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How The Venus Flytrap Snaps: A Poroelastic Buckling: Y. Forterre
How The Venus Flytrap Snaps: A Poroelastic Buckling: Y. Forterre
Y. Forterre
IUSTI CNRS UMR 6595, Universit de Provence, 5 rue Enrico Fermi, 13453 Marseille Cedex 13, France
J. Dumais
DOEB, Harvard University Biological Laboratories, 16 Divinity Avenue, Cambridge MA 02138, USA
ABSTRACT: We present a mechanical study of the snapping closure of the carnivorous plant Venus flytrap
(Dionaea muscipula), which exhibits one of the fastest motion in the vegetable kingdom (typically 100 ms). Using
high-speed video measurements and non-invasive microscopy techniques, we propose that the fast closure of the
Venus flytrap results from a snap-buckling instability, which is damped by the flow of water through the tissues.
A simple model for the snapping of a poroelastic shell and experiments on the mechanical properties of the leaf
tissue are completely consistent with this picture.
1 INTRODUCTION
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Figure 3. a) Measured strain field of the outer surface due
Figure 2. a) Spatially averaged mean curvature m as a func- to closure (averaged over 6 different leaves), where pos-
tion of time. Inset: 3D stereoscopic reconstruction of the leaf itive strains correspond to extension and negative strains
in the open and closed states. b) Spatially averaged gaussian correspond to contraction of the surface in the principal
curvature g as a function of time. The plant is triggered at directions as shown. b) Typical local strain measured on the
t = 0. The closure dynamics is characterized by three phases outer surface of the leaf inside the dashed region, super-
(IIII) (see text). The solid line corresponds to the poroelastic posed on a micrograph of the leaf surface. The polar plot
model (see 5.3). shows the local strain in the direction , (l/l)(), defined
by (l/l)() = 1 cos( 0 ) + 2 sin( 0 ) where 1 and
2 are the principal values of the strain tensor and 0 is the
the open state and curved inward (concave) in the angle of the principal axes with respect to the reference frame
closed state (Fig. 2a). The leaf shape can be naturally of coordinates. Scale bar: 0.1 mm.
characterized in terms of its spatially averaged mean
curvature, m , and its spatially averaged gaussian cur-
vature, g . The snapping motion itself is characterized
by three phases (Fig. 2a): a slow initial phase (20% of 2.2 Strain measurements
total displacement in 1/3 s); a rapid intermediate phase To understand the origin of these curvature changes,
(60% of total displacement in 1/10 s); and finally a sec- we measure the local strains by recording the position
ond slow phase (20% of total displacement in 1/3 s). of fiducial markers on the entire outer surface of the
Most of the leaf displacement occurs in the interme- leaf before and after closure (Green et al. 1991). Our
diate phase, during which the leaf geometry changes measurements of the strain (Fig. 3) are consistent with
from convex to concave (Fig. 2b) shows g as a func- earlier point-wise measurements (Hodick & Sievers
tion of time. We see that g is not constant and also that 1989) but go beyond these by characterizing the spatial
g changes slowly and then rapidly as it passes through structure of the strain field over the entire leaf. Figure
a minimum. As changes in g correspond to stretch- 3a shows that the maximum strain perpendicular to the
ing the mid-plane of the leaf (Love 1944), the first midrib (x-direction) is six times the maximum strain
observation implies that closure involves the storage parallel to the midrib (y-direction). Furthermore, we
and release of elastic energy; while the second obser- find that the strains on the inner surface of the leaf are
vation implies that closure is characterized by a slow 1%, implying that closure is triggered primarily by
storage of elastic energy followed by a rapid release. differential strains in the x-direction.
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Figure 4. Cutting the closed leaf along the dotted lines
shows the natural curvatures in the x-direction (right) and
the y-direction (left). Scale bar: 1 cm.
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Copyright 2005 Taylor & Francis Group plc, London, UK
than small weakly curved leaves. In order to test this,
we measured the speed of the closure as a function of
the geometrical parameter for many different leaves
and plants. (Fig. 6a) shows a clear positive correla-
tion between the maximum speed of the snap and .
Interestingly, the time delay between the beginning of
the motion and the snap increases with the parameter
as shown in (Fig. 6b), just as the maximum speed
of snapping does. This is probably because the delay
reflects the time required to cross the stretching barrier,
which increased with . For a typical leaf L 1 cm,
0.1 cm1 , h 0.1 cm so 1 c , i.e. the leaves
are poised at the transition between smooth closing and
snapping. Since the plant is clearly best served by (i)
minimizing the delay between triggering and closure,
(ii) snapping rapidly rather than closing smoothly, this
choice of may not be a coincidence.
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where P
2 h2 /2 kE is the poroelastic time. This
expression arises from the solution of a Darcy-type
diffusion equation for the fluid pressure, which in turn
gives rise to a moment resisting the bending:
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speed from the cellular to the organ level by natures Hodick, D. & Sievers, A. 1989. On the mechanism of
consummate hydraulic engineers, plants, shows how trap closure of Venus flytrap (Dionaea muscipula Ellis).
controlling elastic instabilities in geometrically slen- Planta 179:3242.
der objects provides an alternative to the more common Juniper, B. E., Robins, R. J. & Joel. D. M. 1989. The
carnivorous plants. London, San Diego:Academic Press.
muscle-powered movements in animals. Love, A. E. H. (4th ed.) 1944. A treatise on the mathematical
theory of elasticity. New York:Dover.
ansfield, E. H. (2nd ed.) 1989. The bending and stretching of
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