First published online December 22, 2003
Journal of Experimental Biology 207, 535-544 (2004)
Published by The Company of Biologists 2004
doi: 10.1242/jeb.00789
Strikes and startles of northern pike (Esox lucius): a comparison of muscle activity and kinematics between S-start behaviors
Julie E. Schriefer1 and
Melina E. Hale1,2,3,*
1 Department of Organismal Biology and Anatomy, University of Chicago,
Chicago, IL 60637, USA
2 Committee on Neurobiology, University of Chicago, Chicago, IL 60637,
USA
3 Committee on Computational Neurobiology, University of Chicago, Chicago,
IL 60637, USA

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Fig. 1. Electrode placement. Northern pike (Esox lucius) were implanted
with seven electrodes to measure white epaxial muscle activity: one in the
left jaw and bilaterally in anterior, midbody and posterior positions in the
middle of the epaxial white muscle. Bars represent approximate range in the
positions of electrodes among the fish. The asterisk indicates the center of
mass (CM).
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Fig. 2. Movements of a strike and a startle of the northern pike (Esox
lucius). The feeding strike (AG) is characterized by isolated
caudal movement (CE) followed by a forward lunge accompanied by the
opening of the jaw (F,G). The startle response (HN) is characterized by
an initial S-shaped bend (J) followed by a stronger bend (L-bend; K) and a
propulsive tail stroke (LN) with no jaw opening. A comparison of the
feeding strike and escape startle demonstrates that initial movement is
restricted to the caudal region of the body during the feeding strike
(CE) while the startle involves significant rostral and caudal movement
(IK). Scale bar, 10 cm.
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Fig. 3. The angle of head movement during behavioral stages of strike and startle
responses. Values are means ± S.E.M.. The angles of head
movement during the S-bend and from the maximum S-bend to the maximum L-bend
are significantly greater for the startle than for the strike. There was no
significant difference in the angle of head movement in stage 2 of strike and
startle behaviors.
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Fig. 4. Durations of kinematics stages of S-start behaviors for the northern pike
(Esox lucius). Values are means ± S.E.M.. The
durations of the combined stages 1 and 2 movements, the S-bend and stage 1
were significantly greater for strikes than for startles. By contrast, there
was no significant difference in L-bend or stage 2 durations between strikes
and startles.
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Fig. 5. Mean peak center of mass velocity (A) and acceleration (B) of S-start
behaviors for the northern pike (Esox lucius). Values are means
± S.E.M.. Mean peak linear velocity and linear acceleration
were measured during the propulsive movement in stage 2. Neither velocity nor
acceleration was significantly different between strike and startle
behaviors.
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Fig. 6. Examples of electromyogram (EMG) patterns from S-start strike and startle
behaviors. EMGs from rostral, midbody and caudal axial white muscle on both
sides of the body and from left side jaw adductor muscle are shown. S, L and
End indicate the end of the S-bend, L-bend and stage 2, respectively. The
y-axis for jaw adductor EMGs of strikes and startles and for left
caudal activity of the strike range from 1.0 mV to 1.0 mV while all
other graphs range from 0.6 mV to 0.6 mV.Traces are aligned to first
onset of activity with a vertical broken line. During the strike, caudal
muscle activity (right side) occurs prior to more rostral activity; during the
startle, there is near simultaneous activity of rostral and midbody muscle on
one side of the body, in this case the left, with caudal muscle on the
opposite side. During the strike, jaw muscle activity onset is considerably
delayed from first onset of axial activity, while during the startle the jaw
muscle is active from initiation of axial muscle activity.
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Fig. 7. Summary of the electromyographic recordings of S-start strike and startle
muscle activity. The left margin of each bar is mean onset of activity and the
right side is mean offset. Trials were aligned by setting the first onset of
muscle activity to zero. Error bars represent the standard error of onset time
(left) and duration (right). There is no standard error for initial caudal
activity during the strike because this activity was, without exception, the
first activity recorded for the strike. During the strike, initial caudal
activity is followed by rostral and midbody activity on both sides of the
body. Jaw adductor muscle activity is delayed relative to axial activity.
During the startle response, jaw muscle is coactive with contralateral caudal
muscle on one side of the body and rostral and midbody ipsilateral muscle on
the opposite side. The initial activity is immediately followed by ipsilateral
caudal muscle and stage 2 contralateral activity.
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Fig. 8. Frequency distributions of adductor muscle electromyogram (EMG) onset
times. There were non-overlapping distributions of adductor muscle onset time
between strike and S-start escape behaviors. Onset of adductor muscle activity
during the startle took place within 5 ms of the initiation of axial activity
while during the strike the delay between axial and cranial activity was
longer and more variable.
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Fig. 9. Diagram of motor control and function of fast-start behaviors, modified
from fig. 5 of Domenici and
Blake (1997 ). The designations
of whole-body S-start versus caudal S-start reflect the differences
in rostral bending during the initial movements of strike and startle
behaviors. This work confirms the role of an independent S-start behavior that
functions as a startle and demonstrates differences in the motor control of
strike and startle behaviors. While there are indications that these behaviors
are controlled by different, but possibly overlapping, neural circuits, neural
mechanisms have yet to be identified.
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© The Company of Biologists Ltd 2004