Pantodon is a lovely
little fish that lives just below the water's surface in Nigeria. As a
consequence of its optical ecology, its eye views into air and water simultaneously.
We are currently studying two specific behaviors, feeding and the shadow
reflex, and relating them to its brain organization and the visual electrophysiology
of cells at different loci of its visual pathway. In particular, we have
found a diencephalic nucleus (nucleus RostroLateralis) that is concerned
only with vision in air. This nucleus receives direct visual input from
the ipsilateral ventral retina and indirect visual input from cells of
the optic tectum (bilaterally) that are postsynaptic to the ventral retina.
This nucleus is found in less than a dozen of the >20,000 species of fish.
Interestingly, it is found in Anableps anableps, another and unrelated
species that simultaneously sees in air and water.
The Eye of Pantodon
The eye of Pantodon
is unusual because its visual field is closely reflected in eye and retinal
structure. The fundus of the eye is bisected by a modified falciform process
that extends from the nasal pole almost to the temporal pole. Blood vessels
emerge perpendicular to the falciform process to run on the surface of
the retina. A small region of continuity extends from ventrotemporal retina
to dorsotemporal retina at the temporal pole of the ora serrata. The embryonic
fissure extends from the nasal pole to the optic disc. I am currently studying
the non-symmetrical postembryonic addition of cells in the retina as a
probe to understand the morphological pressures that might account for
the extension of the falciform process into the temporal hemiretina. The
ratio of temporal / nasal >>3.
The visual field
is tripartite. The ventral hemiretina views into the air. The water surface
area through which all aerial rays pass is known as Snell's window. The
dorsal retina views into the water column and the region between (which
has an unusual retinal structure, see below) views the aquatic environment
as seen in a reflection from the aquatic side of the air-water interface.
The retina reflects this tripartite visual field in the following manner:
The falciform
process bisects the retina in the nasal hemiretina (left) and sits on the
surface of the temporal retina. The black tissue is due to highly melanized
chorioid epithelial cells that penetrate the retina via the optic nerve
at the disk. Cones are wider and larger in the aquatic viewing retina (right,top)
than the aerial viewing retina (right, bottom).
Behaviorally Relevant Neurology
Along with Dr. Hong Y. Yan of the T.H. Morgan School
of Biology, University of Kentucky, we have been studying the feeding behavior
of this fish using videotape techniques. We found that the fish feeds monocularly
and it only feeds on targets at the water surface. [In the numerous years
of observing this fish, I have seen only 1 example of feeding within the
water column.] It only captures a target from within 1 cm of an eye. It
is fairly clear that the sensory information involved with feeding is a
combination of vision and lateral line with visual input under bright illumination
acting in an inhibitory manner. Either vision alone or lateral line alone
are sufficient for feeding purposes, but the combination in low illumination
provides a more favorable sensory environment. Bright light seems to inhibit
the feeding process.
Although the fish will jump at targets in the air,
it does not jump at the targets. It jumps at the image of the target in
Snell's window. The figure at the left just below demonstrates that the
fish does not jump at the hanging cricket in air. Rather, as you can see
in the image below, the fish jumps at the image of the cricket in
Snell's window.
Each frame is a composite of a top view (above) and a side view
(below) taken simultaneously (with the aid of an overhead mirror). The
small arrows
in frame 4 identify the hanging cricket. The large arrows in figure
4 identify the vector of movement. As can be seen in frame 5, the fish
(albeit blurred),
is jumping at a target adjacent to its eye (ie., within Snell's window).
Since feeding only occurs from stimulation of the ventral retina, we
searched for a neurological correlate or neural element that can be found
in the visual pathway concerned with the superior visual field. In the
optic tectum, we found a distinctive neuron (below) whose dendrites appear
to integrate visual and lateral line input. Since these cells are predominately
(>85%) found in the tectual representation of the aerial visual field,
we feel that these cells signal a potential feeding target. Interestingly
and as a corollary, we also studied the distribution of these cells in
a fish (goldfish) that feeds throughout its visual field and sure enough,
the cells are found throughout the tectum. We are currently searching through
the tectum of other fish with uniquely peculiar feeding habits for the
tectal distribution of these cells.
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for further information (an abbreviated cv) about the studies of this lab, send email to me at saidel@crab.rutgers.edu