Q1.

Distinguish between equilibrium and disequilibrium paradigms of plant succession

According to Powell (2000), landscapes and the ecosystems that comprise them age through
time. The series of changes resulting in forest ageing is called plant succession. Plant succession,
which begins with colonization or reoccupation of a disturbed area by vegetation, refers to
temporal changes in both species abundance and vegetation structure. Once initiated, plant
succession may follow a variety of pathways (directions) and can occur at varying rates of speed
(Drury and Nesbit, 1973). Since the composition, structure, and function of an ecosystem can
change as plant succession progresses, it is important that land managers understand the many
ways in which succession influences forest development to ensure that management activities are
placed on a sound ecological foundation (Powell, 2000). The equilibrium and non-equilibrium
paradigms establish the ecological basis for the plant succession debate. The equilibrium
paradigm has been in existence since the beginning of the scientific inquiry, while the non-
equilibrium paradigm is of more recent origin (Egerton, 1973). The two paradigms represent
unique interpretations of ecosystem behaviour in response to disturbance.

The classical equilibrium paradigm as articulated by Clements (1916) believed that plant
succession caused ecosystems to develop in a predictable sequence of steps, much the same way
as a human infant matures into an adult. Proponents of this super-organism philosophy
maintained that individual species were linked together in mutually beneficial systems exhibiting
properties greater than the sum of their parts (Clements, 1916, Egerton, 1973). Clements
contended that nature was orderly and that its order was for the most part stable and self-
regulating. He assumed that the normal condition of ecosystems was a state of homeostasis or
equilibrium, a forest grows to a mature climax stage that becomes its naturally permanent
condition (Clements, 1916). Disturbance was a transitory phenomenon, a foreign intrusion into
an otherwise balanced and harmonious

In other words, the equilibrium paradigm and its associated metaphor, ‘the balance of nature’,
are founded on the assumption that ecosystems possess the capacity for internal regulation
through negative feedback mechanisms, including intense intra- and interspecific competition
and plant-animal interactions (Powell, 2000). The capacity for internal regulation is assumed to
contribute to the predictable and directional response of equilibrium vegetation dynamics
because succession must pass through similar stages to a single equilibrium point (Pickett and
Ostfeld, 1995). Therefore, equilibrium systems are assumed to return to their pre-disturbance
state (homeostasis) or to their pre-disturbance trajectory (homeorhesis) when the disturbance has
ceased (Powell, 2000).

Many contemporary ideas about the environment are based on Clements’ (1916) notion that
nature is capable of retaining its inherent balance more or less indefinitely if only humans could
avoid disturbing it (Cronon, 1996). Clementsian ideology about the balance of nature is still
espoused today: “we must not disturb the hierarchical balance of nature and the food chain. The
earth has a natural system of interacting homeostatic mechanisms similar to the human body. If
one system is diseased, then other systems develop abnormalities in function”. Clements and his
followers viewed disturbance as a rare phenomenon and believed that most sites in North
America supported climax vegetation prior to Euro-American settlement. The classical
equilibrium paradigm of plant succession, a dominant ecological philosophy was based on the
following principles (Clements, 1916, Pickett and Ostfeld, 1995):

 Plant succession was orderly, unidirectional, always passed through the same phases, and
was therefore predictable and deterministic.
 Ecosystems were viewed as closed, self-regulating, and subject to a single stable equilibrium.
 Plant succession occurred only when one plant community modified the physical
environment to an extent allowing another community to get established.
 Plant succession always culminated in a relatively stable community (the climax plant
association).
 The stable endpoint of plant succession (climax) was the norm, with disturbances being
nothing more than aberrant disruptions of the normal successional process.
 Humans were not included on the roster of normal ecological factors, either as an ecosystem
component or as a process (disturbance agent, etc.).

On the other hand, ecologists began to criticize the equilibrium paradigm early in the 20th
century for several reasons, including (i) limited evidence to support the occurrence of
equilibrium systems, (ii) inability to account for the dynamic behaviour of various ecological
systems and (iii) the implication that historical events play only a minor role in ecosystem
dynamics (O’Neill et al, 1986; Wu and Loucks, 1995; O’Neill, 2001). However, it was not until
the 1970s that the theoretical basis for plant succession deviated from the equilibrium paradigm,
when several prominent ecologists promoted the existence of multiple steady or equilibrium
states (Briske, 2003). Disturbances were assumed to force one stable community across a
threshold to a subsequent stable community on the same site. However, the validity of the
multiple steady-state concept has been questioned on the basis of various conceptual and
experimental issues (Walker and Wilson, 2002).

By the mid-1950s, ecologists began to realize that natural systems are not nearly so balanced or
stable as Clements would have us believe, and his habit of talking about ecosystems as if they
were organisms, with a life cycle much like that of a living plant or animal, was far more
metaphorical than real (Wu and Loucks, 1995). This change in attitude gradually led to a modern
non-equilibrium or dynamic equilibrium paradigm of plant succession. The non-equilibrium
model considers ecosystem structure to be determined by long-term forces such as ecological
(plant) succession and fluctuations in climate, and by the more immediate effects of disturbance
events.

Individual communities that have reached a stable condition (the climax endpoint of plant
succession) are rare although a mosaic of different communities at a landscape scale can exhibit
a stable frequency distribution of vegetative states. For example, many landscapes exist as a
shifting mosaic of patch types (Bormann and Likens, 1979). This concept suggests that an
overall balance of patch births and deaths can produce a dynamic equilibrium – there is local
change, but the total number of young and old communities remains relatively constant in
response to disturbance processes and plant succession (Powell, 2000).

With the advent of the non-equilibrium paradigm, the emphasis has shifted from viewing
disturbance as a rare and unpredictable event to treating it as a natural process operating at a
variety of spatial and temporal scales (Wu and Loucks, 1995). Non-equilibrium conditions are
particularly common when the spatial scale of a disturbance process approaches or exceeds the
size of a typical landscape unit. In areas where landscape-scale fires, windstorms (hurricanes),
disease epidemics, or insect outbreaks occur regularly, forests are unlikely to reach a steady
state.

The non-equilibrium paradigm and its associated metaphor, ‘the flux of nature’ (Pickett and
Ostfeld, 1995), are founded on the assumption that ecosystems possess a limited capacity for
internal regulation (Wu and Loucks, 1995). This implies that the behaviour of non-equilibrium
systems is more vulnerable to external disturbances, compared with those of equilibrium
systems. Consequently, the behaviour of non-equilibrium systems is characterized as more
dynamic and less predictable than equilibrium systems (Powell, 2000). The non-equilibrium
paradigm emphasizes ‘event-driven’ vegetation dynamics, which implies that the greatest
potential for vegetation change is associated with the occurrence of periodic and often stochastic
climatic events However, the non-equilibrium paradigm does not imply that ecosystem
behaviour is unconstrained by functional, historical or evolutionary limits (Powell, 2000).

In summary, the non-equilibrium model is based on the following principles (Cook, 1996,
Pickett and Ostfeld, 1995):

 Disturbance is frequent enough to exert a significant influence on vegetation dynamics, with

the result that many ecosystems will never reach a climax state.
 Random influences (chance events) play a significant role in plant succession.
 Life history characteristics (see appendix two) can have a direct bearing on plant succession,
causing it to vary whenever the species that occupy a disturbed area change.
 Different ecological mechanisms may drive plant succession on closely related sites or at
different points during a sere’s development.
References

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