chemical composition increases the surface tension of the cell sufficiently
to provoke its contraction into a more spherical form.
Ostwald has introduced a very useful conception for the study of this
question of surface energy. A liquid surface contains a quantity of energy
equal to its surface tension multiplied by its area, hence any variation
either of area or of tension corresponds to a variation of its energy. This
novel conception constitutes a valuable addition to the experimental study
of the physiology of muscular action, since it gives us some idea of the
mechanism by which chemical energy may be transformed into muscular
contraction.
Whatever the mechanism of transformation in the animal machine, we have to
consider the same quantities as in other motor machines. These are: (1) the
efficiency; (2) the potential energy; (3) the power; (4) the energy given
up to the medium under the form of heat; (5) the temperature.
Muscles, then, are merely transformers which change chemical energy into
mechanical work, the diminution of stored-up energy in a muscle being
expressed by the sensation of fatigue. A muscle may be studied in four
different phases: (1) in repose; (2) in a state of tension; (3) when doing
positive work; (4) when work is being done on it.
When a muscle is in a state of tension, as when a weight is sustained by
the outstretched arm, the muscle is producing no external work. The entire
work done is converted into heat; just as it is in a dynamo or steam engine
which is prevented from turning by a brake. Muscular contraction produces
fatigue even when it does no external work. It is impossible for the muscle
to support even the weight of the outstretched arm itself for any
considerable time.
A muscle is doing positive work when it is raising a weight or moving a
body from one point to another. {110}
The fourth state of muscular contraction is when the muscle is doing
negative work, _i.e._ when work is being done on it, as for instance when
we go downstairs, or when a descending weight forces down the opposing arm
which attempts to support it. In this case the muscles receive a portion of
the energy lost by the descending weight, and this energy shows itself in
the muscle in the form of heat. This increase of heat in a muscle doing
negative work has been clearly demonstrated by the calorimetric experiments
of Hirn and the thermometric experiments of Beclard. Hirn's observations on
muscular calorimetry show a production of heat corresponding to 150
calories per hour when in repose, 248 calories per hour during positive
work, and 287 during negative work. Beclard's thermometric measurements
also show that the temperature of a muscle rises each time that it
contracts, and that the rise of temperature is greatest when the muscle is
doing negative work, least during positive work, and intermediate when in a
state of tension.
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