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.