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Food chains and food webs--
The pathways whereby the energy and matter produced
by plants are distributed among other organisms in a habitat are known
as food webs. Food webs are made up of a network of food chains each of
which connects an autotroph, at the lowest trophic (i.e., feeding) level,
to the herbivores (i.e., primary consumers) and then to the various carnivores
(i.e., secondary consumers, tertiary consumers, and so on).
The efficiency with which energy is transferred from one level to the
next determines the length of food chains. Each step in the food chain
involves loss of efficiency due to the costs of capturing and digesting
food. As a rough approximation, the efficiency of conversion from one
level to another is 10%. (The exact value depends on the P/R ratio and
can be quite variable. It is generally lower in vertebrates than invertebrates,
and lower in birds and mammals than in fishes.) This means that it takes
about 100 grams of plants to create ten grams of herbivore; when eaten,
that ten grams of herbivore will result in one gram of carnivore. Because
of the inefficiency of energy transfer (due to the expenditure of energy
on metabolism), food chains tend to be limited in length; that, with three
levels of carnivores, is likely to be the longest food chain sustainable
in many habitats.
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In some circumstances, much of the plant material is
not eaten by herbivores. This may be because it cannot be eaten (most
herbivores cannot digest wood), or because the herbivore population is
kept low by secondary consumers. In that case the plant (or its parts)
dies, and is colonized by the microorganisms responsible for decay forming
detritus (i.e., particulate organic material derived from the decomposition
of plant and animal remains). Detritus provides food for a range of consumers
(called detritivores) and a food chain results. In most habitats, the
detrital food chains and those chains based on herbivores eating live
plants combine to produce a food web in which the animals at higher levels
eat elements from both types of chain. These webs can be very complex,
such as in the Mekong River where a large number of interacting species
are present.
The origins of organic material--
The relative amounts of energy passing through the autotroph to herbivore
pathway and through the detritus to detritivore pathway will vary among
habitats or different parts of a river. Some indication of their relative
importance can be estimated from if we know the origin of organic material
in the environment. Autochthonous material originates from within the
aquatic ecosystem; for example, primary production by algae.
Allochthonous material originates outside and is transported into the
aquatic ecosystem; for example, tree leaves that fall into rivers.
In general, most of the organic material in small streams
is allochthonous, originating from the overhanging terrestrial vegetation
or washed in from the surrounding forest floor. There is some autochthonous
primary production by periphyton (see Section 6) but shading by riparian
vegetation may limit the light needed for photosynthesis. Macrophytes
are scarce. In general, therefore, autochthonous primary production is
estimated to be of relatively minor importance (for details, see Cummins
1974)
The amount of allochthonous input varies greatly in
space and time, but inputs of around 1 kg/m2/yr
in the upstream course of shaded rivers can be expected. The timing of
inputs will depend on the schedule of leaf loss and replacement of terrestrial
plants. There is less seasonality in this regard in Southeast Asia than
in the deciduous forests of Northwest Europe and North America.
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The amounts of autochthonous production increase downstream
as the channel widens and shading is reduced (see next section); in larger
rivers, phytoplankton may be present in the main channel and floodplain
pools (see also Section 6). The biomass of detritus is typically less
than upstream, but some decomposing material carried downstream by the
current will be present, regardless of whether there are substantial direct
inputs from the riverbank. The particle size and quality of this transported
detritus may differ considerably from that entering the river from the
surrounding land.
There
is some difficulty in interpreting the origin of the material on floodplains
because of the alternation of aquatic and terrestrial phases over the
flood cycle. Terrestrial material (nutrients and energy) can stimulate
aquatic production by plants and animals during the flood phase. This
is described in more detail under Section on 'Flood Pulse').

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