◆❡✉r❛❧ ❞❛t❛❜❛s❡ ❛♥❛❧②s✐s ✐♥ ▼❛t❧❛❜ ✇✐t❤ P❛♥❞♦r❛✬s ❚♦♦❧❜♦① ❈❡♥❣✐③ ●ü♥❛② ❝❣✉♥❛②❅❡♠♦r②✳❡❞✉ ❉❡♣❛rt♠❡♥t ♦❢ ❇✐♦❧♦❣②✱ ❊♠♦r② ❯♥✐✈❡rs✐t② ❖❦✐♥❛✇❛ ❈♦♠♣✳ ◆❡✉r♦s❝✐✳ ❈♦✉rs❡ ✷✵✵✾ ❈✳ ●ü♥❛② ✭❊♠♦r② ❯♥✐✈❡rs✐t②✮ ◆❡✉r❛❧ ❞❛t❛❜❛s❡ ❛♥❛❧②s✐s ✇✐t❤ P❛♥❞♦r❛ ❖❈◆❈ ✷✵✵✾ ✶ ✴ ✶✸
❲❤② ✇♦✉❧❞ ■ ✉s❡ t❤❡ P❛♥❞♦r❛ t♦♦❧❜♦①❄ P❛♥❞♦r❛ ❝❛♥ ❜❡ ✉s❡❞ ❢♦r✿ ✶ ❊①tr❛❝t✐♥❣ ❡❧❡❝tr♦♣❤②s✐♦❧♦❣✐❝❛❧ ♣r♦♣❡rt✐❡s ❡✳❣✳✱ s♣✐❦❡s ❢r♦♠ ❛ ♠❡♠❜r❛♥❡ ✈♦❧t❛❣❡ tr❛❝❡ ✷ ❉❛t❛❜❛s❡ ❛♥❛❧②s✐s ♦❢ ♠♦❞❡❧ ♦r ❡①♣❡r✐♠❡♥t❛❧ ❞❛t❛ ❈r❡❛t✐♥❣ ❛ ❞❛t❛❜❛s❡ ❢r♦♠ ❛♥② ❞❛t❛ ❢♦r q✉❡r②✐♥❣ ❛♥❞ ♣❧♦tt✐♥❣ ✳ P✉tt✐♥❣ r❡s✉❧ts ❢r♦♠ ❛♥❛❧②s✐s ♦❢ ✈♦❧t❛❣❡ tr❛❝❡s ♦❢ ♠✉❧t✐♣❧❡ ♠♦❞❡❧s ✐♥t♦ ❛ ❞❛t❛❜❛s❡✳ ❆❞✈❛♥❝❡❞ ♦♣❡r❛t✐♦♥s ✇✐t❤ ❛ ❞❛t❛❜❛s❡✿ st❛t✐st✐❝s✱ ♠✉❧t✐✈❛r✐❛t❡ ❛♥❛❧②s✐s ✱ ❡t❝✳ P❛♥❞♦r❛✬s ♣❧♦tt✐♥❣ ❢✉♥❝t✐♦♥s ✭❝❛♥ ❛❧s♦ ❜❡ ✉s❡❞ ♦✉ts✐❞❡ ♦❢ ❞❛t❛❜❛s❡s✮✳ ❙❡❡ ●ü♥❛② ❡t ❛❧✳ ✭✷✵✵✾✮ ◆❡✉r♦✐♥❢♦r♠❛t✐❝s ❀ ❛❧s♦ ❅ ■◆❈❋ ❙♦❢t✇❛r❡ ❈❡♥t❡r ❛♥❞ ❖❈◆❈ ❢♦r✉♠✳ ❈✳ ●ü♥❛② ✭❊♠♦r② ❯♥✐✈❡rs✐t②✮ ◆❡✉r❛❧ ❞❛t❛❜❛s❡ ❛♥❛❧②s✐s ✇✐t❤ P❛♥❞♦r❛ ❖❈◆❈ ✷✵✵✾ ✷ ✴ ✶✸
❲❤❛t ✐s t❤❡ P❛♥❞♦r❛ t♦♦❧❜♦①❄ ▼❛✐♥ ❝♦♠♣♦♥❡♥ts ♦❢ P❛♥❞♦r❛✿ ❈✳ ●ü♥❛② ✭❊♠♦r② ❯♥✐✈❡rs✐t②✮ ◆❡✉r❛❧ ❞❛t❛❜❛s❡ ❛♥❛❧②s✐s ✇✐t❤ P❛♥❞♦r❛ ❖❈◆❈ ✷✵✵✾ ✸ ✴ ✶✸
▲♦❛❞✐♥❣ ❛ ♠❡♠❜r❛♥❡ ✈♦❧t❛❣❡ tr❛❝❡ P❛♥❞♦r❛ ❝❛♥ r❡❛❞ t❤❡ ✜❧❡ ❢♦r♠❛ts ❢r♦♠✿ ✶ ❙✐♠✉❧❛t♦rs✿ ◆❡✉r♦♥ ●❡♥❡s✐s ✷ ❉❛t❛ ❛❝q✉✐s✐t✐♦♥ ♣r♦❣r❛♠s✿ ❆❧❧ ◆❡✉r♦❙❤❛r❡✲❝♦♠♣❛t✐❜❧❡ ❛❝q✉✐s✐t✐♦♥ ❞❡✈✐❝❡s ✭❆❧♣❤❛ ❖♠❡❣❛✱ ❈❛♠❜r✐❞❣❡ ❊❧❡❝tr♦♥✐❝ ❉❡s✐❣♥✱ ◆❡✉r♦❊①♣❧♦r❡r✱ P❧❡①♦♥✱ ❘✳❈✳ ❊❧❡❝tr♦♥✐❝s ■♥❝✳✱ ❚✉❝❦❡r✲❉❛✈✐s ❚❡❝❤♥♦❧♦❣✐❡s✱ ❛♥❞ ❈②❜❡r❦✐♥❡t✐❝s ■♥❝✳✱ ❡t❝✳✮ ✸ ❖t❤❡r✿ ❍✐❡r❛r❝❤✐❝❛❧ ❞❛t❛ ❢♦r♠❛t ✭❍❉❋✺✮ ❈✳ ●ü♥❛② ✭❊♠♦r② ❯♥✐✈❡rs✐t②✮ ◆❡✉r❛❧ ❞❛t❛❜❛s❡ ❛♥❛❧②s✐s ✇✐t❤ P❛♥❞♦r❛ ❖❈◆❈ ✷✵✵✾ ✹ ✴ ✶✸
❆♥❛❧②③✐♥❣ ❛ ♠❡♠❜r❛♥❡ ✈♦❧t❛❣❡ tr❛❝❡ ❇② ❡①tr❛❝t✐♥❣ ❡❧❡❝tr♦♣②s✐♦❧♦❣✐❝❛❧ ❝❤❛r❛❝t❡r✐st✐❝s ▼❡❛s✉r❡ s♣✐❦❡ s❤❛♣❡ ❛♥❞ ✜r✐♥❣ r❛t❡ ♣r♦♣❡rt✐❡s ▼❡❛s✉r❡ s❛❣✱ s♣✐❦❡ ❛❞❛♣t❛t✐♦♥ ❛♥❞ ❝✉rr❡♥t r❡s♣♦♥s❡ ♣r♦♣❡rt✐❡s ❈❛♥ ❜❡ ❞♦♥❡ r❡♣❡t✐t✐✈❡❧② ❢♦r ❛ ❧❛r❣❡ ♥✉♠❜❡r annotated spike characteristics ♦❢ ♠♦❞❡❧s −60 voltage [mV] −70 ❈❛♥ ❜❡ ❡♥t❡r❡❞ ✐♥t♦ ❛ ▼❛t❧❛❜ ❞❛t❛❜❛s❡ −80 −90 ❉❊▼❖ −100 1450 1500 1550 time [ms] ❈✳ ●ü♥❛② ✭❊♠♦r② ❯♥✐✈❡rs✐t②✮ ◆❡✉r❛❧ ❞❛t❛❜❛s❡ ❛♥❛❧②s✐s ✇✐t❤ P❛♥❞♦r❛ ❖❈◆❈ ✷✵✵✾ ✺ ✴ ✶✸
❈r❡❛t✐♥❣ ❛ ❞❛t❛❜❛s❡ ❢r♦♠ ❛r❜✐tr❛r② ❞❛t❛ ❈r❡❛t❡ ❛ ✷ × ✷ ❞❛t❛❜❛s❡ ♠❛tr✐①✿ ❲✐t❤ ▼❛t❧❛❜ ❝♦❞❡✿ ❃✗❃ ❞❜❴♦❜❥ ❂ t❡sts❴❞❜✭❬✶ ✷❀ ✸ ✹❪✱ ④✬❝♦❧✶✬✱ ✬❝♦❧✷✬⑥✱ ④✬r♦✇✶✬✱ ✬r♦✇✷✬⑥✱ ✬❛ ✷①✷ ❉❇✬✮ ❈✳ ●ü♥❛② ✭❊♠♦r② ❯♥✐✈❡rs✐t②✮ ◆❡✉r❛❧ ❞❛t❛❜❛s❡ ❛♥❛❧②s✐s ✇✐t❤ P❛♥❞♦r❛ ❖❈◆❈ ✷✵✵✾ ✻ ✴ ✶✸
❈r❡❛t✐♥❣ ❛ ❞❛t❛❜❛s❡ ❢r♦♠ ❛♥❛❧②s✐s ♦❢ tr❛❝❡s ❯s✐♥❣ ❛ ❞❛t❛s❡t✿ ✶ ❃❃ ♠②❴❞❛t❛s❡t❴♦❜❥ ❂ ♠②❴❞❛t❛s❡t❴❝❧❛ss ✭ ✬ ❞❛t❛ ✴ ∗ ✳ ❜✐♥ ✬ ✱ ❛r❣✉♠❡♥ts ✳ ✳ ✳ ✮ ✸ ❃❃ ♠②❴❞❛t❛❜❛s❡❴♦❜❥ ❂ ♣❛r❛♠❴t❡sts❴❞❜ ✭ ♠②❴❞❛t❛s❡t❴♦❜❥ ✮ ✺ ❃❃ s♦rt❡❞❴♦❜❥ ❂ s♦rtr♦✇s ✭ ♠②❴❞❛t❛❜❛s❡❴♦❜❥ ✱ ✬ ❆P❴❛♠♣❧✐t✉❞❡ ✬ ✮ ❈✳ ●ü♥❛② ✭❊♠♦r② ❯♥✐✈❡rs✐t②✮ ◆❡✉r❛❧ ❞❛t❛❜❛s❡ ❛♥❛❧②s✐s ✇✐t❤ P❛♥❞♦r❛ ❖❈◆❈ ✷✵✵✾ ✼ ✴ ✶✸
❉❛t❛❜❛s❡ ❛♥❛❧②s✐s✿ ◗✉❡r②✐♥❣ ❃❃ ❞❜❴♦❜❥✷ ❂ ✷ ❞❜❴♦❜❥ ✭✶✿✶✵ ✱ ④ ✬ ♥❡✉r♦♥❴✐♥❞❡① ✬ ✱ ✬ ❢ ✐ r ❡ ❴ r ❛ t ❡ ✬ ⑥✮ ❃❃ ❞❜❴♦❜❥✷ ❂ ✹ ❞❜❴♦❜❥ ✭ ❞❜❴♦❜❥ ✭ ✿ ✱ ✬ ♥❡✉r♦♥❴✐♥❞❡① ✬ ✮ ❂❂ ✹✻ ✱ ✿ ✮ ❃❃ ❞❜❴♦❜❥✷ ❂ ✻ ❞❜❴♦❜❥ ✭ ❛♥②❘♦✇s ✭ ❞❜❴♦❜❥ ✭ ✿ ✱ ✬ ♥❡✉r♦♥❴✐♥❞❡① ✬ ✮ ✱ ❬ ✹ ✻ ❀ ✺✻❀ ✶✷❪✮ ✱ ✿ ✮ ✽ ❃❃ ❞❜❴♦❜❥✷ ❂ ❞❜❴♦❜❥ ✭ ❞❜❴♦❜❥ ✭ ✿ ✱ ✬ ♥❡✉r♦♥❴✐♥❞❡① ✬ ✮ ⑦❂ ✹✻ ✫ ✶✵ ✭ ❞❜❴♦❜❥ ✭ ✿ ✱ ✬ ❈■P ✬ ✮ ❃ ✶✵✵ ⑤ ❞❜❴♦❜❥ ✭ ✿ ✱ ✬ r ❛ t ❡ ✬ ✮ ❁❂ ✺✵ ✮ ✱ ✿ ✮ ✱ ✶✷ ❃❃ ❞❜❴♦❜❥✷ ❂ ♠♦❞❡❧❴❞❜❴♦❜❥ ✭ ❛♥②❘♦✇s ✭ ♠♦❞❡❧❴❞❜❴♦❜❥ ✭ ✿ ✱ ✬ r ❛ t ❡ ✬ ✮ ✱ ✶✹ ♥❡✉r♦♥❴❞❜❴♦❜❥ ✭ ✿ ✱ ✬ r ❛ t ❡ ✬ ✮✮ ✱ ✿ ✮ ❈✳ ●ü♥❛② ✭❊♠♦r② ❯♥✐✈❡rs✐t②✮ ◆❡✉r❛❧ ❞❛t❛❜❛s❡ ❛♥❛❧②s✐s ✇✐t❤ P❛♥❞♦r❛ ❖❈◆❈ ✷✵✵✾ ✽ ✴ ✶✸
▼✉❧t✐✈❛r✐❛t❡ ❛♥❛❧②s✐s ❚❡tr♦❞♦t♦①✐♥ ❜❧♦❝❦ ❡✛❡❝ts ♦♥ ✜r✐♥❣ r❛t❡ ♦❢ ❣❧♦❜✉s ♣❛❧❧✐❞✉s ♥❡✉r♦♥s ✇✐t❤ ❝✉rr❡♥t ✐♥❥❡❝t✐♦♥ steady−rate (+100pA) [Hz] with 0 to 7 nM TTX [Hz] 30 −3 rate change 28 n=3 −4 26 −5 24 −6 22 n=3 20 −7 0 7e−09 1 TTX [M] ❈✳ ●ü♥❛② ✭❊♠♦r② ❯♥✐✈❡rs✐t②✮ ◆❡✉r❛❧ ❞❛t❛❜❛s❡ ❛♥❛❧②s✐s ✇✐t❤ P❛♥❞♦r❛ ❖❈◆❈ ✷✵✵✾ ✾ ✴ ✶✸
▼✉❧t✐✈❛r✐❛t❡ ❛♥❛❧②s✐s ✭■✮ ❙✐❢t✐♥❣ t❤❡ ❞❛t❛❜❛s❡ t♦ ✜♥❞ ❡✛❡❝ts ♦❢ ♣❛r❛♠❡t❡rs ❙❛♠♣❧❡ ✇✐t❤ ✸ ◆❡✉r♦♥s✿ P✐❝r♦❚① ✵ . ✵✵✵✶ ✵ . ✵✵✵✶ ✵ . ✵✵✵✶ ✵ . ✵✵✵✶ ✵ . ✵✵✵✶ ✵ . ✵✵✵✶ ❑②♥❆❝✐❞ ✵ . ✵✵✶ ✵ . ✵✵✶ ✵ . ✵✵✶ ✵ . ✵✵✶ ✵ . ✵✵✶ ✵ . ✵✵✶ ❚❚❳ ✵ ✼ ❡ − ✵✾ ✵ ✼ ❡ − ✵✾ ✵ ✼ ❡ − ✵✾ ❆♣❛♠✐♥ ✵ ✵ ✵ ✵ ✵ ✵ ❞r✉❣ ✹❆P ✵ ✵ ✵ ✵ ✵ ✵ ◆❡✉r♦♥■❞ ✶✵✼ ✶✵✼ ✶✵✽ ✶✵✽ ✶✶✵ ✶✶✵ ❉✶✵✵♣❆ st❡❛❞② r❛t❡ ✷✺ . ✾✾✽✷ ✶✾ . ✻✵✺✻ ✷✾ . ✾✻✼✸ ✷✷ . ✼✻✷✽ ✷✸ . ✽✹✹✸ ✷✵ . ✾✼✹✹ ❋♦❝✉s ♦♥ ❝❤❛♥❣❡s ✇✐t❤ ❚❚❳✿ P❛❣❡ ✶ P❛❣❡ ✷ P❛❣❡ ✸ ❚❚❳ ✵ ✼ ❡ − ✵✾ ✵ ✼ ❡ − ✵✾ ✵ ✼ ❡ − ✵✾ ❉✶✵✵♣❆ st❡❛❞② r❛t❡ ✷✺ . ✾✾✽✷ ✶✾ . ✻✵✺✻ ✷✾ . ✾✻✼✸ ✷✷ . ✼✻✷✽ ✷✸ . ✽✹✹✸ ✷✵ . ✾✼✹✹ ❘♦✇■♥❞❡① ✶ ✷ ✸ ✹ ✺ ✻ ❈✳ ●ü♥❛② ✭❊♠♦r② ❯♥✐✈❡rs✐t②✮ ◆❡✉r❛❧ ❞❛t❛❜❛s❡ ❛♥❛❧②s✐s ✇✐t❤ P❛♥❞♦r❛ ❖❈◆❈ ✷✵✵✾ ✶✵ ✴ ✶✸
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