TORSION IN PROTOPLASM

William Seifriz
From the Botanical Laboratory, University of Pennsylvania, Philadelphia, Pa.

Received August 8, 19.&

Several years ago, in a discussion on the spiral paths followed by

organisms in movement and growth, I expressed the belief that a state
of torsion is established in protoplasm when it is in motion, provided
certain simple conditions are met. These conditions exist in a freely
suspended strand of protoplasm in which flow is only in one direction
at any given moment. A thread of the plasmodium of a myxomycete
satisfies the foregoing requirements.
Experimental verification of the assumption that a state of torsion
exists in protoplasm under specified conditions was accomplished by the
following method. One end of a freely hanging strand of the myxomycete
Physarum polycephalum was affixed, by adhesion, to the under side of a
cover slip. To the lower end of the strand a delicate glass needle was
attached and adjusted to a horizontal position. The needle served as a
pointer to indicate the direction and extent of twist and, therefore, of
torsion in the thread. With the freely hanging protoplasmic thread and
its horizontal needle thus suspended from a cover slip, the slip was
placed in position over a small moist chamber, the protoplasmic thread
hanging within the chamber. The swing of the needle and the flow of
the protoplasm were observed through the microscope.
There would be no evident reason for the swing of the needle, first
in one direction and then in another, were there not an alternating twist
in the protoplasmic thread, and there would be no reason for the twist
were not some force bringing it about. This force is the flow of protoplasm.
Obviously, movement alone is not sufficient. The flow of a liquid in a
plastic tube will bring about a twisting of the tube only if the wall is
spirally wrapped or fluted as in the case of the rifling of a gun barrel.
The protoplasm of the slime mold Physarum is in a constant state of
flow; the movement is first in one direction and then in the opposite
direction. This rhythmic motion of protoplasm has been the subject of
much investigation (9), and is now recognized as a fundamental form of
movement in living matter. The average time of flow in one direction in
Physarum was established by Kamiya (3) at 47.5 seconds.
27
28 WILLIAM SEIFRIZ

When a horizontal needle, which is attached to a freely hanging

thread of protoplasm, is observed through the microscope after a lapse
of time sufficient to permit the dismembered protoplasm to resume
normal streaming, the needle will be seen to swing rhythmically to the
right and to the left. Many irregular movements occur. These are due
to disturbances caused by severance of the strand, support of a needle,
and readjustment necessary when so small an amount of protoplasm is
subjected to such extreme conditions. But even more significant than
experimental factors in causing irregular movements are the disturbances
frequently occurring in healthy plasmodia. The 45 to 50 second rhythm
in protoplasmic flow is maintained only when the slime mold is progressing
continuously in one dierection. Change in direction of locomotion, or
change in the internal physiological state of the plasmodium is sufficient
to cause an upset in the normal rhythm of progression. There are also
interference phenomena so well analyzed in terms of harmonics by
Kamiya (3). In spite of these disturbances there is a surprising regularity
in protoplasmic flow and, therefore, in twist and torsion of the proto-
plasmic strand, as revealed by the swing of the needle. This rhythm is
shown in Fig. 1.
The experimental work here reported progressed in two stages. There
was first the direct observation that what had been predicted on the basis
of theory was in fact true. A state of torsion is set up in protoplasm when
that protoplasm is in a state of flow. This fact was then placed on a
quantitative basis.
Quantitative observations necessitated somewhat more intricate
technique. Means were needed by which the precise degree of swing of
the needle could be measured at stated intervals. To accomplish this,
the moist chamber containing the hanging thread of protoplasm with
its attached needle was placed on a revolving microscope stage. The
needle was kept in one position by rotating the stage, first to the right
and then to the left, as the needle swung left and right. The vernier
readings were noted at convenient intervals of time, including always
the moment of reversal in direction of swing. The latter readings are
important because they establish the points of return, the peaks and
troughs of the curve. Intermediate readings determine the shape of the
curve and reveal unexpected irregularities.
The curve in Fig. 1 presents five significant facts: (1) that there is a
rhythmic twist in the protoplasmic thread; (2) that the amplitude of
the arc over which the needle travels attains a maximum of 60’; (3) that
the swing is always slightly greater in one direction than in the other;
(4) that there are irregularities in movement which come in groups
between periods of perfect rhythm; and (5) that the beat is 25 seconds,
exactly half of the expected 50 seconds. Of the foregoing facts, the first,
 10 15 16 17 I6 19 20 21

FIG. 1
Graph of the Torsion Rhythm in Protoplasm.
The amplitude in degrees, and the direction in twist of the protoplasmic thread,
are plotted as ordinates against time as abscissae.
29
30 WILLIAM SEIFRIZ

that a state of torsion is set up in moving protoplasm, was my primary

interest. Once this prediction was experimentally verified, interest then
centered in the greater swing in one direction, and the relation between
50 second periods of flow and 25 second periods of twist.
That there is a progressive increase in the amplitude of swing as the
protoplasm flows first in one direction and then in the other in its shuttle
movement, is shown by the gradual incline and decline in the peaks of
the curve (Fig. 1). The protoplasm, therefore, increasingly tightens,
then loosens, its spiral twist. The first portion of Fig. 1 shows a downward
trend of the curve peaks; the right half of the central portion and the
last third of the curve show the same effect in the opposite direction.
They indicate a greater swing in one direction at each pulsation.
The left half of the central portion of the curve is a period during
which irregularities in movement occur due either to physical or physio-
logical readjustments, or to interference in the several rhythms known to
occur in a plasmodium (3). It is these perfectly normal disturbances, as
well as abnormal experimental ones, which frequently upset the rhythm.
During the preliminary experiments attempts were made to establish
a correlation between the rhythm in swing of the needle and the time of
protoplasmic flow but without success, for there were too many conflicting
observations. Swing of the needle to the right and to the left while the
protoplasm continued to flow in the Same direction was frequently
observed but only later understood. That twist and torsion are due to
protoplasmic movement seemed inevitable, and it also appeared likely
that a swing to the right would occur when flow is in one, up or down,
direction, with a swing to the left when flow is in the opposite direction.
The difficulty was a matter of technique. There is also a disturbing lag in
the swing of the needle. The lags were frequently recorded, thus confusing
the synchronism between flow and twist. These difficulties were later
mastered and the cause of the apparent lack of synchronism understood.
The tidal flow of protoplasm is a shuttle movement. If such a move-
ment is the energy source of a mechanism, then the force which it can
exert is greatest midway between the two terminals, for the speed is
greatest there. A tidal flow will, consequently, exert an ever increasing
force until its maximum speed is reached at the mid point, and from
there on, though the protoplasm is still flowing in the same direction,
the available force decreases until it reaches zero at the expiration of the
period of flow. The protoplasmic strand will therefore attain maximum
torsion at the mid point. To twist the thread further will require greater
force, but the available energy is now falling off. From the mid point on,
the protoplasm is still moving in the same direction but at an ever
decreasing speed; as a result, the strand will unwrap, the torsion which
 PROTOPLASM TORSION 31

was set up now operating against the decreasing force of protoplasmic

flow.
The following two groups of observations indicate what takes place.

Direction of
protoplasmic flow: swing of needle
down left
up right
up left
down right

down left
down right
up left
up right
It will be noticed that at each reversal in direction of flow, there is
a corresponding reversal in direction of awing of the needle, but there is
also a reversal in awing of the needle while flow continues in one direction.

THEORY

The thought which led directly to the foregoing experimental proof

of a state of torsion in protoplasm was baaed on the knowledge that there
are many spiral tendencies in animate nature, revealed in the shape,
growth, and locomotion of a great variety of organisms and tissues (7).
Asymmetry in form and movement occura in men, trees (8), protozoa (6)
and chromosomes (3). From the work of Aatbury (l), Mark (4), Meyer
(5) and others on the molecular structure of elastic systems, it is possible
to interpret the spiral tendencies of living matter in terms of molecular
configuration. Obviously, not all the constituents of living matter need
have a spiral or asymmetric orientation, but only the fibrous, the struc-
tural proteins (2) ; nor need all macro-spirals in nature be correlatid
with micro-spirals or molecular asymmetry. However, every attempt to
explain the twist in tree trunks in terms of external forces rather than
inner molecular structure proved to be wholly unsuccessful (8). A mo-
lecular interpretation of the spiral form and movement of organisms is
in keeping with modern chemical interpretations of the structure of
organic matter, particularly elastic material. Outward spiral form and
movement are due either to asymmetric molecules or to a spiral orienta-
tion of molecules.
A molecular interpretation, though convincing, in no way affects the
experimental fact that a state of torsion is established in flowing proto-
plasm.
32 WILLIAM SEIFRIZ

SUMMARY

1. A delicate glass needle suspended in a horizontal position on the

end of a freely hanging strand of myxomycete protoplasm swings alter-
nately to the right and to the left through an arc of about 40” when the
protoplasm is in a state of flow.
2. The swing of the needle is the result of a twisting of the living
thread, brought about by streaming of the protoplasm.
3. As a result of the spiral twist produced by streaming, a state of
torsion is set up in the protoplasm.
4. The period of swing of the needle is half the period of rhythmic
flow in. one direction.
5. That the period of swing is but one-half the period of flow is due
to a gradually increasing force brought about by increase in rate of flow
up to the mid point of movement in one direction, after which there is a
gradual lessening in the force applied and a consequent unwrapping of
the protoplasmic thread.
6. A molecular interpretation of the structural basis of spiral form
and movement in protoplasm is given, based on the known asymmetry
of molecules.

REFERENCES

1. ASTBURY, W. T., Ann. Rev. Biochem. 8, 113 (1939); Proc. Roy. Sot. (London) Bl29,
307 (1940).
2. BANGA, I., AND SZENT-GYBRGYT, A., Science 92, 514 (1940).
3. KAMIYA, N., The Structure of Protoplasm (edited by W. Seifria), p. 199, Ames,
Iowa (1942).
4. MARK, H., Chem. Rev. 23, 121 (1939).
5. MEYER, K. H., Die hochpolymeren Verbindungen, IRipzig (1940).
6. SCHAEFFER, A. A., Anut. Record 34, 115 (1927).
7. SEIFRIZ, W., Science 77, 50 (1933).
8. SEIPRIZ, W., Science 78, 361 (1933).
9. SEIFRIZ, W., Botan. Rev. 9, 49 (1943).