Mimosa pudica: Electrical and mechanical stimulation of

plant movementspce_2066 163..173

ALEXANDER G. VOLKOV
1
, JUSTIN C. FOSTER
1
, TALITHA A. ASHBY
1
, RONALD K. WALKER
1
, JON A. JOHNSON
1
&
VLADISLAV S. MARKIN
2
1
Department of Chemistry and Biochemistry, Oakwood University, 7000 Adventist Blvd., Huntsville, AL 35896, USA and
2
Department of Neurology, University of Texas, Southwestern Medical Center, Dallas, TX 75390-8813, USA
ABSTRACT
Thigmonastic movements in the sensitive plant Mimosa
pudica L., associated with fast responses to environmental
stimuli, appear to be regulated through electrical and
chemical signal transductions. The thigmonastic responses
of M. pudica can be considered in three stages: stimulus
perception, electrical signal transmission and induction of
mechanical, hydrodynamical and biochemical responses.
We investigated the mechanical movements of the pinnae
and petioles in M. pudica induced by the electrical stimu-
lation of a pulvinus, petiole, secondary pulvinus or pinna by
a low electrical voltage and charge. The threshold value was
1.31.5 V of applied voltage and 2 to 10 mC of charge for the
closing of the pinnules. Both voltage and electrical charge
are responsible for the electro-stimulated closing of a leaf.
The mechanism behind closing the leaf in M. pudica is
discussed. The hydroelastic curvature mechanism closely
describes the kinetics of M. pudica leaf movements.
Key-words: charged capacitor method; electrical signaling;
motor cells; plant electrophysiology.
Abbreviations: C, capacitance; d, distance between the
edges of the a pair of pulvinules; DPDT, double pole double
throw switch; HEC, a hydroelastic curvature model; I, elec-
trical current; L, the hydraulic coefcient of pore perme-
ability; P, hydrostatic pressure; PXI, PCI eXtensions for
Instrumentation; Q, charge of capacitor; t, time; U, voltage;
t
a
, characteristic time of the pore opening; t
r
, characteristic
time of uid transfer.
INTRODUCTION
Mimosa pudica is a seismonastic plant in which the leaves
close and the petiole hangs down in response to wind, vibra-
tion and touch as a defense mechanism for protection from
animals and some insects (Bose 1902, 1907, 1913, 1918, 1926,
1928). As it was mentioned by Bose (1918) many years ago
in his famous book about the sensitive plant M. pudica, the
phenomenon of movements in plants under the action
of external stimuli presents innumerable difculties and
complications. This plant also responds to electrical
stimulation.
The electrical phenomena in plants attracted researchers
since the 18th century (Bertholon 1783; Bose 1926; Bal-
ushka, Mancuso & Volkman 2006; Volkov 2006). Both
mechanical and electrical stimulation of M. pudica can lead
to the closing of a pinna and the falling down of the petiole.
This is caused by different processes and components in
the primary and secondary pulvini, which involve voltage-
gated potassium and calcium-permeable anion channels
(Abe 1981; Roblin & Fleurat-Lessard 1987; Stoeckel &
Takeda 1993), H
+
-ATPases (Liubimova, Deminovskaya
& Fedorovich 1964; Liubimova-Engelgardt et al. 1978;
Fleurat-Lessard et al. 1997), aquaporins (Temmei et al.
2005), actin (Kameyama et al. 2000; Yao, Xu & Yuan
2008), Riccas factor (Ricca 1916) or gallic acid 4-O-
(b-d-glucopyranosyl-6-sulfate (Schildknecht & Meier-
Ausgenstein 1990), neurotransmitters (Roshchina 2001) and
osmotic pressure (Tamiya et al. 1988).The actincytoskeleton
in the pulvinus plays an important role in the petiole
bending. Kameyama et al. (2000) found that the actin in M.
pudica is heavily tyrosine phosphorylated and that changes
in the extent of phosphorylation correlate with the degree of
bending of the plants petioles. Liubimova et al. (1964) mea-
suredATPase activity during the thigmonastic movement of
M. pudica. Fleurat-Lessard et al. (1997) studied the distribu-
tion and activity of the plasma membrane H
+
-ATPase in M.
pudica in relation to the leaf movements and found a high
density of H
+
-ATPase inthe phloemandpulvini. Liubimova-
Engelgardt et al. (1978) isolated from the pulvinus (1)
soluble ATPase loosely bound to pulvinus structures;
(2) Ca,Mg-dependent ATPase tightly bound to pulvinus
structures; (3) ATPase bound to the pulvinus membrane
structures. Ca,Mg-dependent ATPase is similar to ATPases
from muscle and non-muscle motile cells (Liubimova-
Engelgardt et al. 1978). Ion channels in the pulvinus are
involved in the redistribution of potassium, chloride and
calcium during the gravitotropically induced movement of
M. pudica (Roblin & Fleurat-Lessard, 1987). Transport of
ions between upper and lower parts of the pulvini leads to
the osmotic owof water through aquaporins and a pressure
gradient is created in the pulvinus, which can result in the
falling down of the petiole.
Correspondence: A. G. Volkov. Fax: +1 256 726 7113; e-mail:
agvolkov@yahoo.com
Plant, Cell and Environment (2010) 33, 163173 doi: 10.1111/j.1365-3040.2009.02066.x
2009 Blackwell Publishing Ltd 163
Mechanical movements inM. pudica were inducedby very
high applied voltages (Ritter 1811; Gardiner 1888; Bose
1918; Jonas 1970; Balmer & Franks 1975). Balmer & Franks
(1975) briey applied 200400 V between the soil and the
primary pulvinus to measure the contractile characteristics
of a petiole. They estimated that the threshold voltage was
about 25 V with any electrode polarity. Jonas (1970) used a
0.5 mF capacitor charged by 50, 100 and 150 V for electro-
stimulation and found that there were oscillations of leaves
and fast petiolar movement after the application of an elec-
trical shock.WhenYao et al. (2008) applied 9 Vto M. pudica,
the petioles bent downwards and the pinnae closed.
In the study reported, we analysed both experimentally
and theoretically the mechanism of mechanical movements
in M. pudica induced by low voltage electro-stimulation
of the pulvinus, petiole or pinna. The charged capacitor
method (Volkov et al. 2008a) is a very efcient tool for the
study of bioelectrochemistry of cells, clusters of cells or
for electro-stimulation of whole plants and evaluation of
biologically closed electrical circuits (Volkov, Coopwood
& Markin 2008b; Volkov et al. 2009a; Volkov, Carrell &
Markin 2009b). The present work aims (1) to use electrically
or mechanically stimulated M. pudica leaf movements as
experimental models; (2) to display a digital description of
the kinetics of these movements; and (3) to provide evi-
dence that these movements are governed by hydroelastic
curvature (HEC) mechanism (Markin, Volkov & Jovanov
2008) that has been previously proposed for the Venus
ytrap closing process.
MATERIALS AND METHODS
Electrodes
All measurements were conducted in the laboratory at
21 C inside a Faraday cage mounted on a vibration-
stabilized table (Fig. 1). Ag/AgCl electrodes were prepared
fromTeon-coated silver wires (A-M Systems, Inc., Sequim,
WA, USA) according to the method described by Ksenzhek
& Volkov (1998). The resistance between two Ag/AgCl
electrodes that are 2 cm apart in 0.1 m KCl solution was
found to be 10 kOhm. The response time of Ag/AgCl elec-
trodes was less than 0.1 ms. As a control experiment, we also
used platinum electrodes instead of Ag/AgCl electrodes
and received the same results. Platinum electrodes were
prepared fromTeon-coated platinum wires (A-M Systems,
Inc.). Following insertion of the electrodes into M. pudica,
the pinnae closed. We allowed plants to rest until the leaves
were completely open. The dependence of the electrical
discharge on time in M. pudica was measured 3 to 25 h after
insertion of the electrodes.
Plant electro-stimulation
PXI (PCI eXtensions for Instrumentation) is a rugged
PC-based platform that offers a high-performance solution
for measurement and automation systems. PXI combines
the Peripheral Component Interconnect electrical bus with
the rugged, modular Eurocard mechanical packaging of
CompactPCI and adds specialized synchronization buses
and key software features. PXI also adds mechanical, elec-
trical and software features that dene complete systems
for test, measurement and data acquisition. A NI-PXI-
4071 digital multimeter connected to 0.2 mm thick non-
polarizable reversible Ag/AgCl electrodes was used to
record the digital data. A NI PXI-4110 DC Power Supply or
C-type 1.5 V electrical battery was the voltage sources for
capacitor charging. The charged capacitor was used for
electro-stimulation of M. pudica.
The Charged Capacitor Method (Volkov et al. 2008a,b,
2009a,b) was used to estimate, with high precision, the
amount of electrical energy necessary to induce a response.
A double pole, double throw (DPDT) switch was used to
connect the known capacitor to the 1.5 V battery during
charging, and then to the plant during plant stimulation.
Since the charge of a capacitor Q connected to the voltage
source U is Q = CU, we can precisely regulate the amount
of charge using different capacitors and applying various
voltages. By changing the switch position, we can instanta-
neously connect the charged capacitor to the plant and
induce a response. We used various capacitors that had a
capacitance ranging from 1 to 100 mF.
Images
Digital video recorders, Sony DCR-HC36 (Sony USA, San
Diego, CA, USA) and Canon ZR300 (Canon USA, Lake
Success, NY, USA), were used to monitor the movements of
the M. pudica and to collect digital images, which were
analysed frame by frame. A photo camera Nikon DX40DX
(Nikon USA Inc., Melville, NY, USA) with AF-S Micro
Nikkor 105 mm 1:2.8 G ED VR lenses was used for the
photography of M. pudica.
Plants
The seeds of M. pudica L. were soaked in warm water
(30 C) for 48 h. Then they were grown in well-drained peat
moss at 21 C with a 12:12 h light : dark photoperiod. After
growing for 2 weeks, the seedlings were transplanted into
Light
Faraday cage
NI PXI-1042Q
Microcomputer
NI PXI-4071
Digital multimeter
LabView software
Figure 1. Experimental setup.
164 A. G. Volkov et al.
2009 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 163173
pots in a plant-growing chamber. The humidity averaged
4550%. The plants were watered every day. Twothree-
month-old plants were used for the experiments. Irradiance
was 700800 mmol photons m
-2
s
-1
. All experiments were
performed on healthy adult specimens.
RESULTS
Mechanical stimulation of M. pudica
Figure 2 shows different shapes and curvatures of pinnules
in open or closed states. During the day, pinnules can have
either concave or convex congurations (Fig. 2). Figure 3
schematically illustrates the different curvatures of pinnule
pairs in the process of opening and closing. Kinetics of
pinnules closing, induced by a gentle mechanical touch of
the midrib of a pinna, is shown in Fig. 4a. The shape of this
time dependence for the distance between the edges of
one pair of pinnules is very similar to the shape of similar
dependencies for the Venus ytrap (Markin et al. 2008;
Volkov et al. 2008a,b, 2009a,b). However, the speed of
closing is about 10 times slower. Figure 5 shows the time
dependence for pinnules opening after being mechanically
stimulated to close. The process of pinnules closing takes
place in 45 s, whereas the opening of the pinnules occurs in
about 600 s.
Electrical stimulation of M. pudica
We found that the closing of the pinnules and the petiole
bending can be electro-stimulated by the application of a
low voltage to the pulvinus (Figs 6 & 7). Instead of applying
a non-physiological high voltage of 200400 V (Ritter 1811;
Jonas 1970; Balmer & Franks 1975), we applied 1.5 V to
the pulvinus. The minimum threshold voltage was 1.3 V.
Kinetics of the petiole bending obtained on three different
M. pudica plants after electro-stimulation is shown in
Fig. 8. There is a 0.50.7 s delay in response to the electrical
stimulus, and fast petiole bending occurs in 4 s. The time
dependence for petiole relaxation after pulvinus electro-
stimulation by 1.5 V (+ on the top and - on the bottom of
the pulvinus) compared with the initial state is shown in
Fig. 9. Figure 10 shows the kinetics of a petiole relaxing
to the initial state 10 min after electro-stimulation and
subsequent bending of the petiole.
Figure 11 shows that the closing of the pinnules can be
stimulated by a charged capacitor connected to a secondary
pulvinus and a rachis. If a capacitor of capacitance C is
discharged through a parallel resistor R, the dependence of
the capacitors voltage U
c
on time is
U t U
t
c
( )
( )
0
exp

(1)
where
RC (2)
Figure 2. Photos of Mimosa pudica:
(a) open pinnules; (b) pinnules in the
process of closing, (c) and (d) usual state
of leaves with open and partially
closed pinnules.
(a) (b)
(d)
(c)
Pinnules
Pulvinule
Rachis
Petiole
Secondary
pulvinus
Pinna
A B C D
Pulvinule
Pinnule
5 10 s
10 min
Figure 3. Different curvatures of pinnules in (a) open state,
(b) in the process of closing, (c) in a closed state, (d) in the
process of opening.
Electrical and mechanical stimulation of plant movements 165
2009 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 163173
and U
0
is initial voltage. The circuit time constant t governs
the discharging process. With increasing capacitance, the
time of the capacitors discharge increases according to
Eqns 1 and 2. Since charge Q = CU, a closing charge can
be estimated from the time dependence of a capacitor
discharge.
The closing of the pinnules after electro-stimulation by a
charged capacitor is shown in Fig. 12. Similar effects of pin-
nules closing can also be induced by a charged capacitor if
the polarity of the electrodes is changed (Fig. 13). However,
a higher charge is required for pinnules closing.
The closing of the pinnules can be induced by applying a
DC current to the petiole from a DC power supply
(Fig. 14). Since charge Q = I t, it is possible to nd the
closing charge. The closing charge depends on the location
and polarity of the electrodes and varies from 2 to 10 mC.
DISCUSSION
Theoretical: The HEC model
The sensitive plant M. pudica belongs to the exclusive
type of plants that display active movements of their parts.
Especially interesting was the insect-catching mechanism
of the Venus ytrap. Throughout the years, a number of
explanations for this phenomenon were put forth (Darwin
1875; Lloyd 1942; Williams & Bennet 1982; Hodick &
Sievers, 1989; Forterre et al. 2005). Recently, the novel
mechanism based on a HEC model was proposed by
Markin et al. (2008). Pinnules in M. pudica and lobes in
the Venus ytrap move as a result of changes in the shape,
curvature and volume of the cells. During the closing or
opening of the pinnules in M. pudica or lobes in the Venus
ytrap, curvature varies between concave and convex
Figure 4. Kinetics of pinnules closing,
mechanically stimulated (MS) by a touch
of the midrib. Points are experimental
data, solid line is the theoretical
dependence estimated from Eqn 3.
d, distance between rims of pinnules in
the middle of pinna; d
max
, maximal
distance between rims of pinnules in the
middle of pinna.
Time (s)
0 2 4 6 8 10
d
/
d
m
a
x
0.0
0.2
0.4
0.6
0.8
1.0
0 0.2 s 0.4 s
1.5 s 2.0 s 5.0 s
0.6 s 0.8 s 1.0 s
MS
d d
max
(b)
(a)
166 A. G. Volkov et al.
2009 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 163173
shapes. Movements in both plants might have similar
mechanism.
The closing of the Venus ytrap represents the non-
muscular movement based on hydraulics and mechanical
elasticity. The nastic movements in various plants involve a
large internal pressure (turgor) actively regulated by the
plants. The leaf of the Venus ytrap is visualized by the
model as a thin, weakly curved elastic shell with intrinsic
natural curvatures that depend on the hydrostatic state of
the two surface layers of cells A and B (Markin et al. 2008),
where different hydrostatic pressures P
A
and P
B
are main-
tained. Two layers of cells, mechanically connected to each
other, behave like a bilayer couple where the in-plane
expansion or contraction of any of them causes a change in
the curvature of the whole leaf. Besides that, the leaf has an
intrinsic mechanical tendency to curve so that the interplay
between mechanical and hydraulic forces provides the
possibility to change the shape and to perform active
movement when hydrostatic pressure changes quickly.
In the open state, the pressure in the upper layer is higher
than the pressure in the lower layer, maintaining the convex
shape of the leaf. The fact that the hydrostatic pressure in
different parts of the plant can vary is very well known. It is
also known (Tamiya et al. 1988; Schildknecht & Meier-
Ausgenstein 1990; Temmei et al. 2005) that stimulation of a
Mimosa plant causes a very fast redistribution of water.
Tamiya et al. (1988) found that after stimulation, water in
the lower half of the main pulvinus is transferred to the
upper half of the main pulvinus. Water movement in con-
junction with Mimosa movement was visualized by a non-
invasive NMR imaging procedure (Tamiya et al. 1988). This
fast water redistribution is obviously driven by the pressure
difference between various parts of the plant, and it occurs
through open pores.
The HEC model assumed that water pores between the
two hydraulic layers are closed at the resting state. Aqua-
porins are likely to play the role of these pores (Volkov
et al. 2008b). The external trigger, whether mechanical or
Figure 5. Kinetics of pinnules opening
after closing by a touch of the midrib.
d, distance between rims of pinnules in
the middle of pinna; d
max
, maximal
distance between rims of pinnules in the
middle of pinna.
1.0 min 1.5 min 2.5 min
3.5 min 4.5 min 5.5 min
6.5 min 7.5 min 12 min
Time (min)
0 2 4 6 8 10 12 14
d
/
d
m
a
x
0.0
0.2
0.4
0.6
0.8
1.0
Electrical and mechanical stimulation of plant movements 167
2009 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 163173
electrical, results in the opening of these connecting pores.
Water then rushes from the upper to the lower layer and the
bilayer couple quickly changes its curvature from convex to
concave and the trap closes.
If the trigger reaches the threshold value at the moment
t
s
and the characteristic time of the pore opening kinetics
is t
a
, then the open probability of the pores is given by
a simple exponential function n
op
(t) = 1 - Exp[-(t - t
s
)/t
a
].
The rate of uid transfer between the two layers was
presented as J = n
op
L(P
A
- P
B
), where L is the hydraulic
coefcient of pore permeability. The characteristic time of
uid transfer was found as t
r
= 1/(2k
r
L).
The trigger signal opens the pores between these layers
and the uid rushes from one layer to another. The leaf
relaxes to its equilibrium state corresponding to the closed
conguration. The closing process was parameterized by
the distance between the edges of the trap that varied with
time as
d t x
t t
x ( ) ( )

_
,

1
]
1

( )

+ 1 1 exp exp

a s
a
s
r r
(3)
Here, x gives the nal position of the lobes. From a com-
parison with the experiment, Markin et al. (2008) were able
to nd typical parameters of the Venus ytrap closing at
t
a
= 20 ms and t
r
= 70 ms. These times can vary depending
on the type of trigger, but they remain very short; this is the
cause for the rapid closing of the Venus ytrap (Volkov
et al. 2008a,b, 2009a,b).
Closing consists of three distinctive phases. Immediately
following stimulation, there is a mechanically silent period
with no observable movement of the plant (t
s
). This is fol-
lowed by a period when the movement of the lobes starts
with acceleration (t
a
). The third period of fast movement is
the actual trapping when the leaves quickly relax to the new
equilibrium state (t
r
). Similar phenomenon was found
during the pinnules closing in M. pudica (Fig. 4).
In conclusion, the goal of the HEC model was to describe
the three stages of the trap closing discovered in the experi-
ments and to elucidate the processes that govern these
stages. The model was deliberately made as simple as pos-
sible so that the mechanical details would not mask the
basic ideas.
The detailed mechanical analysis based on bending
energy consideration resulted in Eqn 3 describing closing
kinetics and consequent discovery of the parameters of the
system (Markin et al. 2008). Of course, M. pudica has a
different anatomy and reacts to external stimuli much more
slowly. Nevertheless, there is reason to believe that hydro-
static effects are essentially involved in the active move-
ments of M. pudica. Due to this reasoning, we decided to
parameterize our quantitative observations of this plants
movement with the HEC model and specically with Eqn 3,
which can be used for describing the displacement of
0 90 ms 210 ms 300 ms 390 ms
480 ms 570 ms 660 ms 1 s
100 s 120 s 180 s
+
-
Pulvinus
Figure 6. Closing of pinnules and a petiole bending triggered by electrical discharge in Mimosa pudica between electrodes in a pulvinus
connected to 1.5 V DC Power Supply during 10 s.
168 A. G. Volkov et al.
2009 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 163173
certain parts of the plant. Looking at Figs 4b, 9 and 10, one
can notice that the movement of M. pudica after triggering
also consists of three stages: a mechanically silent period
with no observable movement of the plant (t
s
), the move-
ment with acceleration (t
a
) and the third period of relax-
ation to the new equilibrium state (t
r
).
This parameterization will permit the nding of the time
parameters of the system and will help to elucidate the
actual structure and mechanisms participating in the M.
pudica movements. That was the reason to formally extend
the hydroelastic model to the case of M. pudica.
Leaf movements in M. pudica are caused by asymmetri-
cal turgor changes between two layers in the cortical cells of
the pulvinus, secondary pulvinus and pulvinule at the base
of each pinnule. Figure 4 shows that the kinetics of closing
the pinnae in M. pudica is similar to the time dependency of
the trap closing in Dionaea muscipula Ellis (Volkov et al.
2008a,b), but 10 times slower. The mechanism of pinnules
closing is probably very similar to the trap closing in the
Venus ytrap proposed by Markin et al. (2008). Mechanical
stimuli generate an action potential (Sibaoka 1962, 1991;
Stoeckel & Takeda 1993) and activate ion transport
0 s 4 s 10 s 12 s
13 s 15 s 17 s 37 s
360 s 1380 s
+
-
Stem
Petiole
Pulvinus
Figure 7. Closing of pinnules and a petiole bending triggered by 5 s electrical discharge in Mimosa pudica between electrodes connected
to 1.5 V DC Power Supply during 10 s and relaxation to the initial state during 20 min.
Time / s
0 1 2 3 4 5 6
/
m
a
x
0.0
0.2
0.4
0.6
0.8
1.0
max
Q
Q
Q
+ +

Stem
Petiole
Figure 8. Kinetics of a petiole bending triggered by 5 s
electrical discharge in Mimosa pudica between electrodes
connected to 1.5 V DC Power Supply.
Electrical and mechanical stimulation of plant movements 169
2009 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 163173
between the lower and upper parts of a pulvinus or/and
in pulvinules (Abe 1981; Samejima & Sibaoka 1982). This
initiates the opening of aquaporins and induces water
transport due to osmosis and a change in turgor pressure
(Tamiya et al. 1988; Temmei et al. 2005). The leaves of M.
pudica actively employ turgor pressure and hydrodynamic
ow for their mechanical movements.
We are indebted to the reviewer of this paper who
suggested discussing the terminology describing plant
movements. The thigmonastic reactions of sensitive plants
like M. pudica or the Venus ytrap to external stimuli are
usually called active movements. However, these reactions
consist of a few steps involving different mechanisms that
might look quite passive rather than active. For example,
0 2 min 3 min 4 min
5 min 6 min 7 min 8 min
9 min 10 min 11 min 14 min
Figure 9. Time dependence of the petiole relaxation after electrical stimulation of the pulvinus.
Time / min
0 2 4 6 8 10 12
/
m
a
x
0.0
0.2
0.4
0.6
0.8
1.0
Q
Q
Figure 10. Kinetics of a petiole relaxation after electro
stimulated petiole bending in Mimosa pudica. Points are
experimental data, solid line is the theoretical dependence
estimated from Eqn 3.
Time / s
0 2 4 6 8 10 12
U

/

V
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
1.6
off Ag/AgCl (+)
Ag/AgCl ()
Figure 11. Time dependence of voltage between two platinum
electrodes located in the secondary pulvinus (+) and a rachis (-)
connected to charged 4.7 mF capacitor and NI-PXI-4071 digital
multimeter. U is the capacitor voltage in volts. A capacitor was
disconnected (off) when the pinnae start to close.
170 A. G. Volkov et al.
2009 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 163173
the most spectacular phase of the Venus ytraps rapid
closing and catching of insects according to the HEC model
is driven by the release of elastic energy stored in the leaves
of the Venus ytrap. This energy is locked in the leaves
forming the trap due to the pressure differential between
two hydraulic layers. After the opening of the water chan-
nels and the redistribution of the water, the latch is
removed and the leaf relaxes to the new equilibrium closed
state. This certainly looks like a passive process, although it
represents an active reaction of the plant to external
stimuli. After digesting the prey, the leaf slowly opens due
to the pumping of water from one hydraulic layer to
Time / s
0 20 40 60 80 100 120 140 160 180
I

/

n
A
160
140
120
100
80
60
40
20
0
Closing
pinnules
Ag/AgCl (+) Ag/AgCl ()
Petiole
Figure 14. Time dependence of DC current between two
Ag/AgCl electrodes located in the petiole connected to 1.3 V DC
Power Supply and NI-PXI-4071 digital multimeter. Distance
between electrodes was 1 cm.
0 1 s 7s 9 s
25 s 35 s 40 s 55 s
Figure 12. Sequence of Mimosa pudica photos after electrical stimulation (2 mC, 1.5 V) using two platinum electrodes located in the
secondary pulvinus (+) and a rachis (-) connected to charged 4.7 mF capacitor and NI-PXI-4071 digital multimeter. A capacitor was
disconnected when the pinnules start to close.
Time / s
0 20 40 60 80 100 120 140
U

/

V
1.6
1.4
1.2
1.0
0.8
0.6
0.4
0.2
0.0
off
Ag/AgCl ()
Ag/AgCl (+)
Figure 13. Time dependence of voltage between two platinum
electrodes located in the secondary pulvinus (-) and a rachis (+)
connected to charged 10 mF capacitor and NI-PXI-4071 digital
multimeter. U is the capacitor voltage in volts. A capacitor was
disconnected (off) when the pinnae start to close.
Electrical and mechanical stimulation of plant movements 171
2009 Blackwell Publishing Ltd, Plant, Cell and Environment, 33, 163173
another. The mechanism of this process is not yet known,
but it is certainly driven by another source of energy, either
osmotic or chemical. All of these stages constitute the entire
hunting cycle of the Venus ytrap described by the term
active movement. One should just remember that this
descriptive term encompasses multiple processes.
Comparison of theoretical and
experimental results
In Figure 4b, we followed the kinetics of pinnules closing as
a function of time. We tted the kinetics with function (3)
(continuous line in Figs 4b and 10). One can see that this
curve very accurately describes the experimental measure-
ments. From the curve tting, we were able to nd the
following parameters for Fig. 4b: t
s
= 0.27 s, t
a
= 0.2 s,
t
r
= 1.7 s and x = 0.243. In Fig. 10, we followed the kinetics
of a petiole relaxing and found t
s
= 1.1 s, t
a
= 0.4 s, t
r
= 3.5 s,
and x = 0.
One can see that the movement of M. pudica is an order
of magnitude slower than the Venus ytrap, although it
proceeds through the same three phases. We believe that
these phases reect consequently (1) the processing of the
triggering signal; (2) the opening of the water channels
between two hydraulic reservoirs; and (3) the transfer of
water between these reservoirs. The next step in the devel-
opment of this mechanism should be establishment of
specic players in these movements.
So, the HEC model for M. pudica is based on the assump-
tion that the pinnules possess curvature elasticity and
consist of outer and inner hydraulic layers where different
hydrostatic pressure can build up. The open state of the
pinnules contains a high elastic energy accumulation due to
the hydrostatic pressure difference between the outer and
inner layers of the pinnule. Stimuli open pores connecting
the two layers; water then rushes from one hydraulic layer
to another and the trap relaxes to the equilibrium congu-
ration corresponding to the closed state. Water transport
leads to extension of the cells on the exterior surface of the
leaf and to a change in the natural curvature of the leaf,
which is likely to drive the closure process.
There are many quick mechanical movements in plants,
and the new HEC theory can be used for understanding
and estimating their exact mechanisms. The non-invasive
method of electro-stimulation permits the study of certain
steps in signal transduction and fast responses in the plant
kingdom. However, the detailed elucidation of the mecha-
nisms of these steps will demand additional biophysical and
biochemical approaches.
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