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Introduction
Stickslip friction generates a vast array of oscillatory
movements, ranging from earthquakes and squeaking doors to
the tuned excitation of bowed stringed instruments (Benade,
1990; Persson, 2000; Rabinowicz, 1995). While the transitions
between static and sliding friction are key to many biological
movements (e.g. Goodwyn and Gorb, 2004; Niederegger and
Gorb, 2006; Scherge and Gorb, 2001), biological sound
production generated through periodic stickslip frictional
movements is both unusual and understudied. Here, we
characterize the acoustic mechanics of stickslip friction in the
California spiny lobster (Panulirus interruptus) (Fig.1) using a
combination of kinematics, acoustic analyses and mechanical
friction tests.
While virtually all translating surfaces experience friction
forces that oppose movement and many surfaces simply slide
relative to each other, stickslip systems are defined by the
presence of oscillations due to shifting magnitudes of friction
and elastic energy storage between the two surfaces (Fig.2).
Most physics examples highlight the principles of friction in
steady-sliding systems; a simple example is a block, which
Eye
File
Stick phase
Plectrum
Plectrum
file
File
File
(1)
Plectrum
Fstatic
Fstatic
Fmin
Fkinetic
Static
Sliding
Sliding
Time
Time
Ff
Fn
V
Force beams
Ff
Plectrum
File
Linear stage
(movement in/out
of page)
Fn
Fig.4. Scanning electron micrographs of the file and plectrum surfaces. Anterior is to the right of the page in all images. (A) The dorsal file surface
is covered with microscopic shingles with their leading edges oriented anteriorly. A groove on the medial edge of the file (medial to bottom of
page) guides a knob on the plectrum such that the plectrum slides anteriorly and posteriorly in a controlled alignment, similar to a sliding door in
its track. Scale bar, 100m. (B) An oblique view of the file shingles shows the distinct separation of the shingles and the additional ridge running
medio-laterally on most shingles. Scale bar, 5m. (C) A dorsal view of file shingles (scale bar, 10m; inset, 2m) shows the more prominent
frictional edges of the shingles on the anterior side, analogous to shingles on the roof of a house. (D) The ventral surface of the plectrum is
composed of soft-tissue plectrum ridges that run parallel to their antero-posterior movement over the file. Scale bar, 100m. (E) The leading
edges (posterior) of the plectrum ridges fuse with the setae on the ventral surface of the plectrum. Scale bar, 50m. (F) The anterior edge of the
hemisphere of plectrum ridges ultimately attaches to the ventral cuticle of the antennal base. Scale bar, 100m.
Relative acoustic
amplitude (V)
Plectrum
Plectrum
velocity (cm s1) displacement (cm)
0.16
0.08
0
20
10
0
0.3
0
0.3
0
0.01
0.02
0.03
Time (s)
0.04
RMS pulse
amplitude (V)
Scaled peak
pulse amplitude
R2
F-ratio
P-value
R2
F-ratio
P-value
R2
0.84
0.82
15.9
3.6
0.0001*
0.06
0.80
0.77
16.0
5.7
0.0001*
0.02
0.25
0.81
0.04
0.8
0.75
0.2
0.6
F-ratio P-value
Scaled RMS
pulse amplitude
R2
F-ratio
P-value
0.43
0.29
36.1
12.3
<0.0001
0.0007
0.16
0.7
0.4
16.0
0.0001
Asterisks indicate significant differences across individuals in two of the tests. Two results are not shown, because they violated statistical
assumptions of the repeated-measures tests (negative-tending variances).
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Normal force
Fn (N)
Frictional force
Ff (N)
Plectrum
displacement
(cm)
0.2
0
2
0
4
Coefficient of
friction
Discussion
Through the integration of acoustics, kinematics,
0
functional morphology and friction mechanics, we
2 D
present the first mechanical characterization of a
bioacoustic stickslip mechanism. We found that P.
1
interruptus has similar morphological and acoustic
features to previously studied spiny lobsters, with soft
0
tissue plectrum ridges that articulate against the
0.28
0.32
0.36
0.40
0.44
frictional edges of cuticular shingles to produce pulsed
Time (s)
stickslip sounds. The kinematic analyses suggested
that
plectrum
movements
influence
signal
Fig.6. Synchronous measurements of plectrum displacement (A) frictional
characteristics,
while
the
friction
results
provided
a
forces (B, Ff; C, Fn) and coefficient of friction (D, ). The oscillating force
mechanical
explanation
for
the
variation
in
plectrum
and coefficient of friction waveforms are a product of the stickslip friction
kinematics and acoustics. Plectrum velocity and the
dynamics of the plectrum moving over the file. Each force trace (Ff and Fn) is
normal force between the plectrum and file played
correlated with the movement of the plectrum. In B, the raw data are depicted
important roles in the acoustic and frictional properties
in blue and the filtered data (low pass filter at 280Hz) in red. Grey regions
label the slip phases of the stickslip cycling. In D (coefficient of friction), the
of this system. This study provides the essential
linear fits of the static friction regions of the waveform are depicted in red.
framework for future stickslip models, not only for
engineering design, but also for understanding the
individual control of sound production and, ultimately,
explaining the evolutionary implications of varying stickslip
P=0.04, non-significant individual effects, P=0.07). Acoustic
frictional properties across spiny lobsters.
measurements of pulse rate were not significantly positively
California spiny lobsters shared the general acoustic and
correlated with plectrum speed (P>0.05).
morphological features as reported in other palinurids
Given the average plectrum velocity and shingle dimensions,
(Panulirus argus, Palinurus elephas and Panulirus longipes)
shingles were excited at approximately 1530shingless1, which
(Fig.4) (Meyer-Rochow and Penrose, 1974; Meyer-Rochow
was distinct from the average acoustic pulse rate of
and Penrose, 1976; Patek, 2002; Patek and Oakley, 2003;
90pulsess1.
Phillips et al., 1980; Smale, 1974). The mediallateral ridges
Stickslip friction measurements
present on each shingle in P. interruptus were similar to other
Panulirus species (Meyer-Rochow and Penrose, 1976;
The static coefficient of friction (s) ranged from 0.43.5
Moulton, 1957; Moulton, 1958; Patek, 2002), although they are
(mean 1.70.3). As predicted by the stickslip model (Fig.2),
not present in Palinurus (Patek, 2002) or Puerulus species
the coefficient of friction decreased rapidly when the plectrum
(S.N.P., personal observations). These surface asperities are
started to move (Fig.6). The maximum changes in over the
likely to increase the friction between the plectrum and file
course of each stickslip cycle were on average 1.10.3 and
surfaces compared to the smooth shingle surfaces of other
ranged from 0.13.6.
palinurids. The plectrum ridges appeared to be derived from
Slip velocity, stickslip frequency (equal to acoustic pulse
setae, given that the posterior limits of the plectrum ridges were
rate), s and change in varied as we applied a range of normal
constructed of fused setal bases (Fig.4E). The acoustic signal
forces from 0.55N. Although the stage was driven at a
features of P. interruptus were in the range of other spiny
constant velocity (10cms1), the plectrum slipped at velocities
lobsters (Patek and Oakley, 2003) with short duration pulses
ranging from 7.675.7cms1 (mean: 28.07.7cms1).
(7.9ms) and moderate pulse rates (90pulsess1).
Plectrum velocity was significantly positively correlated with
Given the basic principles of stickslip friction, can lobsters
Fn in six out of the nine individuals, but tests for individual
behaviourally control acoustic signal features and does variation
effects on the overall dataset were significant (whole model:
in shingle anatomy across species lead to differences in
R2=0.6058; F=14.01, P<0.0001; significant effects of Fn,
stickslip acoustics? One possible avenue for control of the
individual, and individual by trial). The change in was
system is through plectrum velocity and normal force. The
positively correlated with velocity, again with significant effects
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