We owe the word ecosystem to Arthur Tansley who
used it in 1935. It has since become the central organizing
idea in ecology. As a model of interrelatedness in nature, it
presents both the biological and non-biological aspects of the
environment as one entity, with strong emphasis on measuring the
cycling of nutrients and the flow of energy in the system—
whether it be a pond, a forest, or the earth as a
whole.
An ecosystem is defined as the unit of plants and
animals which is functionally integrated through nutrient and
energy fluxes.
Its boundaries are defined as the habitat but the
selection of these limits is more subjective. Within the
International Biological Programme it was conventional to delimit a
type hectare, though the size of the type depended upon vegetation
homogeneity. Only non-destructive measurements, such as
temperature, rainfall or C02 fluxes, were made within
this area. The effects of trampling, a very significant factor in
delicate systems such as the tundra, were confined to strictly
prescribed paths or board walks. The type hectare was surrounded by
an undisturbed buffer zone, the importance of which, again, should
not be underestimated in any ecological studies. Around the outside
of this zone sampling areas were delimited ranging from 1 ha to 10
ha for vegetation and soil analyses to much larger areas for mammal
and bird studies.
The definition of an ecosystem, within this
programme of research, was an area which is self-contained in terms
of its primary production and nutrient cycling. That is to say, the
extent of nutrient fluxes across the boundary was small with
respect to the internal pathways. It is possible to recognize a
pond, lake or the oceans as falling within this definition but
flowing water systems tend to have a major through-flux of
nutrients and little internal production and cycling. Streams and
rivers may therefore be regarded as open ecosystems- or more
strictly, like the soil, as a subsystem of a larger complex . The
precise definition of the ecosystem is thus open to debate but the
principle of the interacting sub- components of the biological
system is the essential nature of the concept.
This holistic concept of ecosystem structure and
functioning was developed through the work of a number of eminent
ecologists in the early part of this century. Charles Elton (1927),
for example, described the biotic structure of ecosystems by
allocating the organisms to a number of broad trophic levels
(plants, herbivores and carnivores). A graphical presentation of
the trophic structure of most ecosystems results in pyramids of
numbers or biomass reflecting the decreasing abundance of organisms
at higher trophic levels, e.g. foxes < mice < grass on a
biomass or density basis per unit area. Subsequently, in 1942,
Raymond Lindeman drew attention to the quantitative relationships
which exist between these different trophic levels in terms of
energy and matter transfers which maintain the functional integrity
of the system. This 'trophic dynamic' concept, as it was called,
stimulated much of the quantitative analysis of ecosystems which is
fundamental to our present understanding of this aspect of
ecology.
A generalized model of ecosystem structure
usually has a minimum of three subsystems, plant, herbivore and
decomposition, and their main compartments or trophic levels. The
functioning of the ecosystem may be described in terms of energy or
nutrient fluxes but both approaches have their value and
limitations. Ecosystem energetics (as well as at community and
population levels) are of particular value for comparing different
systems since plant and animal trophic levels, and their
interactions, can be expressed and compared in a single unit of
energy, the joule or calorie (1J = 0.24 cal). In many situations
where the availability of mineral nutrients, rather than energy,
limits the rate at which the system functions, the investigation of
nutrient flux pathways may be more meaningful. Mineral nutrient
budgets also emphasize the integrity of the ecosystem and the
feedback loops operating within the boundaries which regulate and
maintain this functional integrity.
There have been few complete studies of energy
flux through ecosystems but two classical examples are forest and
marine bay.
A forest ecosystem is dominated by the the large
stock of stored energy of the plant biomass in comparison with the
gross primary production. The energetic cost of maintaining this
biomass is low due to the predominance of metabolically inert woody
tissues. The herbivore biomass is small and the main energy fluxes
are autotroph and decomposer respiration.
The marine bay plankton community has, by
comparison, a minute autotroph biomass supporting a proportionately
higher energy flux through the larger herbivore subsystem. This
phenomenon is characteristic of planktonic communities and is known
as an inverted biomass pyramid.
Carbon and energy are closely linked in the
biosphere, and ecosystem structure and functioning can be similarly
described in terms of carbon fluxes. Carbon flow is a good
illustration of the functional integration of adjacent terrestrial
and aquatic ecosystems. Both of the types of energy budgets
considered above can be seen as integral components of the larger
system. The open nature of the stream system should also be noted
with respect to its definition as an ecosystem.
The following generalizations can be made on the
fate of net primary production in natural ecosystems.
(1) In terrestrial ecosystems part of
the NPP is stored in perennial tissues and contributes to biomass.
In grass dominated or aquatic ecosystems this increment is usually
negligible but in immature forests or plantations it may contribute
20-60 per cent of NPP.
(2) A minor proportion of terrestrial
NPP is consumed by herbivores. Even in intensively grazed
grasslands this fraction rarely exceeds 25 per cent of the total
NPP because, although more than half of the above ground biomass
may be removed, the annual production of roots may be equal to or
exceed above ground production. In forest systems insects or
browsing animals rarely consume more than 10 per cent of the NPP,
while in aquatic systems 80 per cent or more production by
phytoplankton may be consumed by herbivorous zooplankton.
(3) Material not allocated to plant
growth or the herbivore subsystem, together with the remains of the
herbivores, predators and excretory products, enters the decomposer
subsystem. In ecosystems such as mature forests, which no longer
continue to accumulate biomass, a major proportion of net
production is shed as plant litter and is processed by the
decomposers. Fungi and bacteria are the main agents of
decomposition but animals which feed on detritus, fungi and
bacteria, as well as their predators, are important components of
this community. Material is recycled between the decomposer
organisms until carbon mineralization is complete.
The empirical analysis of ecosystem structure
requires the delimitation of its boundaries. Sometimes these are
defined naturally by the habitat but in a region of more or less
homogeneous terrestrial vegetation, or on a vegetation gradient,
the affinities to one another than woodlands. Thus within a biome
we can define a typical assemblage of plant and animal types or a
series of specific assemblages of animal and plant species. The
unit of plants and animals which is functionally integrated through
nutrient and energy fluxes is the ecosystem. The environment
of an ecosystem is formed by adjacent ecosystems with which it
interacts through fluxes of materials across their boundaries. The
measurement of these fluxes can provide considerable insight into
the internal dynamics of the ecosystem and also allow a more
precise definition of its identity.
Within the ecosystem boundary there are three
functional compartments — the green plants (autotroph
subsystem), the animals, together with their predators, which
feed on living plants (the herbivore subsystem) and the
organisms decomposing dead plant and animal remains, together with
their predators (the decomposer subsystem). Nutrient
elements taken up from the soil and air by plants are synthesized
into tissues using light energy (primary production).
The total amount of material and energy fixed by
the autotrophs is called Gross Primary Production
(GPP).
Some proportion of the GPP is respired (R) by the
plants and the remaining Net Primary Production (NPP = GPP
- R) is available to the heterotrophs.
The production of tissues by the growth and
reproduction of heterotrophic organisms in the herbivore and
decomposer subsystems is known as secondary
production.Ultimately all material is processed by the
decomposers and the elements released in a mineral form suitable
for re- utilization by plants. It is useful from an ecological
point of view to distinguish between relatively unavailable
nutrients in a storage pool and the active, exchangeable
reservoir or exchange pool. The capital of nutrient elements
accumulated in living and dead tissues (standing crop, as
distinct from biomass which consists of living material
only) is referred to as the utilized pool. An essential
feature of an ecosystem is that while nutrients are recycled within
its boundaries, energy enters as light and leaves as metabolic heat
in a one-way flux.