Hollmann, M., Engelmann, J., and G. von der Emde. 2008. ÒDistribution, density, and
morphology of electroreceptor organs in mormyrid
weakly electric fish: anatomical investigations of a receptor mosaic.Ó Journal
of Zoology, 276, 149-158.
The
electric fish Gnathonemus petersii uses
electroreception instead of vision for orientation and foraging presumably due
to environmental restrictions on vision (it lives in murky African streams). It
does so through a process called electrolocation,
during which the fish produces an electrical field around its body using its
electric organ (located in the tail) and detects and analyzes distortions in
the field caused by surrounding objects using electroreceptor organs
distributed across the surface of its body. Both active and passive electrolocation is used in this system. Gn. petersii [sic]
has three kinds of electroreceptors: ampullary organs (which detect low-frequency electric
fields, and which are used passively), ÔKnollenorgansÕ
(which respond to high frequencies and which are used when communicating with conspecifics), and mormyromasts
(which respond to the fishÕs own electric organ discharges [EODs] and changes
to the fishÕs electric field caused by nearby objects; used for active electrolocation). The mormyromasts
are the most abundant type found in Gn. petersii.
Previously
it had been proposed that this species possessed two Ôelectrosensory
foveae,Õ one of which was at the chin area (called the ÔSchnauzenorganÕ)
and the other of which was between the mouth and nares
(called the Ônasal regionÕ). It was known that these two regions had high
electroreceptor densities and were overrepresented in the brain. The authors were
interested in illuminating the spatial patterns of mormyromast
distribution across the bodies of five different species of mormyrids.
They used 22 specimens of Gn. petersii and one specimen each of four other species.
Using electron microscopy, the researchers counted the number of mormyromast pores on the Schnauzenorgan,
the nasal region, and the back, and measured their inner and outer diameters.
They found
that the Schnauzenorgan of Gn. petersii was significantly more mormyromast-dense than other regions, possessing up to 65
receptors per square millimeter at the tip (which also contained many ampullary organs and Knollenorgans).
The dorsal side of the Schnauzenorgan had significantly
more mormyromasts than the other three sides, which
did not differ significantly from each other. They also found that the nasal
region was rich in mormyromasts as well, but did not
have as many on average (4.6 per square millimeter). The density of these
receptors decreased with distance from these two regions, and total number of mormyromasts did not vary much between individuals despite
size differences (resulting in decreasing density as body length increases). Though
other species had differing numbers of mormyromasts,
the spatial distribution of them across their bodies followed the general
pattern observed in Gn. petersii.
Additionally,
the researchers found that the outer diameters of the mormyromasts
on the dorsal side of the fish were significantly wider than those at the Schnauzenorgan, but did not differ significantly nasal
region mormyromasts. The inner diameters were also
wider on the dorsal side of the fish compared to other regions (with the
exception of the lateral sides of the head, where inner pore diameter did not
significantly differ from that of the dorsal pores). The authors found a highly
significant correlation between outer and inner diameters of the mormyromast pores.
The
authors suggest that one cause for the difference in mormyrmast
diameters (increasing diameter towards the back of the fish, though density
decreases) is increased sensitivity (as a size effect: larger inner and outer
diameters correspond to larger-diameter canals and total size of the organ,
respectively). The larger overall organs on the dorsal side of the fish may be
more sensitive (increased number of receptor cells) than those at the Schnauzenorgan. The authors conclude that their results are
in concordance with the hypothesis that two electrical foveae are present in Gn. petersii,
one at the Schauzenorgan and one in the nasal region.
However, they could not claim that these two foveae were really discrete, as
the density of receptor organs decreases continuously from the anterior to
posterior end of the fish. They do suggest that, because the Schnauzenorgan is moved from side to side and over objects
during foraging or exploration, while the nasal region is held Òat a fixed
angle while the fish is swimming,Ó it may be possible that the Schnauzenorgan and nasal region are functionally distinct.
They go on to hypothesize that the Schnauzen organ is
a fovea for near-field prey detection/identification Òequipped with a focusing
mechanism (funneling of current and movement of the chin),Ó while the nasal region
is for long-range detection of objects.
This
studyÕs findings are in agreement with information presented in class about
weakly electric fish: the high frequencies of the fields they use for electrolocation and communication, the concentration of electroreceptors at the head region, and the use of
spatially differing electric field disturbances across the surface of the body
to ÒmapÓ images. Additionally, this study suggests that decreasing mormyrmast density from the anterior to posterior end of
the fish is accompanied by increasing sensitivity of each mormyrast
in the same direction, the significance of which is not yet clear.