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<br />synaptic potential. . ., polarity inversions due to openings
<br />of the circuit, etc." However, some of the concepts
<br />established by the Biarritz researchers are difficult to
<br />understand and have been questioned by other
<br />researchers (Hume, 1984; Sharber, personal
<br />communication).
<br />In 1991 during an electrofishing-injury workshop in
<br />Bozeman, Montana, N.G. Sharber introduced another
<br />explanation for the responses of fish to an electric field.
<br />According to this theory, often referred to as the Bozeman
<br />paradigm, the observed responses of fish, including
<br />muscular seizures resulting in spinal and related injuries,
<br />are comparable to responses of humans and other animals
<br />subjected to electroconvulsive therapy and can be
<br />similarly explained as phases of epilepsy, specifically
<br />automatism, petit mal, and grand mal (Sharber and Black,
<br />1999; Sharber et aI., 1994, 1995; Sharber, personal
<br />communication).
<br />How the underlying concepts of the Biarritz para-
<br />digm fit in the context of the Bozeman paradigm, and vice
<br />versa, has yet to be well explored. Because the phases of
<br />epilepsy are understood to be disorders of cerebral func-
<br />tion, Sharber et al. (I 994), and Sharber and Black (I999)
<br />suggested that the electric-field responses observed in
<br />fish are due to various levels of overstimulation of the
<br />central nervous system, either directly to the brain or
<br />short-circuited through the spinal cord. However, other
<br />researchers, including Haskell et al. (I 954), Vibert (I 963,
<br />1967b), Lamarque (I 967a, 1990), Edwards and Higgins
<br />(I 973), and Wydoski (I980), concluded that the various
<br />responses elicited in fish by an electric field are the result
<br />of direct stimulation of not only the central nervous sys-
<br />tem, which controls voluntary reactions, but also the au-
<br />tonomic nervous system, which controls involuntary
<br />reactions, and muscles themselves. Haskell et al. (I 954)
<br />and Lamarque (I967a, 1990) demonstrated that tetany in
<br />DC and muscular bends of the body toward the anode
<br />upon circuit closure in DC, or repeatedly in PDC, can be
<br />induced by direct overstimulation of efferent nerves or
<br />nerve endings associated with muscles. In those experi-
<br />ments, either efferent nerves were severed from the spi-
<br />nal cord, or the spinal cord was destroyed or removed
<br />prior to electric-field exposure. Muscular bends of the
<br />body often resulted in what was or would have been
<br />movement (taxis) towards the anode.
<br />There is surely some truth to both paradigms, and
<br />perhaps a better understanding of the responses of fish
<br />to electric fields will require an integration of the two
<br />(possibly along with aspects of the power-transfer theory
<br />discussed below). The major intensity-dependent re-
<br />sponses of fish described by both paradigms (reactive
<br />detection, undirected or inhibited swimming and taxis,
<br />and narcosis and tetany) are illustrated in Fig. 11 and
<br />discussed later in more detail. The electro-physiological
<br />
<br />SNYDER 23
<br />
<br />mechanisms involved in epilepsy and electroconvulsive
<br />therapy might or might not be much better understood
<br />than those for the responses of fish to electric fields. In
<br />either case, a collaboration of biologists, including ex-
<br />perts in neuro-physiology, should be fruitful for both dis-
<br />ciplines. Certainly, the observed results of the Biarritz
<br />experiments and others mentioned above are valid under
<br />the conditions in which they were performed, but a much
<br />more complete and defmitive understanding of the electro-
<br />physiological mechanisms involved is needed to better
<br />determine what electrical-field parameters and conditions
<br />will optimize desired electrofishing responses and mini-
<br />mize injury and other adverse effects.
<br />
<br />Theory of Power Transfer
<br />from Water to Fish
<br />
<br />Kolz and Reynolds (1989a) suggested that
<br />electro fishing should be viewed as a power-related phe-
<br />nomenon. More specifically, they hypothesized that the
<br />responses of fish to electric fields are directly related to
<br />the magnitude of power density (product of voltage gra-
<br />dient and current density) in the fish and that the in-fish
<br />power-density threshold for each response is constant
<br />(fixed) and independent of water conductivity. Accord-
<br />ing to their theory of power transfer (Kolz, 1989a), when
<br />water conductivity (cw) equals effective fish conductiv-
<br />ity (Cj), 100% of the power density in the water is trans-
<br />ferred to the fish (applied power density in the water, Dw,
<br />equals power density in the fish, Dr). But, as water con-
<br />ductivity either increases or decreases relative to the ef-
<br />fective conductivity of the fish (conductivity mismatch),
<br />power transfer to the fish is progressively less efficient.
<br />To establish or maintain a desired level of power density
<br />in the fish under conditions of conductivity mismatch
<br />(perhaps just above the threshold for a specific response),
<br />power density in the water must be progressively in-
<br />creased beyond that of the match condition in accord
<br />with the relation Dwl Dj = (I + qi I (4q), where q = cwl cf
<br />(the conductivity mismatch ratio; Kolz, 1989a). Subscript
<br />frepresents effective in-fish values which match corre-
<br />sponding in-water values, subscript w, at the minimum of
<br />the curve represented by this equation.
<br />When plotted on the log-log graph of power-density
<br />ratio versus water-conductivity ratio in Fig. 12, or the
<br />unique four-way log graph (Figs. 6 and 13) used by Kotz
<br />and Reynolds (1989b), the above relation yields what Kolz
<br />and Reynolds referred to as a normalized curve for pre-
<br />dicting the increase in applied power-density needed to
<br />maintain a constant level of power-density in a fish (the
<br />curve minimum) as water conductivity changes. Note that
<br />the curve is symmetrical with a rounded bottom, like an
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