Thinking of Biology
Biological organization
T he cell theory, or cell doc- ria of self-control and then viewing
trine, which states that all or-
ganisms are composed of simi-
lar units of organization, called cells,
was first enunciated in 1839 and has
remained one of the foundations of
modern biology. This idea predates
other great paradigms in biology,
such as Darwin's theory of evolution
(1859), the rediscovery of Mendel's
laws of inheritance (1900), and the
establishment of comparative bio-
chemistry (1940). Although ultra-
structure research and molecular bi-
ology have added much to the cell
theory, and it has retained its emi-
nent status in biology, the cell theory
faces two ongoing problems.
First, the cell theory began as, and
to a great extent remains, a struc-
tural idea. This structural view,
which is found in most textbooks,
describes the components of a cell
and their fate in cell reproduction.
Today, however, biology focuses on
DNA and its informational features,
and the description of a cell needs to
catch up with this contemporary view
of life. The cell is, and needs to be
described as, a unit of self-control.
That is, the description of a cell needs
to incorporate ideas about how in-
formation is converted to structure.
The second problem with the cell
theory is historical: It first gained
prominence along with the organismal
theory, which is the idea that the
organism has its own structural fea-
tures apart from those of cells. The
ensuing debate over which of the
two structural theories better ex-
plains biological organization, espe-
cially in protists, has never been re-
solved sufficiently by structural
criteria.
A resolution to both problems can
be found by restating the cell and
organismal theories in terms of crite-
by Robert W. Korn
January 1999
A new
an old problem
them as existing at different levels of
biological organization. Levels of
control is the subject of hierarchical
theory; hence, organisms can be bet-
ter described by a hierarchical than a
mainly structural perspective. In this
article, I explain the relationship
between the cell theory and hierar-
chical theory. I begin by developing
both theories and then try to recon-
cile them.
The cell theory-organismal
theory controversy
In 1838, Theodor Schwann and Mat-
thias Schleiden were enjoying after-
dinner coffee and talking about their
studies on cells. According to his
biographer (Fredericq 1884), when
Schwann heard Schleiden describe
plant cells with nuclei, he was struck
by the similarity of plant cells to cells
he had found in animal tissues. The
two scientists went immediately to
Schwann's lab to look at his slides.
Schwann published his book on ani-
mal and plant cells (Schwann 1839)
the next year, a treatise devoid of
acknowledgments of anyone else's
contribution, including that of
Schleiden (1838). He did, however,
summarize his findings into three
famous conclusions about cells:
* The cell is the unit of structure,
physiology, and organization.
* The cell retains a dual existence as
a distinct entity and a building block
in the construction of organisms.
* Cells form by free-cell formation,
similar to the formation of crystals.
The first two statements are still ac-
ceptable, although the third is clearly
wrong. Before Schwann's publica-
tion, there were two rival theories on
how cells form-one claiming free-
cell formation and the other claim-
ing cell division. Schwann picked the
is collaborating with JSTOR to digitize, preserve, and extend access to
look at
wrong one-free-cell formation-be-
cause he failed to consider all avail-
able theoretical ideas. The correct
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interpretation of cell formation by
division was finally promoted by
others and formally enunciated in
RudolphVirchow'spowerfuldictum
Omnis cellula e cellula ("All cells
only arise from pre-existing cells";
Wilson 1896).
The cell doctrine reached its
present-day eminence in 1896 with
the publication of E. B. Wilson's The
Cell in Development and Heredity,
which was an accumulation of what
was known about the roles of cells in
embryology and chromosomal be-
havior.Nevertheless,some biologists
held stubbornlyto the idea that nerve
organizationremainedan important
exception to the cell theory because
some fibers appeared to come from
nonliving substances. But even this
final resistance to the cell theory
succumbed with Ross Harrison's
convincingdemonstration(Harrison
1907) that neurons in culture arise
only from other nerve cells.
The cell is usually investigated by
three approaches: characterizing
structural components, identifying
the biochemical activity of the com-
ponents, and noting how the compo-
nents multiply and segregate during
cell division. In 1952, a fourth ap-
proach to the study of the cell began
when the mathematician John von
Neumann developed a theory of self-
reproducing automata, in which he
viewed cells as self-regulating ma-
chines. Although others had previ-
ously described cells as machines, it
was von Neumann who emphasized
the importance of internal control
and who provided the formalism
behindthis feature.He proposedthat
such a machine would have three
components: an assembler, to make
another automaton; an information
tape, to direct the activity of the
assembler; and a tape recorder, to
51
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make a new copy of the tape.
Because the cell is the smallest
unit of reproduction, the question is
then: Where are these three compo-
nents in a cell? Certain cell organelles
can be recognized as von Neumann's
components if they are viewed from
a generic perspective. The cytosol,
with its batteries of enzymes for syn-
thesis and breakdown of many
biomolecules, functions as the as-
sembler; the chromosomes are, col-
lectively, the information tape; and
the set of DNA polymerases consti-
tutes the tape recorder. To complete
these correspondences, the scrapyard
in which the automaton lives be-
comes the nutrient-rich external en-
vironment of the cell, which includes
at least inorganic materials. von
Neumann had a problem working
out the logic behind the system, par-
ticularly with the type of material
needed in the scrapyard, and so he
abandoned the problem. Only later
were his writings salvaged and ed-
ited by Arthur Burks (von Neumann
1966).
Clearly, von Neumann was aware
that the biological cell was the only
example of self-reproduction known,
and so it probably served to guide
him in constructing his theory. As a
proliferating system, a cell must in-
clude at least some central DNA (as
in prokaryotes) or a nucleus (as in
eukaryotes), and some cytoplasm
with assorted enzymes and mono-
meric building blocks.
In contrast to the smooth devel-
opment of the cell theory over the
past 150 years, the history of the
organismal theory has been fraught
with inaccuracies and misunder-
standings. The organismal theory
proposed that a living thing, or or-
ganism, has a complex organization
and is often, but not necessarily,
composed of cells. When Carl von
Siebolt found, in 1845, that protozo-
ans were similar to cells of multicel-
lular organisms, they were called
unicellular organisms by some (e.g.,
von Siebolt himself and Max Ver-
worn) and noncellular organisms by
others (e.g., Thomas Henry Huxley,
Charles Otis Whitman, Clifford
Dobell). Even metazoans were, for
several reasons, considered noncel-
lular (Singer 1959). First, some early
chordate embryos appeared to have
a free-nucleate stage-a "feature"
52
based on errors in observations. Sec-
ond, many adult animal tissues ap-
peared to have no cell boundaries, a
conclusion that also reflected poor
observation. Third, plant cells are
connected by plasmodesmata and
animal cells have gap junctions; these
seemed to create a continuum of pro-
toplasm instead of separate cells, an
error in interpretation because cells
can still be individuals even though
they are connected. Fourth, some
animal parts cannot be regenerated,
as Wilhelm Roux found when the
destruction of one cell of a two-
celled frog embryo led to a partial
embryo (Roux 1888). This result
suggested to some embryologists that
parts, rather than cells, constitute an
organism-an erroneous assumption
because regulative and mosaic pat-
terns of development were just be-
ginning to be recognized. Moreover,
in 1866, Ernst Haeckel, a dominant
figure in science, claimed that a plant
begins as a cell that simply enlarges
and forms compartments by plate
formation. Adding to the difficulties
for the organismal theory was its
association with vitalism, the phi-
losophy that immaterial forces gov-
ern biological activities, which was
espoused by such eminent thinkers
as General Jan Smuts and Alfred
North Whitehead, among others
(Beck 1957).
To most biologists of the late nine-
teenth century, the organismal theory
offered little for understanding how
life is organized. There were, how-
ever, some positive contributions
from the organismal theory, such as
the construction of a fate map for the
chicken embryo by Wilhelm His. In
this approach (His 1874), various
cytoplasmic regions of the egg were
viewed as destined to become spe-
cific structures of the adult, indepen-
dent of any cellular organization.
Similarly, Kaplan and Hagemann
(1991) cited a number of cases in
which plant form is independent of
cell size, shape, and orientation; they
concluded that at least part of an
organism's morphology must come
from something other than the build-
ing blocks of cells.
After contributing significantly to
the cell theory, von Neumann pro-
posed a new idea about organisms.
His friend, the mathematician Stanis-
law Ulam, suggested that complex
activity such as self-reproduction
might come from a set of similar
units working as a "team." von
Neumann took up this idea and redi-
rected his energy to what he called
"cellular automata." He defined a
cellular automaton as an entity that
can assume various states as speci-
fied by its own encoded information.
Although the cellular automata of a
group can have copies of the same
program, they assume different
states, depending on their status in
the group. Integrated group behav-
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ior comes from the close interaction
between individual automata, and
because individual actions change,
new and interesting behavior pat-
terns arise, including life cycles.
Around 1970, John Conway's game
Life (Gardner 1970), which dealt
with a special brand of cellular au-
tomata, became the rage at com-
puter centers throughout the world.
This little game, carried out by com-
puter or by hand, provided a visual
representation of how complex group
behavior could emerge from a simple
set of rules and a few individual
automata.
A multicellular organism is simi-
lar to a group of cellular automata in
three ways. First, each automaton
has the same program, just as an
organism is composed of cells with
the same genotype. Second, an au-
tomaton can assume any one of a
number of different states encoded
in the program, just as a cell can
differentiate into any one of many
cell types that are encoded in the
genotype. Third, specific ensembles
of automata can pass through cyclic
patterns of structure and behavior
that are parallel to an organism's life
cycle. Although von Neumann's idea
of self-reproduction supported the
cell theory, his concept of cellular
automata gave new meaning to the
organismal doctrine in that it showed
how a small group of cells with a
simple common program could gen-
erate complex behavior.
Just as a biological cell is but one
example of the self-reproducing phe-
nomenon, so the multicellular or-
ganism is but one type of cellular
automata collective. A cell is charac-
terized by a set of nucleo-cytoplas-
mic relationships (involving many
different components, such as chro-
mosomes and enzymes), whereas a
BioScience Vol. 49 No. 1
multicellular organism is composed
of many similar components-cells.
The distinction between the cell and
organismal theories is that, accord-
ing to the cell theory, cells organize
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
pattern (Figure 1).
Hierarchicaltheory
The term "hierarchy" was coined by
Pseudo-Dionysius in the sixth cen-
tury to formalize the relationships
between levels of Christian orders,
or sacred rulers (Osborn 1967). Al-
though the idea of a hierarchy began
as a description of a sacred chain of
authority, it was eventually extended
to other human organizations, such
as government and armies. It also
came to mean a type of classification
in which different groups are included
under a common heading. This in-
clusive approach has appeared spo-
radically in biology. August Comte
in 1854 saw a "hierarchy of sci-
ences," Virchow in 1858 noted a
"hierarchy of pathological condi-
tions," and Hughling Jackson in 1881
described a "hierarchy of nervous
centres in man" (Whyte 1969).
Joseph Woodger first described
biological organization as "hierar-
chical," using a taxonomic approach
(Woodger 1929). First, he classified
the known biological levels of cells,
tissues, and organs; he then exam-
ined how these levels are structurally
related. As part of an organism, a
cell has many relationships to other
cells. These relationships no longer
exist when a cell is isolated from its
normal neighbors; thus, it will be-
have differently. To understand cells,
Woodger insisted, they must be stud-
ied either in their normal environ-
ment or in artificially reconstructed
conditions. Because Roux's half-
embryo of one cell has a neighbor
missing, it will not make exactly half
an adult. This view of the biological
hierarchy of cells, tissues, organs,
and organisms has been passed on
through biology textbooks, starting
with Simpson and Beck (1967). The
January 1999
fertilized egg
organism organism
h
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Cell doctrine Organismal doctrine
Figure1. The differencebetweenthe cell and organismaldoctrines.The cell doctrine
considersa cell as an identifiablesystemwith processesfor supportingitself as well as
processesthat contributeto an intercellularpattern.The organismaldoctrineconsiders
cells as unitscontributingdifferentfeaturesto an overallorganismalpatternby which
cells are sustained.
scheme of a nested classification of inclusive view of hierarchies is the
cells, tissues, and organs (e.g., Table emphasis on the more passive, mem-
1) is now commonly included not bership aspects of organizations.
only in textbooks (e.g., Raven and I have modified Pattee's idea by
Johnson 1996) but also in most schol- emphasizing that in hierarchies, con-
arly descriptions of biological sys- straints are nonreciprocal (Korn
tems (Weiss 1971, Guttman 1976, 1994). For example, the constraint
MacMahon et al. 1978, Koestler of gravitational pull is shared equally
1979, Lieberman and Vrba 1995). by the sun and the earth, but the pull
It was Howard Pattee, a physicist, influences the small mass of the earth
who pointed out that hierarchies can more than the large mass of the sun.
be either physical (meaning, in fact, Therefore, the earth revolves around
physical, chemical, or biological) or the sun. Such nonreciprocal con-
descriptive (cognitive) and that it is straints can be easily identified in vari-
the physical hierarchies that are ous aspects of plant organization (Korn
formed by members at a higher level 1986, 1994, in press). Table 1 shows
constraining members at a lower level more levels of organization than are
(Pattee 1969). Because energy is re- traditionally cited, and the emergence
quired to forge and, often, to main- of these levels during evolution can
tain a physical constraint, physical be understood by the insertion of
hierarchies are dynamic systems. new features accompanied by new
Even Pseudo-Dionysius's original constraints over these new features.
idea that an organization is based on This dynamic view of a hierarchy is
authority implies constraint, because illustrated in Figure 2.
authority dictates constrained behav- Only on rare occasions does the
ior. The problem with the traditional, idea of nonreciprocal constraint ap-
Table 1. Traditional classificational view of the hierarchyof higher organisms.a
Level Plant Animal
Organism Dandelion Mouse
Organ system Shoot Integumentary
Stem Skin
Organ
Tissue complexb Vein Dermis
Tissue Phloem Epidermis
Cell complexb Loading system Hair follicle
Cell Companion cell Keratinocyte
aLower levels (from atom to organelle) are not included.
bThese levels are not typically included in the traditional classificational view.
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
to shoots. The persistence of this
double gradient can be interpreted
as an organismal control constraint
that is important in maintaining in-
determinategrowth and dictatingre-
generationafterinjury.In insecteggs,
gradientsof RNA and protein estab-
lish the three body regions of head,
thorax, and abdomen and subse-
quently the 14 segments (Nusslein-
Volhard 1991). In higher animals,
an egg gradient is often visible by
yolk concentrationor in the location
of the nucleus that is probably re-
placed by special regions for pri-
mary, secondary,and tertiaryinduc-
tions. The adult nervous system
eventually replaces the inductive
fields of the embryo as the means of
organismal maintenance.
Organismalgradients can also be
found in unicellularorganisms, such
as the siphonaceous algae and ciliate
protozoans. The marine alga Ac-
etabularia, or mermaid's umbrella,
has the cell featureof a single nucleus
and some cytoplasm (Figure 4). It
also has a stalk and rhizoid system
and, later, a cap system, each of
which can be regenerated,indicating
the presenceof a regenerationgradi-
ent as an organismalfeature. Certain
structures in Acetabularia serve to
constrain other structures that are
comparable to organs in higher
plants. As examples, the rhizoid
serves for anchorage, analogous to
the root of higher plants; the stalk
functions in support, as does the
stem;and the cap ray has a reproduc-
tive function similar to ovuliferous
scales of cones and stamens in flow-
ers. Because tissues are well-packed
cell multiples,Acetabulariahas some
tissue-likestructures,such as the sys-
tem of numerouschloroplasts in the
stalk and the whorl of hairs around
the stalk. At a higher level, organ
systems are sets of integratedorgans
for spatial deployment, such as the
stem and root systems in plants.
Analogously, Acetabularia has or-
gan system-like groups found as the
ensemble of rays arranged as a cap
and the set of rhizoids spread out as
a holdfast.
Amongthe ciliateprotozoans,Vor-
ticella has cell, tissuelike, organlike,
and organismal features (Figure 4).
This unicellular organism has the
cell feature of a nucleus, some cyto-
January1999
Figure 4. Levels
of organizationin
unicellularorgan-
isms. org, organ-
ism;os, organsys-
tem; o, organ; t,
tissue;c, cell.
plasm,andmost
likely the orga-
nismal feature
of a head-to- d rhi
foot gradient, rh
as demonstrat- Acetu
ed by regenera-
tion studies in
other ciliates (Stevens 1903, Sonne-
born 1963). Faure-Fremiet (1954)
consideredboth unicellularand mul-
ticellular organisms to have a gradi-
ent in their cortical cytoplasm and
suggested that this gradient deter-
mines cytoplasmic specialization in
unicells and cell types in metazoa.
Other unicellularorganismscan also
be characterized by an organismal
gradient.In bacteria,polarity occurs
in wall synthesis, spore formation,
and, sometimes, motility. Simple al-
gae, fungi, and protozoa also ex-
pressasymmetricorganizationalfea-
tures similar to those in bacteria.
Multicellular organisms can also
be given a new hierarchicaldescrip-
tion, not by the traditional approach
of levels but by the identification of
descendent constraints. The organ-
ism is the body and is representedat
some times by a gradient. Organs,
such as leaves in plants and bones in
chordates, are spatially deployed
structuresof the body that were origi-
nally constrained during their for-
mation by regions of the organismal
gradient.Tissues are similartypes of
cells packedtogetherandconstrained
by smallergradientregions or fields,
such as that which specifies the epi-
dermis in both plants and animals.
Other structural levels are often
found in multicellular organisms,
such as the cell complex that is in-
serted between cell and tissue in the
cases of the sieve tube-companion
cell association and the mammalian
hair follicle. Comparableintermedi-
ate levels in unicellular organisms
also seem to be present in the transi-
tory whorl of hairs in Acetabularia
and the pellicle striations in Vorti-
cella, which are equivalent to tissues
of multicellular organisms.
Vorticella
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Genes are responsible for control
of processesat the cell andorganismal
levels. Somegenes specify regulatory
proteins for metabolism and cell di-
vision, whereas others specify the
production of an intercellulargradi-
ent. All genes are nucleotide se-
quences, regardlessof the hierarchi-
cal level at which their products
function. However, genes for cell
function interact with one another
indirectlythrough protein products,
whereas genes for organismal activ-
ity interactwith regions of the gradi-
ent, also indirectly, through their
protein products.
The presence of an intercellular
organismalgradientraises two inter-
estingdevelopmentalquestions.First,
how does the gradientbecome estab-
lished? This question is another way
of asking when an individual organ-
ism begins. Individualsemergewhen
eggs and spores (and, experimen-
tally, pollen and isolated plant cells)
disengage from the organismal gra-
dient in the form of cells that are free
to initiatetheir own gradient.For ex-
ample, in Pelvetia, a close relativeof
Fucus,Kropfet al. (1986) found that
the developmentalaxis forms during
the first cell cycle following fertiliza-
tion. Traditionally, an individual is
viewedas the resultof a genotype,and
a new individual is thought to arise
when a new genotypeforms (Ehrlich
and Holm 1963). Asexual organisms
are problematic by this line of rea-
soning becausethey reproducewith-
out genetic recombination. But by
hierarchicalreasoning,asexualforms
are considered individuals because
each has its own organismal gradi-
ent. A connected clone of cattails
and a Portuguese Man-of-War are
examples of connected individuals.
55

(E)4 ?
+ ()
?(D)
e--
' C ,,
Figure 5. Evolution of hierarchicalsystems. c, cell; 0, organism.
A second developmental question
raised by hierarchical theory con-
cerns the emergence of tissues and
organs during development. The cell
is built of parts put together by en-
zymes, along with self-assembly, and
is capable either of cell division (if
embryonic) or assisting other cells in
dividing (if specialized). By contrast,
the organism dictates, through hier-
archical constraints, where and when
cell division occurs and what materi-
als form in support cells to help other
cells divide. Such specialized mol-
ecules as hemoglobin and chloro-
phyll, although synthesized by cells,
are higher-level features because they
influence cell division only indirectly.
In plants, an organismal gradient
forms several smaller regions for
determining organs. For example,
the region of leaf inception fraction-
ates to determine various tissues ex-
pressed in a leaf, including epider-
mis, procambium, and other kinds
of specialized cells. Animals seem to
have an organismal gradient, por-
tions of which are active in establish-
ing body regions and body segments;
induction mechanisms then take over
for determining organs and tissues.
Generally, cellular features emerge
through ascending syntheses, with
monomers enzymatically combined
into polymers that are self-assembled
into macromolecular complexes. By
contrast, higher levels come from a
descending cascade of determination,
from the organismal level, through
the organismal gradient, which is
subdivided into organ-determining
subregions, which generate tissue-
forming interactions. Green's (1987)
description of plant cell elongation
easily fits into this ascending-de-
scending scheme. That is, microtu-
bules are synthesized from the "bot-
56
unicellularorganisms
multicellular organisms
tom up" in that amino acids form
tubulin monomers that form macro-
molecular assemblies of microtu-
bules. Microtubules are also con-
strained from the "top down" in that
organ growth takes place at the
domed tip of the cell, which pro-
duces directional and localized stress
that, in turn, specifically orient mi-
crotubules at the cell level.
Hierarchical theory also opens the
door to a better understanding of
evolution. A simple primitive cell
has a cell level (c) for multiplication,
and an organismal level (0) for de-
termining how and when a cell moves
and divides. Together, these levels
create a unicellular organism (0).
Evolution can then progress in two
major directions. In one, cell divi-
sion and the organismal gradient re-
main tightly coupled. This coupling
is elaborated in unicellular organ-
isms, such as protozoans, siphona-
ceous algae, and bacteria. In the sec-
ond direction, cell division and the
organismal gradient become un-
coupled, and groups of cells are con-
strained by a common gradient (Fig-
ure 5).
A new perspective
Hierarchical theory attempts to iden-
tify various levels of an organization
as well as the types of descendent
constraints. In biology, these con-
straints occur in the form of bond-
ing, support, enclosure, and control
(Figure 2; Korn 1994).
Applying these hierarchical ideas
to organisms, the following points
can be made:
* The fertilized egg has both cell and
organismal levels.
* Ciliated protozoa, siphonaceous
algae, bacteria, and other microor-
ganisms have both cell and organis-
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-
tures that modify the ascending cell
features.
It is of more than passing interest
that this last feature-ascending syn-
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theses and descending specializa-
tion-parallels what is presented in
most introductory biology courses.
Generally, the fate of atoms is to first
be part of new molecules that, in
turn, become part of new macromol-
ecules and eventually belong to new
organelles and cells. At the same
time, an organism begins develop-
mentally at fertilization, first delim-
iting organs, then tissues, and finally
specialty molecules (e.g., collagen,
hemoglobin, and chlorophyll). This
developmental portrait is essentially
the ontogenetic rule of von Baer
(1828), who asserted that develop-
ment passes from general to specific
traits-essentially a downward hier-
archical description. Both instruc-
tors and researchers accept this hier-
archical paradigm based on their own
experiences but often lack the con-
ceptual knowledge for integrating
the levels.
The two problems stated at the
beginning of this paper can now be
solved directly. The first problem
concerns how the antiquated struc-
tural description of a cell might be
updated to that of an information
system. von Neumann's approach is
to view a cell as self-regulated by
virtue of its ability to process infor-
mation and as capable of self-divi-
sion. This description goes well be-
yond early ideas of cells as machines,
in which internal control is not ex-
plicitly specified. The feature of self-
description is therefore the opening
for introducing the broader idea of
information processing and the mod-
ern view of a cell. The second prob-
lem-how to distinguish between a
cell and an organism, especially
among unicellular forms-is resolved
by a hierarchical explanation. Uni-
cellular organisms are both cells, in
having nucleo-cytoplasmic activities,
BioScience Vol. 49 No. 1
and organisms, by virtue of the pres-
ence of molecular gradients and struc-
tural polarity. The cell and organism
exist at two different levels of the
living hierarchy, vertically interact-
ing but distinctly separate.
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Robert W. Korn (RKorn@Bellarmine.edu)
is a professor of biology at Bellarmine
College, Louisville, KY40205. A develop-
mental botanist, he is interested in model-
ing the geometries of plant cells and tissues
and the hierarchical implications of these
levels of organization. C 1999 American
Institute of Biological Sciences.
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