Geotropism. An introduction.

By
Poul Larsen.
With 28 figures.

Historical sketch.
As pointed out by SACHS! (1875, 1906) the most conspicuous plant move-
ments were noted already by ancient authors. In the early days of scientific
botany the orientation of roots, main stems, and branches, on the other hand,
seems to have been regarded more as a problem in morphology than in physio-
logy, and the quiescent behavior of these organs evidently has not given rise to
any impression of movement until fairly late in the history of plant science. When
movements in these and other plant parts became recognized, it was also dis-
covered that they could be brought forth by a number of external (and internal)
factors. One of these, when expressed in the most general way, is exposure
under certain angles to a field of mass acceleration. Gravity is the natural repre-
sentative of stimuli of this type, and the best known responses provoked by
such stimuli are the geotropic movements.
The term geotropism was introduced by FRANK (1868) in analogy with the
term heliotropism introduced by DE CANDOLLE in 1832. When "heliotropism"
was later superseded by "phototropism" , "geotropism" should logically have
been replaced by "barytropism" (Greek barytes: weight), but the term geo-
tropism has remained in use till our day.
Most of the history of geotropism before SACHS is contained in works which
are not easily accessible, and the literature before KNIGHT does not seem to
have been reviewed in the last 50 years. The following brief review of the early
history of geotropism is based on information derived mainly from the works
of ELFVING (1879), PFEFFER (1904, 1906, 1912), RAWITSCHER (1932), SACHS
(1875,1906) and ZIMMERMANN (1927b). In most cases, unfortunately, the writer
has not been in a position to consult the originalliterature ; and most of
the references, pertaining to the early history are cited from the exhaustive
bibliography by MARIE CHRISTIANSEN (1917-1929). Other surveys of the early
studies on geotropism are those by CIESIELSKI (1871) and SCHOBER (1899).
The first known treatment of geotropic reactions as a scientific problem is
credited to DODART (1703). He made systematic investigations on the direction
of growth of roots and shoots of bean (Phaseolus) seedlings derived from seeds
which were laid out to germinate with their micropyle pointing in various direc-
tions. He noted that root and shoot oriented themselves in a verticalline regardless
of the orientation of the seed. The same orientation was reached in air and in
soil. DODART made similar experiments with Quercus, and in seedlings of this
plant he noted the sharp curvature produced by a root which was first allowed
to grow in the normal direction and then inverted. He also noted the negative
geotropic curvatures in shoots of trees and herbs which had been brought out
of their normal position.
1 See also BÜNNING: Einführung und Übersicht, this volume, Part 1, p. 1.
L. Aletsee et al. (eds.), Physiology of Movements / Physiologie der Bewegungen
© Springer-Verlag OHG / Berlin · Göttingen · Heidelberg 1962
 Historical sketch. 35

DODAR'I' admits bis ineapaeity to indieate the eauses of these movements,

but, hesitantly, suggests a hypothesis : desieeation of the upper side and moistening
of the lower side of the aerial parts of plants might lead to eontraetion of the
fibers of the upper side of the organ and extension of those on the lower side.
For roots, the reverse might then be the ease.
ASTRUC (1709) seems to be the first to suggest that gravity had something
to do with the upward curvatures of aerial parts of plants. He assumed that
the plant juice on account of its density was transported predominantly on
the lower side of such plant parts when not in the vertical position. The lower
side would thus receive better nourishment than the upper side and thereby
grow faster.
A rather peculiar concept was expressed by DE LA HmE (1709). He imagined
that the sap in stems was light ("plus fin et plus volatile") and would, therefore,
ascend in the plant. The growth of the plant part should then follow the ascending
juice. The sap in roots was assumed to be of high specific weight, a property
which presumably guided the growth of these organs. The downward growth
of roots was thus regarded as a passive bending, an idea later adopted by BAZIN
(1741) and several other physiologists.
DUHAMEL DE MONCEAU (1758) rejected DODART'S hypothesis on basis of
experiments with seedlings accomodated in surroundings of uniform moisture
conditions (moist sponges, moist soil), so that unilateral desiccation or moistening
was avoided. Against ASTRUC'S ideas, DUHAMEL points to the fact that the
tip of an inverted main stem will perform an upward curvature although its
nutrient sap must be uniformly distributed at the start of the experiment.
DE LA HillE was dismissed with aremark on the speculative character of his ideas.
Admitting the futility of attempting to find the cause of the different diree-
tions of growth of roots and stems on basis of the knowledge of his day, DUHAMEL
reeommended experimental studies of the conditions under whieh the various
geotropie reactions oeeur. Without formulating any new hypothesis he expresses
his inclination to plaee great signifieanee in the role of vapors inside and outside
the plant.
Real progress in the study of the eauses of the eharaeteristie orientation
of stems and roots was not made until KNIGHT (1806) earried out his famous
experiments. KNIGHT referred to suggestions by eertain philosophers (presumably
ASTRUC, DE LA HmE and BAZIN, see above) that gravity might be the faetor
responsible for the orientation of these organs. KNIGHT was the first to eonclude
that if this is true, a eentrifugal force should have the same effeet as gravity,
and he proeeeded to prove this by experiment.
In his paper (1806) KNIGHT deseribes how he constructs a water-wheel that
operates a seeond wheel revolving in a vertieal plane. The diameter of this
wheel was 11 inches. Several water-imbibed seeds of garden bean (Phaseolus )
were fixed to the rim of the wheel, the embryonie radieies pointing in various
direetions in spaee. The wheel with the seeds was eovered by a box and pro-
tected by wire-netting. The water was admitted, and the wheel made more
than 150 turns per minute. KNIGHT remarks that the [unilateral] effeet of gravity
on the plants must have been completely eliminated under these eonditions.
When the seeds started to germinate, KNIGHT had the pleasure to note that
all the roots, regardless of their initial orientation, pointed their tips away from
the rim of the wheel, growing at right angles to its axis. The stems, on the eon-
trary, grew in the opposite direetion, and after a few days the stem tips inter-
twined around the hub. In this experiment, the centrifugal force was about
3.5 times gravity or stronger.
Handbnch d. Pflanzenphysiologie, Bd. XVIJ/2. 3a