First published online August 30, 2006
Journal of Experimental Biology 209, 3636-3651 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02403
Modeling the electric field of weakly electric fish
David Babineau1,
André Longtin1 and
John E. Lewis2,*
1 Department of Physics
2 Department of Biology, University of Ottawa, Ottawa, Ontario K1N 6N5,
Canada

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Fig. 1. Electric field models. (A) Complete view of the model geometry, composed of
an aquarium, grounding and reference electrodes and the model fish. (B)
Close-up of morphologically accurate `fish' model consisting of a thin skin
layer, an electric organ (EO) and an interior body component (between the EO
and skin). The EO is 15.47 cm long and 0.08 cm thick, the skin is 0.01 cm
thick and the fish is 21cm long and 1.4 cm wide. (C,D) Geometrically simple
models used for studying (C) different fish tapers (see text for explanation)
and (D) various body and skin properties. To achieve different tapers in C,
the left side of the model (here shown for a width of 1.82 cm) is varied. The
EO length and skin thickness for C and D are the same as in B. Model C is
referred to as the `taper' model and model D is referred to as the `box'
model. x and y axes, as well as grounding and reference
electrodes, are not to scale.
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Fig. 2. Optimal model parameters. (A) The optimal EO current density profile (red),
called `skewed', is the sum of two Gaussian curves: a narrow (dotted green)
sink in the tail region and a wide (dotted blue) source in the mid-body.
Rostral curve is centered 15 cm from the tip of the head and has a standard
deviation of 4.46 cm; caudal curve is centered 18.04 cm from the tip of the
head with a standard deviation of 0.5 cm. The ratio of the peak amplitudes of
the two curves is 1:8.38. (B) Optimal conductivity values for the EO (blue),
skin (green) and body (red). Optimal parameter values are normalized to one
and the errors associated with each optimal parameter value are set to zero.
Abscissa values are given as ratios of the optimal parameter value (for each
respective curve) and ordinate values are given as field RMS% errors above
minimal error (28.6%; see text for details on error measure). Optimal
conductivity values are: EO, 0.927 S m-1; body, 0.356 S
m-1; skin, 0.0017 S m-1. Although all parameters here
were varied homogeneously, it should be noted that the optimal skin
conductivity is not uniform along the length of the fish (see
Fig. 8A).
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Fig. 3. Model calibration. (A) 2D field potential surrounding the fish. Top:
experimental data obtained from Assad
(Assad, 1997 ). Bottom:
simulated values obtained with optimal parameters (including non-uniform skin
conductivity). Color maps represent potential with respect to an electrode
placed laterally to the fish, near its zero-potential line (as in
Fig. 1A). All values below
approx. -7.6 mV have been mapped to dark blue in order to show a better
contrast between the positive and negative regions of the dipolar field
(measured potential in tail region reaches approx. -30 mV). The zero-potential
line is shown in yellow. (B) Un-weighted % error and (C) absolute potential
differences between data and simulated values found in A. Broken lines show
cross-sections at which the potentials are plotted in (D) and (E). Potential
differences in C greater than 5 mV are all mapped to dark red. (D) Potential
along dotted line near head (5 cm caudal from the tip of the head) in B for
model (red) and data (blue). (E) Potential along dotted line near tail (20 cm
caudal from the tip of the head) in B for model (red) and data (blue).
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Fig. 4. Electric field characterization: study of the fish's filtering properties
and comparison with an ideal voltage divider. (A) Potential values at the
electric organ (blue) and at the interior (green) and exterior (red) skin
boundaries along the EO segment (zero corresponds to the rostral end, one to
the caudal end), for a 5-cycle sinusoidal current density (fish model). (B)
Right axis, green trace shows the normalized energy of the exterior skin
potential curve as a function of rostro-caudal position along the fish body
(fish model; solid line was obtained using the `shape-preserving interpolant'
fitting function in MATLAB). This energy quantifies the level of `smoothness'
of a given trace (see text for details). Left axis, blue trace shows the
filtering along the EO segment for a 50-cycle sinusoidal current density (fish
model). Filtering quantifies how much the energy has decreased from the EO to
the skin (see text for details). The red line represents the start of the EO
in the fish model (x=4.42 cm; see
Fig. 1B). (C) Filtering along
the EO segment for the fish model (green) and for the taper model with low
(black) and high (blue) taper values. Red broken line represents the location
of taper change in the fish model. (D) Comparison between the fish and an
ideal voltage divider (taper model, taper=0.05). Theoretical (green) and
simulated (blue) transdermal potentials along the EO segment for a 5-cycle
sinusoidal current density. Red trace shows the difference between simulation
and theory (see text for details).
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Fig. 5. Effect of object location on electric images (fish model). (A) Electric
images for an object located 2 cm (red), 2.5 cm (blue), 3.5 cm (green), 5 cm
(black) and 10 cm (pink) lateral of the fish's midline (5 cm caudal from the
tip of the head). The electric image is calculated as the change in
transdermal potential caused by the object. (B) Electric images for an object
located 5 cm (red), 7.5 cm (black), 10 cm (green), 12.5 cm (blue), 15 cm
(pink), 17.5 cm (orange) caudal from the start of the head at a lateral
distance of 3 cm (from the fish's midline). The simulated object for A and B
is a metal disc (brass; conductivity=2.13x107 S
m-1; 1 cm radius).
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Fig. 6. Effect of EO current density profile on electric images (fish model). (A)
Normalized current densities versus normalized EO position. The green
trace shows the 1-cycle sinusoidal current density; the blue trace shows the
optimal skewed current density (sum of two Gaussian curves); the red trace
shows the `impulse' current density (single Gaussian function offset in order
to have a mean of zero). All current densities are zero-mean. (B) Normalized
electric images produced by a metal disc located in the middle of the fish
(black vertical line), 3 cm lateral from the midline (see inset), for skewed
(blue), impulse (red) and sinusoidal (green) current density profiles. The
green markers `xR', `x0' and `xC' illustrate the
`x' or rostro-caudal positions of the three points that characterize
bimodal electric images: the rostral peak, the zero-crossing (located between
rostral and caudal peaks) and the caudal peak, respectively.
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Fig. 7. Positions and amplitudes of the bimodal electric image's characteristic
points (`xR', `x0' and `xC') for different
rostro-caudal object locations (box model; optimal uniform skin conductivity;
object centered 3 cm away from the fish's midline). (A,C) Normalized peak
positions for sinusoidal (A) and skewed (C) current densities. Peak positions
and object locations are normalized with respect to EO coordinates, with the
rostral side corresponding to zero. Blue and green traces are the positions of
the rostral and caudal peaks, respectively, while red traces show the
positions of the zero-crossings. The solid black curve shows the identity
line, where the location of the electric image's dominant peak at the skin
corresponds exactly to the rostro-caudal location of the object. Black broken
lines delimit zones in which certain characteristic curves are closest to the
solid black trace: e.g. in C, the blue trace (xR) is closest to the
identity line in the rostral zone, i.e. the bimodal image's rostral peak is
closest to the object's actual location in this zone. (B,D) Absolute
potentials of the bimodal electric image's rostral and caudal peaks for
sinusoidal (B) and skewed (D) current densities. Blue traces show absolute
potential values for the rostral peak while caudal peak values are shown in
green.
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Fig. 9. Effect of object size and lateral distance on bimodal electric images (box
model; skewed current density; optimal uniform skin conductivity). Lateral
distance is measured with respect to object centers. (A) Un-normalized (actual
amplitudes) and (C) normalized (with respect to the caudal peak's amplitude)
electric images produced by three different-sized metal objects located half
way along the fish's body, 4 cm away from the midline: 0.5 cm (blue), 1.1 cm
(green) and 2 cm (red) radius (see inset). Both images are plotted
versus the normalized box model length, in which the rostral side
corresponds to zero. (B) Peak-to-peak potentials of the electric images found
in A as a function of lateral distance away from the fish's midline for the
three different objects (object sizes same as in A; see inset). (D) Delta,
defined as the difference between the rostral and caudal peak locations, found
in either A or C, as a function of lateral distance from the fish's midline
for the three different objects; delta for the 1.1 cm object is shown in A and
C. Black broken lines in B and D show the distance (4 cm away from the fish's
midline) at which the objects were located for the images shown in A and
C.
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Fig. 10. Effect of object size and lateral distance on bimodal electric images. All
panels are the same as in Fig.
9 except that lateral distance is measured here with respect to
the object's edge. (A) Un-normalized and (C) normalized electric images
produced by three different-sized metal objects located half way along the
fish's body, with the object edges 2 cm away from the midline: 0.5 cm (blue),
1.1 cm (green) and 2 cm (red) radius (see inset). Red traces are the same as
in Fig. 9A,C since a lateral
edge distance of 2 cm for the 2 cm-radius object corresponds to a lateral
object center distance of 4 cm. (B) Peak-to-peak potentials of the electric
images found in A as a function of lateral distance away from the fish's
midline for the three different objects (object sizes same as in A; see
inset). (D) Delta found in either A or C, as a function of lateral distance
from the fish's midline for the three different objects; delta for the 1.1 cm
object is shown in A and C. Black broken lines in B and D show the distance (2
cm away from the fish's midline) at which the objects were located for the
images shown in A and C.
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© The Company of Biologists Ltd 2006