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Levels of Biological Organization: An Organism-Centered Approach

Author(s): James A. MacMahon, Donald L. Phillips, James V. Robinson, David J. Schimpf

Source: BioScience, Vol. 28, No. 11 (Nov., 1978), pp. 700-704
Published by: University of California Press on behalf of the American Institute of Biological Sciences
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Levels of Biological Organization:
An Organism-Centered

James A. MacMahon, Donald L. Phillips,

James V. Robinson, and David J. Schimpf

The concept of "Levels of Organiza- Odum 1959, Weisz 1959). Since then, ward along each of the four lines of rela-
tion" in the biological sciences has had a schemata of various forms have ap- tionship types. Some categories might be
checkered history. The general nature of peared in many texts, sometimes depict- skipped; e.g., for unicellular organisms,
this concept is one of increasing com- ed in more complexity (e.g., Weisz tissue-level relationships are not mean-
plexity from molecule through cell, orga- 1966). ingful nor is the deme concept appropri-
nism, and ecosystem, though details Guttman (1976) doubted the veracity ate for an asexual form. Our categories
vary among authors. Most contemporary and usefulness of the levels-of-organiza- are not exhaustive, but merely the most
biologists accept the concept, the useful- tion concept in biology, pointing out that commonly used; others, such as sub-
ness of which has been stated by Orians the composition and interactions of each family or organelle, can be used.
(1973, p. 3): level in the hierarchy are not determined
Since biological investigations range merely by systems of the level immedi-
from determiningthe structureof some ately below; systems of all levels inter- Physiological-Anatomical
act, and some levels may be absent. Al- Relationships
importantmolecule to measuring the
transfer of solar energy through com- though Guttman's objections may make
plex systems of interactingorganisms, the concept less simplistic, they do not
some conception of the level of organi- violate the principles of hierarchy theory The various relationships from the or-
zation at which a problemis relevantis (Bossort et al. 1977). Hence, they do not ganism through the subatomic particle
essential. At any given level the kinds are by far the most often discussed in the
preclude the existence of a hierarchical
of questions that can be asked about structure in the biological world worthy context of "levels of biological organiza-
every problem must also be under- of study. tion." One reason this line has had scien-
stood. tific appeal is that all levels are com-
We believe that an organizational con-
Early treatises concerned systems at cept of this type has heuristic value, pro- posed of aggregations of units of lower
levels in the hierarchy. For example,
the level of the organism or below. At viding insights to both students and re-
the same time, basic concepts of hier- searchers. we here molecules are composed of atoms and in
turn subatomic particles; cells are com-
archies above the organism level were propose an alternative schema which,
being formulated by ecologists, though because of our biases, elaborates on the posed of subcellular structures, mole-
in a comparatively diffuse manner. Rowe supraorganismal levels. Our concept em- cules, atoms, and subatomic particles.
The physiological-anatomical line of
(1961) provides an extensive review of phasizes the organism as the pivotal unit,
the history of "levels-of-organization" not just another rung on the ladder, and relationships is particularly reasonable
because most levels are clearly bounded
concept in the ecological context. In ad- changes the emphasis from "levels of or-
in space and time, i.e., contained within
dition, he alludes to the importance of ganization" to "biological relationships
of the organism." Our nontraditional use a living organism. Important exceptions
the individualistic approach (a la Glea-
son 1927). Increased interest in levels of of some biological terms might initially occur when some of an organism's mole-
cause conceptual dyspepsia, but we be- cules or other constituents are used out-
organization may have been due to sche-
matic presentations in textbooks (e.g., lieve consistancy in our definitions will side of the individual, as is the case with,
offset any novelty of the usages we pro- e.g., plant allelochemics, bacterial exo-
pose. We will discuss briefly each branch enzymes, insect pheromones, or fish
of our schema (Fig. 1) and allude to some electric fields.
The orderof authorshipis strictlyalphabetical.This of its implications, especially those in This line represents a hierarchy of gen-
paperis a contributionfromthe Departmentof Biol- erally increasing structural complexity
ogy and the Ecology Center, UMC 53, Utah State ecology.
University, Logan, UT 84322. Currentaddress for culminating in the organism. Each level
Phillipsis Departmentof Biology, EmoryUniversi- of complexity appears to have some
ty, Atlanta,GA 30322,and for Robinsonis Depart- emergent properties, but structure and
mentof Biology, Universityof Texas, Arlington,TX
76010. ? 1978AmericanInstituteof BiologicalSci- Our organism-centered diagram (Fig. function are not determined merely by
ences. All rightsreserved. 1) should be read from the organism out- these emergent properties.

700 BioScience Vol. 28 No. 11

Although structure and function are
usually thought of as intimately associat-
ed, there is no clear functional hierarchy
represented in the physiological-anatom- DIVISION
ical line. In fact, this line, as much as any
of our relationship lines, shows that a CLASS
given level may coact with other levels
above or below it in the series. All such ORDER POPULATION
coactions determine function. For ex-
ample, a tissue's functions are based on 2 FAMILY 3 4
molecules affecting it, the state of its
constituent cells and extracellular ma-
trix, and the milieu of the organ and or-
gan system of which it is a part. I
We could subdivide the connections SPECIES DEME ECOSYSTEM
between any two of our levels if a specif-
ic area of interest required this closer
scrutiny; e.g., there are important in-
creasing structural complexities going
from a nucleotide to a nucleotide pair to
a triplet, etc., in a DNA molecule. The
possibility of such refinements does not ORGANSYSTEM BI"
of the
negate the existence of the more general I ORGANISM
pattern, which we choose to emphasize ORGAN Physiological-Anatomical
for simplicity. I 2 Phylogenetic
Phylogenetic Relationships 1 I Coevolutionary
Historically, this is probably the oldest CELL 4. Matter-Energy Exchange
and least modified series of relationships
of the organism, at least in its structure. SUBCELLULAR
Conceptually, however, the nature of I
each of these relationships is highly con- MOLECULE
troversial. Recently, new kingdom ar-
rays have been proposed (Whittaker
1969, Woese and Fox 1977), while the
species concept is periodically reas-
sessed, even to the point of calling the SUBATOMIC
organism the "species" (the hyper- Fig. 1. Biological relationships of the organism.
modern species concept of Ghiselin
1974, Platnick 1977). Moreover, the very
active controversies concerning cladis- phylogenetic relationships will alter the tionary relationships. Coevolution is the
tics, phenetics, and evolutionary system- fact that the individual organism has a subset of evolution resulting from selec-
atics continue (Mayr 1974). Despite phylogeny and is variously related to tive forces produced by interactions with
these philosophical paroxysms, the orga- other organisms with some level of ge- other organisms. Traditionally, the term
nism through genetic descent from com- netic difference involved. Regardless of coevolution has referred to the result of
mon ancestors remains variously related what one calls the levels (ours are tradi- interspecific interactions, and most com-
to other organisms. tional) or how one determines member- monly reciprocal effects have been im-
This continuum of relatedness is arbi- ship in the level, the organism is related plied. Since we see no qualitative dif-
trarily partitioned by a hierarchy of to others phylogenetically. ference in the nature of the interactions
terms (Fig. 1). In the ideal case, the ter- Parenthetically, we note the lack of a between different species or within the
minology will reflect levels of phyloge- universally accepted formal term for the same species, we suggest that coevolu-
netic relationship. The problems in the ultimate taxon, i.e., one containing sev- tionary interactions can occur between
phylogenetic line are generally those of eral kingdoms. Since the erection of such organisms, demes, or populations of the
choosing appropriate characters to es- a level implies a decision as to a common same species and may affect only one of
tablish levels. The characters should re- origin for two or more kingdoms, we the interactants.
flect degree of difference. Certainly there gladly leave this problem for others. The combinations of positive, nega-
is no current consistency in application tive, and neutral effects on expected re-
of terms. (Are two passeriform bird fami- productive success are embodied in the
lies as distinct from each other as two Coevolutionary Relationships
familiar concepts of competition, mu-
araneomorph spider families?) Organisms are associated with other tualism, commensalism, etc. (Burkhold-
We do not enter any of these con- organisms in ways that affect each oth- er 1952). These are specific kinds of coe-
troversies. We merely suggest that none er's expected reproductive success; we volutionary interactions. Though usually
of the fluidity of conceptualization in the will term all such interactions coevolu- depicted for pairs of organisms or popu-

November 1978 701

lations, these interactions may be in- (1971) stated: "... community in the Matter-EnergyExchange
direct, i.e., mediated through other orga- ecological sense (sometimes designated Relationships
nisms, demes, or populations. Although 'biotic community') includes all of the
the interactions occur between orga- populations occupying a given area." Our greatest departure from tradition-
nisms directly, it is often useful to treat Conceptually, we do not use a spatial al thinking comes in the removal of com-
statistically aggregated groups of orga- boundary, but rather a coevolutionary- munity and population from in-line, sub-
nisms as though the groups interact effect boundary. Although it is a truism ecosystem levels. The reasons for this
directly. Organisms and multiorganism that a community occupies a finite space, include the fact that the term interaction
aggregations known as demes, popu- the only a priori spatial boundaries for has had at least two different meanings
lations, and communities constitute the communities we study are those of for ecologists. One meaning is in the
coevolutionary units. the biosphere itself. A recent definition sense of effects of one organism or popu-
When organisms interbreed with a of community (Connell and Slatyer 1977) lation on the expected reproductive suc-
high frequency, they may essentially implies some of what we propose here: cess of another. In the preceding section,
meet the criteria for panmixis and thus "We define a community as the set of or- we suggested that such interactions char-
constitute a deme. Demes are not neces- ganisms that occur together and that sig- acterize coevolutionary systems. The
sarily genetically isolated from each oth- nificantly affect each other's distribution second meaning suggests that environ-
er. At a lower frequency of genetic ex- and abundance. It is the interactions that mental interactions consist of matter or
change (interdemic), we term the make a community a worthy unit of energy flow across the surface of an or-
aggregate of interacting conspecific study" (see also Curtis 1959, Menge and ganism, e.g., ". . . in the functional
demes apopulation. Populations can, of Sutherland 1976). sense, an interaction represents the
course, be comprised of coevolutionarily This emphasis on interaction repre- transfer of energy or mass between an
interacting conspecific hermaphrodites sents a common, though often unspoken, organism and its environment" (Spomer
or nonsexual organisms. Groups of inter- assumption in definitions of community 1973).
acting populations, among which no and population. In our definition, inter- Simply stated, an ecosystem is a set of
gene exchange is taking place but whose specific coevolutionary interaction is a organisms and inanimate entities con-
demography or gene pools are affected requirement for membership in a com- nected by exchanges of matter or ener-
by the interaction, are termed commu- munity (intraspecific interaction for pop- gy. The entities exchanging and the
nities. In the usual case, these are gener- ulation). Any a priori spatial bounda- items being exchanged together consti-
ally not conspecific populations. By this ries, other than those of the biosphere, tute the ecosystem components. In gen-
definition, the simplest community are strictly arbitrary. Organisms might eral, the strongest interactions occur
would be comprised of two genetically co-occur in space and time yet not be among spatially proximate ecosystem
isolated, but coevolutionarily inter- connected by coevolutionary inter- components.
acting, non-conspecific populations. De- actions; they, then, do not belong to the Atmospheric and oceanic circulation
pending on the exact criteria one uses to same community (or population) as we tie all the earth's organisms into a com-
define species, one can conceive of the define it. mon ecosystem known as the biosphere.
situation where two populations of the The choice of coevolutionary-effect The biosphere is an open system, e.g.,
same species are essentially reproduc- boundaries depends on the scientific radiation impinges on the earth from out-
tively isolated, yet co-occur, e.g., in- questions addressed by the investigator. er space and in turn reradiates away
cipient sympatric speciation (White The vagility, longevity, and other life from the earth. All lesser ecosystems
1978). These populations might affect history characteristics of the organisms are, therefore, also open systems; spatial
each other's reproductive success and of interest to the researcher will combine boundaries for them are strictly arbi-
are, thus, acting qualitatively no dif- with the strength of interaction he or she trary. By our definition, ecosystems con-
ferently from the interspecific inter- deems significant to determine the space- taining a single organism are conceiv-
actions we use to define community. De- time dimensions studied. Conspicuous able. This is contrary to the traditional
spite this, we opt to designate the organisms of low vagility that have short view (Odum 1971, Whittaker 1975) that
community as a multispecies entity. generation times (relative to the period an ecosystem consists of a community
The expanded sense of our definition of observation) and are involved in inter- plus its environment (which is true in
of community implies that if every orga- actions easily observed are those most some cases).
nism in the world is linked with every favorable for studies of coevolutionary The biosphere is not merely the largest
other organism through coevolutionary relationships. terrestrial ecosystem. It differs qualita-
interactions, no matter how remote, they Arbitrary truncation of a coevolution- tively from lesser ecosystems in that
all belong to a large, but scientifically un- ary unit in space may be acceptable if the biotic flux across its boundaries is negli-
interesting, community. As we detail be- investigator adequately accounts for the gible, though abiotic flux is not. The bio-
low, it is coevolutionary interactions interactions crossing the boundaries. In sphere also differs from any "world com-
more proximate than those implied many studies, the community is parti- munity" in that the latter, by definition,
above that are of interest and measurable tioned taxonomically, and one phyloge- is a closed system.
by scientists, but the possibility of netic group (taxocene) is studied. Such As it is for the community, the choice
a "world community" cannot be dis- studies are also acceptable, as long as of ecosystem boundaries, in practice, is
missed a priori. The existence of a world the investigator is mindful that the inter- investigator-dependent. Scientific gener-
community is probably untestable and of actions with organisms outside that tax- alization depends on the recurrence of
little meaning in the time frame that we ocene may be coevolutionarily impor- similar units in space or time. Ecosystem
observe organisms. tant. The resultant increased tractability units are frequently chosen by spatial
Our definition of community differs may be worth the reduced coevolution- discontinuities in such features as physi-
from those extant. For example, Odum ary resolution. ognomy, physiography, taxonomic com-

702 BioScience Vol. 28 No. 11

position or any environmental factor. of communities and ecosystems is, in ited space-time units. Such units usually
Temporal discontinuities are manifest in fact, a semantic problem confounded by include truncated demes, populations, or
such regular phenomena as diurnality, arbitrarily delimiting entities in nature communities and not the integral coevo-
lunar cycles, seasonality, etc. Tempo- that are weakly differentiated at best. lutionary units on which the theory is
rally irregular events may be important Thus, if one chooses to study a defined based. These truncated units may not
to some ecosystems, e.g., those follow- plot of ground, say a hectare of desert, to have the same properties as the integral
ing a pulse-reserve paradigm (Noy-Meir elucidate the "structure and function" of units. Furthermore, the space-time unit
1973), but such events are difficult to use a desert ecosystem or community, one may include organisms that are not inter-
for bounding ecosystems. has set an arbitrary boundary for all acting coevolutionarily with the other or-
processes of interest. The organisms af- ganisms; the biotic diversity of such ag-
IMPLICATIONS fecting the matter and energy flow of the gregations may exceed that predicted by
hectare may differ from those organisms theories based on coevolutionary inter-
We suggest that our schema has some that affect the expected reproductive action. This fundamental difference be-
conceptual advantages over those pre- success of the component organisms. tween the nature of effect and dimen-
viously proposed and that our organism- Organisms common to both groups may sional boundaries makes more difficult
centered approach is a step closer to de- still differ in the relative importance of the task of understanding ecological
picting realistically the mode of opera- their ecosystem vs. community effects. patterns.
tion of organisms in nature. For example, pollinators affect the re- Two interpretations of the nature of an
Our selection of the term coevolution- productive success of plants out of pro- organism's community are possible. One
ary relationships (rather than evolution- portion to their impact on the plants' alternative defines a community as an ag-
ary relationships) should not be read as a matter or energy exchange. Hence, one gregation of populations connected by
denial of the importance of abiotic forces should be cognizant that an understand- statistically detectable effects on their
in the evolutionary process. Rather, we ing of ecosystem structure and function size or genetic composition. An orga-
chose our terminology to underscore the need not always provide insight into nism belongs to such a community by
fact that evolutionary change in popu- community properties; the inverse is al- virtue of membership in one of those
lations occurs in a community context, so true. When organisms occupy proxi- connected populations.
not in a biotic vacuum. The term coevo- mate positions in the space-time contin- The second interpretation is more
lution has traditionally been applied to uum, the probability of their being strictly organism-centered. Consider the
adaptations, often complex, resulting coevolutionarily related or connected by case of a bird. In this view, the bird's
from reciprocal biotic interactions. We the flow of matter and energy increases. community is composed of those other
see these as merely the extreme and ob- This is especially true in the latter case organisms that affect its reproductive
vious outcomes of the processes we label because at the very least the decompos- success. In this organism-centered ex-
as coevolutionary. ing remains of dead organisms become a treme, other coevolutionary units af-
Knowledge of the coevolutionary lega- diffuse ecosystem resource. fected by the bird are not members of its
cy from previous generations is impor- In a practical vein, contemporary use community (unless they also happen to
tant to understanding the organization of of phrases like "bird community of affect the bird); however, the bird is a
extant demes, populations, and commu- Green Canyon" are acceptable as long member of their communities.
nities. The Founder Principle (Mayr as one realizes that community in that Both views of the community are use-
1942) in population biology and the role sense is really a partial and truncated ful. The organism-centered extreme is
of historical factors in ecology are ex- community; i.e., the coevolutionary in- the approach preferred in attempts to un-
amples of such legacies. Our concept of teractions probably involve more than derstand coevolution of phenotypic
coevolutionary relationships does not re- just birds and more than just the orga- characteristics, e.g.. life-history traits
quire that the constituent units affect nisms of Green Canyon itself. For con- (Steams 1977). Coevolutionary units not
each other reciprocally, only that each venience, one can truncate the commu- affecting a bird's population are not im-
unit affects or is affected by another. An nity in space or time or partition it portant for its evolution. Excluding them
extreme interpretation might suggest taxonomically with the proper series of from the coevolutionarily defined com-
that, since previous generations are co- modifying words. munity simplifies the task of understand-
evolutionarily related to current ones via The term for all of the organisms of ing population evolution in a community
one-way interactions, then dead orga- an ecosystem by previous definitions has context. The connected populations or
nisms belong with living ones in coevolu- usually been community. Since we do statistical view is the paradigm of choice
tionary units. Although extant coevolu- not view a community strictly as a subset in attempts to understand community
tionary units cannot be perfectly of an ecosystem, another term must be evolution (Cody and Diamond 1975). In
understood without considering their an- used. The traditional terms biota and as- this case, the focus is on the presence or
cestry, most studies emphasize inter- sociation might seem to obtain but these absence of species in the community and
actions among living organisms. have slightly different connotations. To not on the effects of the community on
Potentially the most controversial por- avoid coining a new word, we will simply the individual, with consequent popu-
tion of our schema, in view of contempo- refer to the organisms as the biotic com- lation evolution.
rary definitions, comes with our use of ponents of the ecosystem. The emphases herein are that the orga-
the terms community and ecosystem.' Deductive theory of the organization nism is the central unit of biological rela-
The reason we feel compelled to break of communities, populations, or demes tionships and participates in all four rela-
with tradition (though one based on im- (e.g., Cody and Diamond 1975, May tionship lines simultaneously. Any line
precise definition and usage) is that we 1976) should apply to the coevolutionary of relationships can influence any other
believe that much disagreement by ecol- relationships line. Testing of such theory through their common component level,
ogists regarding the nature and definition is usually attempted in arbitrarily delim- the organism. The same organisms that

November 1978 703

basis (coevolutionary); or over many munitiesand theirrole in communitystabil-
INDUSTRY, generations (phylogenetic). In fact, these ity and organization. Am. Nat. 111:1119-
GOVERNMENT, are relative time intervals for any given 1144.
ACADEME... organism; the absolute time intervals of Curtis, J. T. 1959. The Vegetation of Wiscon-
sin. University WisconsinPress, Madison.
.. ARE YOU SEEKING importance are specific to the organism
Dawkins, R. 1976. The Selfish Gene. Oxford
BIOLOGISTS? in question. Thus, an event recurring at a
University Press, New York.
given temporal frequency may be dealt Ghiselin, M. T. 1974. A radical solution to the
Your job description can be with physiologically by longer-lived or- species problem.Syst. Zool. 23: 536-544.
listed without charge in the ganisms but may produce marked popu- Gleason, H. A. 1927. Furtherviews on the
AIBS Employment Newslet- lation fluctuations in shorter-lived orga- succession-concept.Ecology 8: 299-326.
nisms (Slobodkin and Rapoport 1974). Guttman,B. S. 1976. Is "levels of organiza-
ter, which is distributed to
In summary, we propose the following tion" a useful biological concept? BioSci-
approximately 300 job- ence 26: 112-113.
seekers. system of definitions for some ecological
aggregations of organisms: Lewontin, R. C. 1970.The units of selection.
The Newsletter is issued six Annu. Rev. Ecol. Syst. 1: 1-18.
* Deme-a group of coevolutionarily in- May, R. M., ed. 1976. Theoretical Ecology.
times annually (February, W. B. Saunders, Philadlephia.
teracting organisms that interbreed in a
April, June, August, October, manner approximating panmixis. Maynard-Smith,J. 1964.Groupselection and
and December) to qualified * Population-a kin selection. Nature 201: 1145-1147.
group of coevolutiona-
persons trained in all disci- Mayr, E. 1942. Systematics and the Origin of
rily interacting demes or non-outcrossing
Species. Columbia University Press, New
plines of the life sciences. organisms, which due to a common de- York.
To take advantage of this scent are genetically similar enough to be 1974. Cladistic analysis or cladistic
considered conspecific, yet do not inter- classification? Zool. Syst. Evol.-forsch. 12:
opportunity to recruit your breed sufficiently to be a panmictic unit. 94-128.
personnel from the widest * Community-a group of coevolutiona- Menge, B. A., and J. P. Sutherland. 1976.
possible selection, send your rily interacting populations connected by Species diversitygradients:synthesisof the
job descriptions to the AIBS effects of one population on the demog- roles of predation, competition, and tem-
Employment Newsletter, raphy or genetic constitution of the poral heterogeneity. Am. Nat. 110: 351-
1401 Wilson Blvd., Arling- other. 369.
ton, VA 22209. Telephone: * Ecosystem-the plexus
Noy-Meir, I. 1973. Desert ecosystems: envi-
ronment and producers. Annu. Rev. Ecol.
(703) 527-6776. both of abiotic entities and at least one
Syst. 4: 25-51.
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exchange matter or energy may affect * Biosphere-the ecosystem which in- 1971. Fundamentals of Ecology, 3rd
each other's expected reproductive suc- cludes all of the organisms on Earth dur- ed. Saunders, Philadelphia.
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at the same time. In our organism-cen- troduction to Biology, 2nd ed. Allyn and
tered view, the environment affects all ACKNOWLEDGMENTS Bacon, Boston.
four lines by acting directly on the Platnick, N. I. 1977. [Review of] C. N. Slo-
We thank our colleagues D. Andersen, bodchikoff, ed. 1976. Concepts of Species.
organism. M. Caldwell, K. Dixon, J. Haefner, I. Syst. Zool. 26: 96-98.
Furthermore, the organism is the fun- Rowe, J. S. 1961. The level-of-integration
Palmblad, and J. Warner for reviewing
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a revised manuscript and offered numer-
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ous useful comments. L. Finchum typed action" and "operationalenvironment"in
also Dawkins 1976). This does not mean
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sible by NSF Grant DEB 75-13996 to Steams, S. C. 1977.The evolution of life his-
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cur. For many organisms, effect may be
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704 BioScience Vol. 28 No. 11