Movements and Bioelectrical Events Induced by

Photostimulation in the Primary Pulvinus of Mimosa pudica

G. ROBLIN

University of Poitiers, Station Biologique de Beau-Site, 86000 Poitiers, France

ReceivedJanuary 18, 1982 . Accepted March 26,1982

Summary
It is well known that different stimulating agents can induce spectacular movements of the
primary pulvinus in Mimosa pudica L. A photostimulation can also evoke the same type of
movement. But in many cases the pulvinus reacts with more gradual curvatures. The bioelectri-
cal variations recorded concomitantly with the movements suggest that there is no release of
any «stimulating substance» analogous to that which spread after wounding the leaf.
Key words: Mimosa pudica, pulvinus movement, photostimulation.

Introduction
Foliar motor organs (or «pulvini») in Mimosa pudica L. that form true articulations
can be stimulated by various and numerous agents (for review, see Roblin, 1979).
Besides the rapid leaf movements induced by mechanical, electrical or thermal sti-
muli, it has been observed that the pulvini can react to a sudden increase of light
intensity (a «photostimulation») over periods as brief as 15 seconds (Darwin, 1882;
Bose, 1931; Fondeville, 1963; Roblin and Gavaudan, 1974). When plants are illu-
minated during the scotophase (darkness) of the photoperiodic cycle, some of the
primary pulvini are stimulated to a rapid downward movement. The latent period of
this response generally ranges from 40 to 120 seconds which is a function of
irradiance (Roblin, 1975). It can also be noted that the response shows a rhythmic
pattern, the maximum number of pulvinar reactions being observed in the middle of
the scotophase. The action spectrum drawn up by Fondeville et al. (1967) has clearly
revealed that the reaction is mediated by a specific photosensitive pigment; given that
the maximum reaction is close to 450nm, these authors postulated that it was a flavin
or a carotenoid. The following experiments further investigate the movements of the
primary pulvini triggered by such sudden lighting and shows the bioelectrical events
accompanying them in order to reconsider the hypothesis on the mechanism of light
action in such stimulation.

Materials and Methods

Plants of Mimosa pudica L. were grown in a mixture of garden earth, heath-mould, sand and
peat in the percentages of 50, 20, 15 and 15 respectively. They were kept in climate controlled

Z. Pjlanzenphysiol. Bd. 106. S. 299-303. 1982.

300 G . ROBUN

chambers under the following conditions : temperature 25.5 ± 0.5 oC, relative humidity
55±5%.
Lighting was strictly regulated according to a sequence of 14h of light followed by 10h of
darkness (scotophase); in order to facilitate observations made near the middle of the sco-
tophase (e.g. between 2 and 4 p.m), the lighting was scheduled from 1900 hours to 900 hours
and was provided by white fluorescent «Phytor» tubes (ACEC, Charleroi, Belgium) giving an
intensity of 6,6 W m -2 in the 400-800 nm band (2,2 W m -2 in the 400-500 nm) at the level of
the leaf studied (generally the 7th-9th from the base, the plants bearing 10 to 12 leaves at the
time of sampling). The stimulation was given by the same light source as that used for the cul-
tures.
The device used to study the pulvinar movement was made up of two transparent pro-
tractors with diameters of 13 cm and 16 cm respectively, positioned in two parallel planes. The
plant was positioned between them in such a way that the primary pulvinus to be observed was
in line with the centre point of the protractors. In order to obtain a more accurate measurement
(to a y. degree), a fine glass needle, 10 cm long, and weighing approximately 10 mgs, was
attached to the peticle; the alignment of the needle with the protractor graduations gave the
value of the ex angle (Fig. 1 A, B).

Fig. 1: A: General morphology of a leaf of Mi·

mosa pudica (P,: primary pulvinus, P 2 : se-
condary pulvinus, P 3 : tertiary pulvinus). - B:
The device used to observe the pulvinus move-
ment. - C: Position of electrodes in the experi-
ment shown in Fig. 3.

The details of the recording method for the bioelectrical events by extracellular electrodes
can be seen elsewhere (Stoeckel and Pfirsch, 1975). That is, 50 I'm diameter 10 % iridium-pla-
tinum wires were inserted in the lower part of the primary pulvini (part L2 in the terminology
of Sibaoka, 1951) or into the petiole (P), the reference (R) being inserted in the earth of the pot
(Fig. 1 C). Electrodes were linked through impedance adaptors to potentiometric recorders
(Kipp and Zonen BD 9). The main advantage of this device is that it permits experiments of
long duration with recordings always taken at the same point. However it does not permit
determination of actual values of intracellular potentials, but only of their variations.

Results
1. Pulvinar movements induced by photostimulation
The curves given in Fig. 2 A show a similarity in the time course of the rapid down-
ward movement whether it was induced by shock or by photostimulation. In Bose's
first observation (1931), the rapid downward curvature was preceded by an upward

Z. Pjlanzenphysiol. Ed. 106. S. 299-303. 1982.

 Photostimulation in Mimosa pudica 301

o
ex _III'. ·.
de rees
1iIII·.·•."1iIII.__
30 60 TIME (min.)
~

ph
30 "'<it

50

70
Fig. 2: Comparison between the time course of
the foliar movement after stimulation of the 90
primary pulvinus.
A: When the large movement is induced by
shock or by photostimulation. - B: In the case
of increasing periods of photostimulation (Ph)
which do not trigger the large movement. -
The duration of photostimulation is indicated
by the white bars.

deviation of about ten degrees; this «positive deviation» was never recorded in our
observations.
We have observed that, when the full movement was not achieved, the pulvinus
still effected a small downward movement lasting between 12-15 min under our
experimental conditions, the complete return to the initial position being achieved in
about 30 min (Fig. 2 B). The curves show that 2 min exposure to light were sufficient
to induce such a movement and that 10 min exposure to light amplified this devia-
tion. It also appears that even under prolonged lighting the leaf had a tendency to
return to its initial position, but at a reduced rate.
In a comparative purpose, the curves drawn in Fig. 2 concern the movements
induced in the same leaf by light, over 4 successive days at the same moment of the
photoperiodic cycle (e.g. 3 p.m). Complementary experiments carried out at the
same time on different individual leaves gave the same general results: some pulvini
responded by a full rapid curvature and others only exhibited the small and slow
downward movement.

2. Bioelectrical events concomitant with the movement

The bioelectrical variations recorded in the lower part of the pulvinus were differ-
ent depending on whether the full rapid movement or the slow type of response is
induced. When the full rapid movement was induced the curves obtained were those
recorded in a pulvinus stimulated by shock (Fig. 3 A): an action potential was follow-
ed by a negative slow variation (phase 3) as previously noted (Roblin, 1975; Stoeckel,
1976). The action potential spreads into the petiole from the pulvinus, as measured
2.5 cm distal of the pulvinus (Fig. 3 C).
In leaves where only the small curvature was induced, a slow depolarization
occurred: exposure to light for 30 s is sufficient to induce such a process. The curves
of bioelectrical variations given in Fig. 3 B, recorded on the same primary pulvinus

Z. Pjlanzenphysiol. Bd. 106. S. 299-303. 1982.

302 G. ROBLIN

o 20 min. o 20min Fig.3: Bioelectrical vanatlons recorded

after photostimulation.
f:.VjL2 ~~ L2 00 A: In the lower part of the main pulvinus

.:: ~
when the large movement is induced. - B:
00 .100J 'ph I In the lower part of the main pulvinus un-
der increasing periods of photostimula-
o ~
, T 01f"--T tion which do not induce the large move-
ment. - C: Action potential spreading

.'j~ ".
;

o 20 min. 0 20min from the pulvinus to the petiole when the

large movement is triggered by a photosti-

v •••..........• .JID
J ... TIME
mulation (solid line) or by a wounding
(dashed line). - The photostimulation
(Ph) is indicated with the white bar show-
ing its duration.
o 2 4 6 min.

which movements are described in Fig. 2, show that the amplitude of depolarization
and the return to the initial potential increase with the irradiation time, in a close cor-
relation with the movement. Moreover, there was no spreading of bioelectrical varia-
tion in the petiole.

Discussion
Photostimulation triggered two modes of reaction in the main pulvinus of Mimosa
pudica.
The first mode, which has previously been described by few observers, is revealed
in a rapid and large curvature; the bioelectrical events are the same as those observed
after shock: an action potential (referred to as the «m-wave») is generated and spreads
into the petiole. Fondeville (1964) has put forward an hypothesis that a «stimulating
substance» was produced under such exposure to light. In fact the bioelectrical
recordings show that this hypothesis is improbable or, at least, that the «stimulating
substance» is different from that released after a wounding which induced character-
istic bioelectrical events, the so called «s-wave» (Houwink, 1935; Sibaoka, 1953;
Umrath, 1959; Roblin, 1979) (see Fig. 3 C).
The second mode of reaction is a slow movement of weak amplitude but with the
same time course as the large curvature (e.g. 20-30 min). Concomitantly, bioelectrical
events remaining localized in the pulvinus are noted. The time course of these bio-
electrical variations suggests that they correspond to the same processes involved in
the «phase 3» which is characteristic of the large movement.
The existence of a lau;nt period before the movement suggests that a sequence of
reaction takes place between the light absorption by a pigment and the membrane

Z. Pjlanzenphysiol. Bd. 106. S. 299-303. 1982.

 Photostimulation in Mimosa pudica 303

processes involved in such a pulvinus movement shown by the escape of water and
ions from the motor parenchyma cells of the organ (Toriyama, 1955; Allen, 1969;
Satter and Galston, 1971).
The question now to be considered is to determine at what step(s) of the sequence,
the reaction is directed either to the rapid and large curvature which is governed by
an all-or-none rule, or to the small and gradual movement, the distinction lying only
in the differing kinetics of the processes.
Acknowledgement
This work was supported by the Centre National de la Recherche Scientifique (ERA nO 701
and RCP nO 474).

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