Research methods in ecologyClements, Frederic E. (Frederic Edward)
Science
Research methods in ecology
Clements, Frederic E. (Frederic Edward)
Plant ecology; Plant ecology -- Methodology
=321. Succession by exhausting the soil.= This is a reaction not at all
understood as yet in nature. A number of phenomena, such as the “fairy
rings” of mushrooms and other fungi, the peripheral growth and central
decay of lichens, _Lecanora_, _Placodium_, _Parmelia_, and of matforming
grasses, such as _Muhlenbergia_, and the circular advance of the
rootstalk plants, indicate that certain plants at least withdraw much of
the available supply of some essential soil element, and are forced to
move away from the exhausted area. It is probable that the constant
shifting of the individuals of a formation year after year, a phenomenon
to be discussed under alternation, has some connection with this. It
will be impossible to establish such a relation, however, until the
facts are exactly determined by the method of quadrat statistics. So far
as native formations are concerned, there can not be the slightest
question that prairies and forests have existed over the same area for
centuries without impoverishing the soil in the least degree, a
conclusion which is even more certain for the open vegetation of deserts
and plains. With culture formations, the case is quite different. The
exhaustion of the soil by continuous or intensive cultivation is a
matter of common experience in all lands settled for a long period.
Calcium, phosphorus, and nitrogen compounds especially are used up by
crops, and must be supplied artificially. The reason for this difference
in reaction between native and culture formations seems evident. In
harvesting, not merely the grain, but the stems and leaves, and in
gardening often the root also, are removed, so that the plant makes
little or no return to the soil. In nature, annual plants return to the
ground every year all the solid matter of roots, stems, leaves, and
fruits, with the exception of the relatively small number of seeds that
germinate. Perennial herbs return everything but the persistent
underground parts. Shrubs and trees replace annually an immense amount
of material used in leaves and fruits, and sooner or later, by the
gradual decay of the individuals or by the destruction of the whole
formation, they restore all that they have taken from the soil. This
balance is further maintained to an important degree by the activity of
the roots, which take from the deep-seated layers of the soil the crude
materials necessary for the formation of leaves and fruits. Upon the
fall and decay of these, their materials are incorporated with the upper
layers of the formation floor, from which they may be absorbed by the
undergrowth, or find their way again into the layers permeated by the
tree roots. From the universal occurrence of weeds in cultivated
regions, the pioneers in impoverished or exhausted fields are uniformly
ruderal plants. As is well known, the seed production and ecesis of
these forms are such that they take possession quickly and completely,
while their demands upon the soil are of such a nature that the most
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