Food web structure and interactions in reservoir ecosystem for fisheries

management
Preetha Panikkar and M. Feroz Khan
Regional Centre, Central Inland Fisheries Research Institute
Hesaraghatta Lake Post, Bangalore 560089

Reservoir ecosystem includes a multitude of organisms occupying different

trophic levels which are interconnected into a complex food web. Food web structure and
interactions have a decisive role in determining the dynamics of an ecosystem.
Management of reservoir fisheries involves a deep understanding of the ecology of the
system and therefore should begin with food web structure. Food webs are temporally
variable in reservoirs due to variation in biota, ontogenic change, feeding strategies of
constituent species and hydrographic parameters.

Food web networks require information on the standing stocks of the living and
non-living constituents, rates of primary production, rates of consumption and production
of heterotrophs, diet of heterotrophs as well as rates of transfer between the various
entities in the food web. In simple ecosystems, energy may flow along a simple food
chain consisting of a primary producer (plants), a primary consumer, a secondary
consumer, ... and a top predator. In most ecosystems, however, the linkages among
organisms are more complex and form a food web.

Organisms represented in food chains

Primary productivity of reservoirs is usually dominated by phytoplankton, which

are harvested by primary consumers such as zooplankton, benthic invertebrates, and
filter-feeding fishes. Another energy pathway comes from organic detritus, which is
consumed by detritivores such as bottom-feeding or filter-feeding invertebrates and
fishes. At a higher trophic level, secondary consumers like predatory invertebrates and
fishes feed on zooplankton and benthic invertebrates. At the top of the aquatic food web
are the large piscivores (fish that eat other fish), aquatic fish eating birds and aquatic
reptiles. In reservoirs, productivity is usually high after initial basin inundation, but
stabilizes at lower levels after 5-10 years. However, microbial food webs may be more
important than previously believed in the carbon cycle of lakes and reservoirs and in the
flow of energy from producers to the top predators

Every known food chain begins with a type of autotroph, whether it is a plant or
some kind of unicellular organism. Primary producers, commonly forming autotrophs,
produce complex organic substances (essentially "food") from an energy source and
materials. These organisms are typically photosynthetic plants, which use sunlight as
their energy source. Organisms that get their energy by organic substances are called
heterotrophs. Heterotrophs include herbivores, which obtain their energy by consuming
live plants; carnivores, which obtain energy from eating live animals. Ultimately
detritivores, scavengers and decomposers may predate living or consume dead biomass.

Organisms are connected to the organisms they consume by lines representing the
direction of organism or energy transfer. It also shows how the energy from the producer
is given to the consumer. Typically a food chain or food web refers to a graph where only
connections are recorded, and a food network or ecosystem network refers to a network
where the connections are given weights representing the quantity of nutrients or energy
being transferred.

Trophic Levels and Food Chains

A food chain is the flow of energy from one organism to the next and to the next
and so on. Organisms in a food chain are grouped into trophic levels, based on how many
links they are removed from the primary producers. The biotic elements that comprise an
ecosystem fall into one of several trophic levels. The trophic level of an organism is its
position in a food chain, the sequence of consumption and energy transfer through the
environment. Trophic levels may contain either a single species or a group of species that
are presumed to share both predators and prey. They usually start with a primary
producer and end with a carnivore. For example, a simple grazing food chain is
comprised of

Primary producer → herbivore → carnivore

At the base of the food chain lie the primary producers. Primary producers like
the phytoplankton and macrophytes in a reservoir ecosystem belong to the first trophic
level. They convert solar energy into organic energy. Herbivores belong to the second
trophic level. Carnivores, predators feeding upon the herbivores, belong to the third.
Omnivores belong to both the second and third. Secondary carnivores, which are
predators that feed on other predators, belong to the fourth trophic level. As the trophic
levels rise, the predators become fewer, larger, fiercer and more agile. At the second and
higher levels, decomposers of the available materials function as herbivores or carnivores
depending on whether their food is plant or animal material.

Food web

A food web is a set of interconnected food chains by which energy and materials
circulate within an ecosystem. The food web is divided into two broad categories: the
grazing web, which typically begins with green plants, algae, or photosynthesizing
plankton, and the detrital web, which begins with organic debris. These webs are made
up of individual food chains. In a grazing web, materials typically pass from plants to
plant eaters to flesh eaters. In a detrital web, materials pass from plant and animal matter
to bacteria and fungi (decomposers), then to detritivores, and then to their predators
(carnivores). Generally, many interconnections exist within food webs.
 A food web extends the food chain concept from a simple linear pathway to a
complex network of interactions. This web makes it possible to show much bigger
animals (like a aquatic birds, catfishes) eating very small organisms (like plankton). Food
sources of most species in an ecosystem are much more diverse, resulting in a complex
web of relationships.

Trophic pyramids

It is often the case that the biomass of each trophic level decreases from the base
of the chain to the top. This is because energy is lost to the environment with each
transfer. On average, only 10% of the organism's energy is passed on to its predator. The
other 90% is used for the organism's life processes or is lost as heat to the environment.
Graphic representations of the biomass or productivity at each tropic level are called
trophic pyramids.

Some producers, especially phytoplankton, are so productive and have such a

high turnover rate that they can actually support a larger biomass of grazers. This is
called an inverted pyramid, and can occur when consumers live longer and grow more
slowly than the organisms they consume.

Energy enters the food chain from the sun. Some energy and/or biomass is lost at
each stage of the food chain as; faeces (solid waste), movement energy and heat energy
(especially by warm-blooded creatures). Therefore, only a small amount of energy and
biomass is incorporated into the consumer's body and transferred to the next feeding
level, thus showing a Pyramid of Biomass.

A pyramid of numbers shows the number of consumers at each level drops

significantly, so that a single top consumer (e.g. a carnivorous fish) will be supported by
literally millions of separate producers (e.g. Phytoplankton).

Energy flow

Energy flows from one trophic level to another through a series of steps of
consuming and being consumed. In a reservoir ecosystem, the aquatic macrophytes and
phytoplankton use light energy from the sun to manufacture carbohydrates for their own
needs. Most of this chemical energy is processed in metabolism and dissipated as heat in
respiration. The remaining energy is converted to biomass. Ultimately, this material,
which is stored energy, is transferred to the second trophic level, which comprises
grazing herbivores, decomposers, and detrital feeders. Decomposers and detritivores
utilize energy from wastes or dead organisms, and so complete the cycle by returning
nutrients to the soil or water, and carbon dioxide to the air and water. Most of the energy
assimilated at the second trophic level is again lost as heat in respiration; a fraction
becomes new biomass. Organisms in each trophic level pass on as biomass much less
energy than they receive. Thus, the more steps between producer and final consumer, the
less energy remains available. Seldom are there more than four links, or five levels, in a
food web. Eventually, all energy flowing through the trophic levels is dissipated as heat.
The process whereby energy loses its capacity to do work is called entropy.

Prey - Predator controls

Bottom up control in ecosystems refers to ecosystems in which the nutrient supply

and productivity and type of primary producers (plants and phytoplankton) control the
ecosystem structure. An example would be how plankton populations are controlled by
the availability of nutrients. Generally bottom up control exists in reservoir ecosystems.
Plankton populations tend to be higher and more complex in areas where upwelling
brings nutrients to the surface. In bottom up control the amount of food consumed is
linearly proportional to prey biomass and independent of predator biomass.

In ecology, top down control refers to when a top predator controls the
structure/population dynamics of the ecosystem. The amount of food consumed is
proportional to the product of the prey and predator abundances The classic example is of
kelp forest ecosystems. In such ecosystems, sea otters are a keystone predator. They prey
on urchins which in turn eat kelp. When otters are removed, urchin populations grow and
reduce the kelp forest creating urchin barrens. In other words, such ecosystems are not
controlled by productivity of the kelp but rather a top predator.

There are many different examples of these concepts. It is not uncommon for
populations to be influenced by both types of control.

Computing food web characteristics

Ecopath software can be used to compute parameters and indices corresponding

to the food web characteristics.
System Omnivory Index: The system omnivory index (SOI) is computed as the average
omnivory index of all consumers weighted by the logarithm of each consumer’s
food intake Q (Christensen et al., 2005). It indicates the allocation of predator to
prey interactions linking each trophic level.

Connectance Index: The connectance index (CI) is the ratio between the number of
actual definite trophic associations among all the groups and the theoretical
possible number of connections.

Trophic impact routine: The ’trophic impact routine’ that could be viewed as a
sensitivity analysis (Christensen & Pauly, 1992) and allows the computation of a
predator - prey matrix in which the impact of an increase in biomass for a
predator group is shown for all prey groups has been utilized here in order to
show the short term effect that an increase in the biomass of a group will have on
the biomass of the other groups in this ecosystem.
Electivity index: The index of electivity describes a predator’s preference for prey. It
scales from -1 to 1; where -1 indicates total avoidance of a prey; 0 indicates that a
prey is taken in proportion to its abundance in the ecosystem; and 1 indicates total
preference for a prey.

Niche overlap: The niche overlap shows for each pair of groups in the model how such
pairs overlap with regards to prey and predators. The niche overlap index can be
used to describe various kinds of niche partitioning. An index value close to 0
indicates that two groups have a low resemblance in terms of food consumed and
vice versa for a value close to 1 (Christensen et al., 2005).

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