Food chains and trophic levels
The transfer of food energy from the
source in plants through a series of organisms with repeated stages of eating
and being eating and being eaten is known as the food chain. In complex natural
communities, organisms whose food is obtained from plants by the same number of
steps are said to belong to the same trophic level (troph =nourishment, the
same root as in autotrophic and heterctrophic). Thus, green plants occupy the
first trophic level (the producer level); plant eaters (herbivores, and so on)
the second level (the primary consumer level); carnivores that eat the
herbivores the third level (secondary consumers), and perhaps even a fourth
level (tertiary consumers). It should be emphasized that this trophic
classification is one of function, and not of species as such; a given spccupy
one, or more than one, trophic level according to the source of energy actually
assimilated. We have already called attention to certain algae that may depend
in part on their own food and in part on food made by other algae; or the populations
of men who utilize food from both plant and animal sources.
Figure 3 -2 is an energy flow diagram of
a food chain. This is a more detailed rendition of the last three modules in
figure 3 -1. In this model we introduce the biological terminology used to
describe come munity metabolism – and the quantities shown are on a daily,
rather than annual, basis. The boxes represent the population mass or biomass,
and the pipes depict the flow of energy between the living units. As already
indicated, about half of the average sunlight impinging upon green plants (that
is, producers) is absorbed by leaves and about 1 to 5 percent is converted into
food energy by productive vegetation. The total assimilation rate of producers
in an ecosystem is designated as primary production or primary productivity (pg or A in figure3 – 2). It is
the total amount of organic matter fixed, including that used up by plant
respiration during the measurement period. Net primary productivity (P n ) is the organic matter
stored in plant tissues in excess of respiration during the period of
measurement. Net production represents food potentially available to heterotrophy.
When plants are growing rapidly under favorable light and temperature
conditions, plant respiration may require as little as 10 percent of gross
production so that net production may be 90 percent of gross. However, under
most conditions in nature net production is a smaller percentage of gross,
usually about 50 percent, as shown in the lower line of figure 3 – 2.
As shown in figure 3 – 2 part of net
primary production may be stored or exported, and part may beome an energy
source for heterotrophy. NU and I, are apportioned within different kinds of
ecosystems. Some portion of food ingested by consumers is usually not
digestible or assimilable, so some of the energy is likely to be egested unused
(NA in the diagram); this component may be stored, exported, or consumed by
microorganisms or other heterotrophy. Consumers as well as producers must
respire (R) a large part of the energy assimilated (A) so as to maintain
structure and function (represented by the boxes in the diagram). Respiratory degradation
of energy ‘’pumps out’’, that is, reduces entropy as is required to maintain a
high level of organization. The R flows in figure 3 – 2 represent heat losses
from biological components that were shown by the more generalized heat sink
symbol in figure 3 – 1. Assimilated energy not respired is available for
production (Pn P2 P3 …….) which can take the from of growth of new tissue
and reproduction and growth of individuals (population growth). Production at
heterotrophic levels is often known as secondary production to distinguish it
from the primary production of plants. Any auxiliary energy or nutrients that reduces
the cost of maintenance (respiration) enhances the rate of production. If the
autotrophic level is subsidized, the increased production may be passed along
the chain; if a consumer level is augmented, the ‘’downstream’’ links will be
mostly affected, although secondary production can ‘’loop back’’ upstream so
that even plants might benefit from organic matter not used by animals.
Roughly
speaking, the reduction of available energy with each link in the food chain
(as required by the second law of thermodynamics ) is about two orders of
magnitude at the first or primary trophic level, about one order of magnitude
thereafter. By order of magnitude we mean by a factor of 10. If an average of
1500 kcal of light energy were absorbed by green plants per square meter per
day, we might expect 15 to end up at net plant production, 1.5 to be
reconstituted as primary consumers (herbivores), and 0.3 as secondary consumers
(carnivores) – provided, of course, that there are adapted organisms present that
can fully utilize these resources. Efficiencies in terms of percent energy
transfer are thus on the order of 1 percent at the first level and 10 – 20 percent
at the heterotrophic levels. Since meat is generally of higher nutritional
quality, percent transfer tends to be higher at meat – eating levels. Since so
much energy is lost at each transfer the amount of food remaining after two or
three successive transfers is so small that few organisms could be supported if
supported if they had to depend entirely on food available at the end of a long
food chain. For all practical purposes, then, the food chain is limited to three
or four ‘’links’’. The shorter the food chain, or the nearer the organism to
the beginning of the food chain, the greater the available food energy. About ten
times more people can be supported by 100 acres of corn of they function as
primary rather than secondary consumers, that is, if they eat the corn directly
rather than feed if to animals and then eat meat. If you prefer to eat meat
then you must plan not to let the population become so dense as to preclude the
meat – eating option. In debating man’s’’ world food problem’’ we must consider
the quality as well as the amount that might be obtained from a given area of
land or water.
It should be emphasized that the scheme
in figure 3 – 2 is a model useful for making comparisons with the real thing. Somewhat
larger, and frequently quite a bit smaller, percentages may actually be
involved under different conditions. Much remains to be learned, not only about
the orders of magnitude in different
ecosystems but also about the upper limits. Since the efficiencies of transfer
seem low in terms of man – made machines, man has often thought that he could
improve on nature by increasing the percent of transfer of light to food, and
food to consumer. However, when we consider the low quality of available
energy, the fact that organisms unlike machines are self – maintaining, and the
need for storage and diversity for future survival than it turns that nature’s
efficiencies are just about optimum. (for further discussion of this important
point, see odum and pinkerton 1955; H.T.odum, 1971.
At this point it would be well to make
clear a few points about units, ratios, and efficiencies. Too often our
thinking is clouded by people who, unintentionally or intentionally, use units
that are not comparable in forming ratios and percentages. Biomass varies in
energy value, both quantitatively and qualitatively. Plant biomass runs about
4.5 kcal per ash – free dry gram, and animal close to 5.5 where energy is being
stored, as in seeds or in bodies os migrating or hibernating animals; the
values approach 7 pr 8 kcal. If plant matter is mostly cellulose and lignin
(wood), its energy (4 kcal/g) is unavailable to most animals. The main point to
remember about ratios and percentages when used as estimates of efficiency is
that they should be dimensionless; that is, the same unit should be used for
both the denominator and numerator of the ratio. This, it should be calories/ calories
not calories/grams. Otherwise comparisons may not be valid. For example,
poultry or catfish farmers may tell you they are getting one pound of meat for
every two pounds of feed, implying a 50 percent efficiency of transfer. However,
since the food Is highly concentrated, dry material worth 5 or more kcal/g, and
the meat s ‘’wet weight,’’ worth only about 2 kcal/g, then the real energy out/
energy in efficiency is more on the order of 20 percent.
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