and some of the
Figure3. An individualwith both cellularand hierarchicalfeatures,exemplifiedby the radicle (Juirgens1995). This polarity
fertilizedFucusegg.Theeggis a cellwith a nucleusandcytoplasmas well as an organism may persist as a double gradient in
with a gradient.Whereasthe cell featuresmultiplyas the embryoforms,the organismal which auxin passes from shoots
featureof a single gradientpersistsinto and throughoutthe embryo. (stems and associated leaves) to roots,

54 BioScience Vol. 49 No. 1

and cytokinin moves up from roots Figure 4. Levels
to shoots. The persistence of this of organizationin
double gradient can be interpreted unicellularorgan-
as an organismal control constraint isms. org, organ-
that is important in maintaining in- ism;os, organsys-
tem; o, organ; t,
determinategrowth and dictatingre- tissue;c, cell.
generationafterinjury.In insecteggs,
gradientsof RNA and protein estab-
lish the three body regions of head, plasm,andmost
thorax, and abdomen and subse- likely the orga-
quently the 14 segments (Nusslein- nismal feature
Volhard 1991). In higher animals, of a head-to- d rhi
an egg gradient is often visible by foot gradient, rh
yolk concentrationor in the location as demonstrat- Acetu
Vorticella
of the nucleus that is probably re- ed by regenera-

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placed by special regions for pri- tion studies in
mary, secondary,and tertiaryinduc- other ciliates (Stevens 1903, Sonne- Genes are responsible for control
tions. The adult nervous system born 1963). Faure-Fremiet (1954) of processesat the cell andorganismal
eventually replaces the inductive consideredboth unicellularand mul- levels. Somegenes specify regulatory
fields of the embryo as the means of ticellular organisms to have a gradi- proteins for metabolism and cell di-
organismal maintenance. ent in their cortical cytoplasm and vision, whereas others specify the
Organismalgradients can also be suggested that this gradient deter- production of an intercellulargradi-
found in unicellularorganisms, such mines cytoplasmic specialization in ent. All genes are nucleotide se-
as the siphonaceous algae and ciliate unicells and cell types in metazoa. quences, regardlessof the hierarchi-
protozoans. The marine alga Ac- Other unicellularorganismscan also cal level at which their products
etabularia, or mermaid's umbrella, be characterized by an organismal function. However, genes for cell
has the cell featureof a single nucleus gradient.In bacteria,polarity occurs function interact with one another
and some cytoplasm (Figure 4). It in wall synthesis, spore formation, indirectlythrough protein products,
also has a stalk and rhizoid system and, sometimes, motility. Simple al- whereas genes for organismal activ-
and, later, a cap system, each of gae, fungi, and protozoa also ex- ity interactwith regions of the gradi-
which can be regenerated,indicating pressasymmetricorganizationalfea- ent, also indirectly, through their
the presenceof a regenerationgradi- tures similar to those in bacteria. protein products.
ent as an organismalfeature. Certain Multicellular organisms can also The presence of an intercellular
structures in Acetabularia serve to be given a new hierarchicaldescrip- organismalgradientraises two inter-
constrain other structures that are tion, not by the traditional approach estingdevelopmentalquestions.First,
comparable to organs in higher of levels but by the identification of how does the gradientbecome estab-
plants. As examples, the rhizoid descendent constraints. The organ- lished? This question is another way
serves for anchorage, analogous to ism is the body and is representedat of asking when an individual organ-
the root of higher plants; the stalk some times by a gradient. Organs, ism begins. Individualsemergewhen
functions in support, as does the such as leaves in plants and bones in eggs and spores (and, experimen-
stem;and the cap ray has a reproduc- chordates, are spatially deployed tally, pollen and isolated plant cells)
tive function similar to ovuliferous structuresof the body that were origi- disengage from the organismal gra-
scales of cones and stamens in flow- nally constrained during their for- dient in the form of cells that are free
ers. Because tissues are well-packed mation by regions of the organismal to initiatetheir own gradient.For ex-
cell multiples,Acetabulariahas some gradient.Tissues are similartypes of ample, in Pelvetia, a close relativeof
tissue-likestructures,such as the sys- cells packedtogetherandconstrained Fucus,Kropfet al. (1986) found that
tem of numerouschloroplasts in the by smallergradientregions or fields, the developmentalaxis forms during
stalk and the whorl of hairs around such as that which specifies the epi- the first cell cycle following fertiliza-
the stalk. At a higher level, organ dermis in both plants and animals. tion. Traditionally, an individual is
systems are sets of integratedorgans Other structural levels are often viewedas the resultof a genotype,and
for spatial deployment, such as the found in multicellular organisms, a new individual is thought to arise
stem and root systems in plants. such as the cell complex that is in- when a new genotypeforms (Ehrlich
Analogously, Acetabularia has or- serted between cell and tissue in the and Holm 1963). Asexual organisms
gan system-like groups found as the cases of the sieve tube-companion are problematic by this line of rea-
ensemble of rays arranged as a cap cell association and the mammalian soning becausethey reproducewith-
and the set of rhizoids spread out as hair follicle. Comparableintermedi- out genetic recombination. But by
a holdfast. ate levels in unicellular organisms hierarchicalreasoning,asexualforms
Amongthe ciliateprotozoans,Vor- also seem to be present in the transi- are considered individuals because
ticella has cell, tissuelike, organlike, tory whorl of hairs in Acetabularia each has its own organismal gradi-
and organismal features (Figure 4). and the pellicle striations in Vorti- ent. A connected clone of cattails
This unicellular organism has the cella, which are equivalent to tissues and a Portuguese Man-of-War are
cell feature of a nucleus, some cyto- of multicellular organisms. examples of connected individuals.

January1999 55
 algae, bacteria, and other microor-
ganisms have both cell and organis-
(E)4 ?
+ () unicellularorganisms
?(D) mal levels.
* An individual is identified as a
living system with its own, separate
organismal gradient.
* Development involves ascending
synthesis of cell features and descend-
ing appearance of higher-level fea-
e-- tures that modify the ascending cell
' C ,, multicellular organisms features.

It is of more than passing interest

Figure 5. Evolution of hierarchicalsystems. c, cell; 0, organism.
that this last feature-ascending syn-

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theses and descending specializa-
A second developmental question tom up" in that amino acids form tion-parallels what is presented in
raised by hierarchical theory con- tubulin monomers that form macro- most introductory biology courses.
cerns the emergence of tissues and molecular assemblies of microtu- Generally, the fate of atoms is to first
organs during development. The cell bules. Microtubules are also con- be part of new molecules that, in
is built of parts put together by en- strained from the "top down" in that turn, become part of new macromol-
zymes, along with self-assembly, and organ growth takes place at the ecules and eventually belong to new
is capable either of cell division (if domed tip of the cell, which pro- organelles and cells. At the same
embryonic) or assisting other cells in duces directional and localized stress time, an organism begins develop-
dividing (if specialized). By contrast, that, in turn, specifically orient mi- mentally at fertilization, first delim-
the organism dictates, through hier- crotubules at the cell level. iting organs, then tissues, and finally
archical constraints, where and when Hierarchical theory also opens the specialty molecules (e.g., collagen,
cell division occurs and what materi- door to a better understanding of hemoglobin, and chlorophyll). This
als form in support cells to help other evolution. A simple primitive cell developmental portrait is essentially
cells divide. Such specialized mol- has a cell level (c) for multiplication, the ontogenetic rule of von Baer
ecules as hemoglobin and chloro- and an organismal level (0) for de- (1828), who asserted that develop-
phyll, although synthesized by cells, termining how and when a cell moves ment passes from general to specific
are higher-level features because they and divides. Together, these levels traits-essentially a downward hier-
influence cell division only indirectly. create a unicellular organism (0). archical description. Both instruc-
In plants, an organismal gradient Evolution can then progress in two tors and researchers accept this hier-
forms several smaller regions for major directions. In one, cell divi- archical paradigm based on their own
determining organs. For example, sion and the organismal gradient re- experiences but often lack the con-
the region of leaf inception fraction- main tightly coupled. This coupling ceptual knowledge for integrating
ates to determine various tissues ex- is elaborated in unicellular organ- the levels.
pressed in a leaf, including epider- isms, such as protozoans, siphona- The two problems stated at the
mis, procambium, and other kinds ceous algae, and bacteria. In the sec- beginning of this paper can now be
of specialized cells. Animals seem to ond direction, cell division and the solved directly. The first problem
have an organismal gradient, por- organismal gradient become un- concerns how the antiquated struc-
tions of which are active in establish- coupled, and groups of cells are con- tural description of a cell might be
ing body regions and body segments; strained by a common gradient (Fig- updated to that of an information
induction mechanisms then take over ure 5). system. von Neumann's approach is
for determining organs and tissues. to view a cell as self-regulated by
Generally, cellular features emerge A new perspective virtue of its ability to process infor-
through ascending syntheses, with mation and as capable of self-divi-
monomers enzymatically combined Hierarchical theory attempts to iden- sion. This description goes well be-
into polymers that are self-assembled tify various levels of an organization yond early ideas of cells as machines,
into macromolecular complexes. By as well as the types of descendent in which internal control is not ex-
contrast, higher levels come from a constraints. In biology, these con- plicitly specified. The feature of self-
descending cascade of determination, straints occur in the form of bond- description is therefore the opening
from the organismal level, through ing, support, enclosure, and control for introducing the broader idea of
the organismal gradient, which is (Figure 2; Korn 1994). information processing and the mod-
subdivided into organ-determining Applying these hierarchical ideas ern view of a cell. The second prob-
subregions, which generate tissue- to organisms, the following points lem-how to distinguish between a
forming interactions. Green's (1987) can be made: cell and an organism, especially
description of plant cell elongation among unicellular forms-is resolved
easily fits into this ascending-de- * The fertilized egg has both cell and by a hierarchical explanation. Uni-
scending scheme. That is, microtu- organismal levels. cellular organisms are both cells, in
bules are synthesized from the "bot- * Ciliated protozoa, siphonaceous having nucleo-cytoplasmic activities,

56 BioScience Vol. 49 No. 1

and organisms, by virtue of the pres- Modelling of Patterns in Space and Time. suchungen iiber die Ubereinstimmung in
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