In order to clarify whether or not the electronegative olfactory mueosal potentials (EOG) are generator potentials, the effects of changed ionic enviroment were studied. The EOG decreased in amplitude and in some cases nearly or completely disappeared, when Na + in the bathing Ringer solution was replaced .by sucrose, Li +, choline +, tetraethylammoninm + (TEA), or hydrazine. In the K+-free Ringer solution, the negative EOG's initially increased and then decreased in amplitude, In Ringer's solution with increased K +, the negative EOG's increased in amplitude. When K + was increased in exchange for Na + in Ringer's solution, the negative EOG's decreased, disappeared, and then reversed their polarity (Fig. 6). Next, when the K + was replaced by equimolar sucrose, Li +, choline +, TEA+, hydrazine, or Na +, the reversed potentials recovered completely only in Na+-Ringer's solution, but never in the other solutions, Thus, the essential role of Na + and K + in the negative EOG's was demonstrated. Ba ++ was found t ° depress selectively the electropositive EOG, but it hardly :decreased and never increased the negative EOG. Hence, it is concluded that Ba ++ interferes only with CI-influx, and that the negative EOG's are elicited by an increase in permeability of the olfactory receptive membrane to Na + and K +, but not to CA-. From the ionic mechanism it is inferred that the negative EOG's are in most eases composites of generator and positive potentials.Since the pioneer work by Hosoya and Yoshida (1937) in the dog, and by Ottoson (1954Ottoson ( , 1956 in the rabbit and frog, the electrical phenomena elicited in the olfactory epithelium by application of odors have been extensively studied and much new information has been obtained. Besides the electronegative slow potentials of the " o n " type found by the above workers, and named "electro-olfactogram ( E O G ) " by Ottoson (1956), electronegative po-
A B S T R A C TIn the negative EOG-generating process a cation which can substitute for Na + was sought among the monovalent ions, Li +, Rb +, Cs +, NH4 +, and TEA +, the divalent ions, Mg ++, Ca ++, Sr ++, Ba ++, Zn++, Cd++, Mn ++, Co ++, and Ni ++, and the trivalent ions, A1 +++ and Fe II z. In Ringer solutions in which Na + was replaced by one of these cations the negative EOG's decreased in amplitude and could not maintain the original amplitudes. In K+-Ringer solution in which Na + was replaced by K +, the negative EOG's reversed their polarity. Recovery of these reversed potentials was examined in modified Ringer solutions in which Na + was replaced by one of the above cations. Complete recovery was found only in the normal Ringer solution. Thus, it was clarified that Na + plays an irreplaceable role in the generation of the negative EOG's. The sieve hypothesis which was valid for the positive EOG-generating membrane or IPSP was not found applicable in any form to the negative EOGgenerating membrane. The reversal of the negative EOG's found in K +-, Rb +-, and Ba++-Ringer solutions was attributed to the exit of the internal K +. It is, however, not known whether or not C1-permeability increases in these Na+-free solutions and contributes to the generation of the reversed EOG's.The ionic mechanisms underlying the generation of the electro-negative and positive "electro-olfactograms" (EOG's) have been studied in previous papers (Takagi and Wyse, 1965;Takagi, Wyse, and Yajima, 1966;Takagi, 1968; Takagi, Wyse, Kitamura, and Ito, 1968). Briefly, the results showed that the negative EOG's primarily depend upon the entry of Na+ and the simultaneous exit of K +, and that neither Li +, tetraethyl a m m o n i u m (TEA+), choline +, hydrazine, nor sucrose can substitute for Na+; the positive EOG's primarily depend upon the entry of CI-and the simultaneous exit of K +, and three anions, Br-, F-, and H C O~ can substitute for C1-.
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