First published online January 31, 2007
Journal of Experimental Biology 210, 561-569 (2007)
Published by The Company of Biologists 2007
doi: 10.1242/jeb.02697
Orienting and avoidance turning are precisely computed by the predatory sea-slug Pleurobranchaea californica McFarland
Liudmila S. Yafremava1,
Christopher W. Anthony2,
Laura Lane2,
Jessica K. Campbell2 and
Rhanor Gillette1,2,3,*
1 Program in Biophysics and Computational Biology, University of Illinois at
Urbana-Champaign, University of Illinois, Urbana, IL 61801, USA
2 Department of Molecular and Integrative Physiology, University of Illinois
at Urbana-Champaign, University of Illinois, Urbana, IL 61801, USA
3 The Neuroscience Program, University of Illinois at Urbana-Champaign,
University of Illinois, Urbana, IL 61801, USA

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Fig. 1. Stimulus application protocol. Nine stimulus loci were distinguished on the
oral veil, with 0 designating midline, and +4 and 4 designating the
right and left tentacle, respectively.
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Fig. 2. Unilateral applications of chemotactile stimulus resulted in linear
relationship between the turn angle and stimulus locus. (A) Responses to the
appetitive stimulus betaine (crosses) and to the aversive stimulus taurine
(filled circles). (B) The slope of the relationship in A for appetitive
stimulus increases with the stimulus concentration. Values are means ±
s.e.m.; for correlation coefficients, see text.
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Fig. 3. Animals overshoot in orienting turns by approximately a factor of 4. (A) If
animals were turning to position the stimulus in the center of their oral
veil, then the angle of turn would relate to the stimulus locus according to
the relationship: angle=arctan (xk/4), where k is the ratio
of the oral veil width to the body length L (solid line without
datapoints in B). (B) The relationship determined from data (solid line with
data points, replotted from Fig.
2A) gives a slope 4 times steeper than the predicted relationship
for k=0.3.
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Fig. 4. Sequential turns stimulated by a single stimulus application. A significant
fraction of animals performed multiple turns in response to unilateral oral
veil stimulation. Three turns were most common, of which the first and third
turn angles had the animals going in a closely similar direction. The second
turn angle, to the opposite direction, was not correlated with the first or
third. See text for further details.
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Fig. 5. (A) Correlation analysis of 17 cases of sequential turns showed that the
angle of the third turn was highly correlated with and was not significantly
different from the angle of the first turn (r=0.84). (B) The second
angle and first angle were not correlated (r=0.08).
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Fig. 6. A summary model of function in turn behavior. Chemotactile afferents of
oral veil are conceived as being integrated in an interneuronal layer that
averages amplitude and laterality of chemotactile input. The output of the
interneuronal layer is translated to the turn motor network through a switch
mechanism regulated by the animal's appetitive state and memory of experience.
The switch acts to invert the laterality of the output to the turn motor
network to result in orienting or avoidance turns.
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© The Company of Biologists Ltd 2007