In Elton's account of the natural community as a simplified economy he suggested the
following
four principles that could describe the economy of nature as it operates everywhere on earth.
Elton called first of his principles the "food chain."
In every community, plants, through the photosynthetic conversion of sunlight to food,
form the first
link in a chain of nutrition. Food, one might say, is the essential capital in the natural economic
order. The remaining links- usually no more than two or three, and almost never more than four-
include the herbivorous animals and their predators.
A typical food chain in North American oak woods might link acorns, quail, and foxes,
or acorns,
mice, and weasels; with some 200 species of birds and mammals alone feeding on the oaks. The
potential number of food chains is extraordinarily large. They could be found in nature by the
thousands, all showing a common pattern but no two alike in every respect. And the bottom of the
chain, rather than the top, is the most important link: The plants make the whole system possible.
Elton referred to the sum total of chains in any community as the "food web",
an exceedingly
complex design of crisscrossed lines of economic activity. Such webs are easiest to analyze in the
relatively unpopulated arctic zones and almost impossible to untangle in the warm, humid tropics,
where life forms abound. In every food chain certain roles must be performed. The plants, for
example, are all "producers." Animals can be described as either first- or second-order
"consumers," depending on whether they eat plants or other animals. Those animals that feed
on
the most numerous plants in a habitat, like the bison on prairie grasses, or copepods on diatoms in
the sea, are the "key industries" in those economies.
In 1926, August Thienemann had introduced the terms "producer," "consumer,"
and "reducer," or
"decomposer," which describe economic roles of organisms in a specific ecological setting;
Elton
now generalised them for every food chain in nature. These labels emphasized the nutritional
interdependence that binds species together, and they became the boxes of resources with which
ecologists would increasingly build models of materials and energy flows through ecosystems.
In general, an animal as large as an elephant cannot survive on food as small and
lively as insects.
It would exhaust itself in the chase and need more food than it could catch. A more substantial,
stationary, and unresisting kind of food, the leaves, twigs and outer branches of trees and shrubs,
is called for.
The giant whale is able to live on tiny crustaceans only because they are so numerous
and easy to
harvest. The law of nature decrees that each species has an optimum food size, and this law
determines the structure of the food chain. "The very existence of food-chains," Elton noted,
"is due
mainly to the fact that any one animal can only live on food of a certain size. Each stage in an
ordinary food chain has the effect of making a smaller food into a larger one, and so making it
available to a larger animal." The chain thus becomes an ascending scale of larger and larger
mouths and stomachs, except where a predator has the advantage of such special weapons as
poison, like the spider, or group tactics, like the wolf pack. Even then, of course, the spider cannot
kill and consume an elephant.
The only species that ignores these rules is civilised man, with his artificial techniques
for more
efficient food gathering. He can kill the largest animals on earth, or he can gather the smallest grain
and seeds, and so eat lower on the food chain. But for Elton, modern man is distinctly an outsider,
not to be confused with the mainstream economy of nature and its workings.
To serve as food for organisms higher up on the food chain, plants and animals near
the bottom
must be more numerous and reproduce more rapidly. Their fertility is a function of their position in
the economic order: The smaller a creature, the more common it is and the faster it reproduces. In
contrast predators at the top of the chain must reproduce more slowly than their prey, or they will
end up with nothing to eat. They must also distribute themselves more sparsely across the land;
hence every tiger looks for his own hill and defends it against other tigers, or he perishes. The
demarcation and defence of private territories for food or breeding insures that each successful
individual will have an adequate base for survival. Such territorialism goes on where it must among
the birds and mammals, which are constantly required to adjust their density to their food supply.
The system of interacting populations resulting from these different behavioural requirements
Elton
called the "pyramid of numbers." A single plot of ground may provide a home for millions of
microscopic soil bacteria, thousands of grass plants and insects, hundreds of trees, a few dozen
rabbits, sparrows, and squirrels and at the apex only a single hawk. As one goes up the chain, the
total weight of protoplasm at each level (the "biomass") declines along with the numbers.
One lion,
for instance, may eat fifty zebras in a year, so that the total physical mass of the predator is but
a
small fraction of the combined weight of its prey. The plants at the food chain's base constitute by
far the greatest part of the mass of living substance on earth.