O papel dos interneurônios inibitórios do bulbo olfatório no processamento de odores:...

(1)

❯♥✐✈❡rs✐❞❛❞❡ ❞❡ ❙ã♦ P❛✉❧♦

❋❋❈▲❘P ✲ ❉❡♣❛rt❛♠❡♥t♦ ❞❡ ❋ís✐❝❛

❉❊◆■❙❊ ❆❘❘❯❉❆ ❋❆❈❈❍■◆■

❖ ♣❛♣❡❧ ❞♦s ■♥t❡r♥❡✉rô♥✐♦s ■♥✐❜✐tór✐♦s ❞♦ ❇✉❧❜♦

❖❧❢❛tór✐♦ ♥♦ Pr♦❝❡ss❛♠❡♥t♦ ❞❡ ❖❞♦r❡s✿ ❯♠ ❊st✉❞♦

❈♦♠♣✉t❛❝✐♦♥❛❧

(2)

❉❊◆■❙❊ ❆❘❘❯❉❆ ❋❆❈❈❍■◆■

❖ ♣❛♣❡❧ ❞♦s ■♥t❡r♥❡✉rô♥✐♦s ■♥✐❜✐tór✐♦s ❞♦ ❇✉❧❜♦

❖❧❢❛tór✐♦ ♥♦ Pr♦❝❡ss❛♠❡♥t♦ ❞❡ ❖❞♦r❡s✿ ❯♠ ❊st✉❞♦

❈♦♠♣✉t❛❝✐♦♥❛❧

❚❡s❡ ❛♣r❡s❡♥t❛❞❛ à ❋❛❝✉❧❞❛❞❡ ❞❡ ❋✐❧♦s♦✜❛✱

❈✐ê♥❝✐❛s ❡ ▲❡tr❛s ❞❡ ❘✐❜❡✐rã♦ Pr❡t♦ ❞❛

❯♥✐✈❡rs✐❞❛❞❡ ❞❡ ❙ã♦ P❛✉❧♦✱ ❝♦♠♦ ♣❛rt❡ ❞❛s

❡①✐❣ê♥❝✐❛s ♣❛r❛ ❛ ♦❜t❡♥çã♦ ❞♦ tít✉❧♦ ❞❡

❉♦✉t♦r ❡♠ ❈✐ê♥❝✐❛s✳

➪r❡❛ ❞❡ ❈♦♥❝❡♥tr❛çã♦✿

❋ís✐❝❛ ❛♣❧✐❝❛❞❛ à ▼❡❞✐❝✐♥❛ ❡ ❇✐♦❧♦❣✐❛✳

❖r✐❡♥t❛❞♦r✿

Pr♦❢✳ ❉r✳ ❆♥tô♥✐♦ ❈❛r❧♦s ❘♦q✉❡ ❞❛ ❙✐❧✈❛

❋✐❧❤♦✳

❱❡rsã♦ ❝♦rr✐❣✐❞❛

❱❡rsã♦ ♦r✐❣✐♥❛❧ ❞✐s♣♦♥í✈❡❧ ♥❛ ❋❋❈▲❘P✲

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✐✐

❆✉t♦r✐③♦ ❛ r❡♣r♦❞✉çã♦ ❡ ❞✐✈✉❧❣❛çã♦ t♦t❛❧ ♦✉ ♣❛r❝✐❛❧ ❞❡st❡ tr❛❜❛❧❤♦✱ ♣♦r q✉❛❧q✉❡r ♠❡✐♦ ❝♦♥✈❡♥❝✐♦♥❛❧ ♦✉ ❡❧❡trô♥✐❝♦✱ ♣❛r❛ ✜♥s ❞❡ ❡st✉❞♦ ❡ ♣❡sq✉✐s❛✱ ❞❡s❞❡ q✉❡ ❝✐t❛❞❛ ❛ ❢♦♥t❡✳

❋■❈❍❆ ❈❆❚❆▲❖●❘➪❋■❈❆

❆rr✉❞❛ ❋❛❝❝❤✐♥✐✱ ❉❡♥✐s❡

❖ ♣❛♣❡❧ ❞♦s ■♥t❡r♥❡✉rô♥✐♦s ■♥✐❜✐tór✐♦s ❞♦ ❇✉❧❜♦ ❖❧❢❛tór✐♦ ♥♦ Pr♦❝❡ss❛♠❡♥t♦ ❞❡ ❖❞♦r❡s✿ ❯♠ ❊st✉❞♦ ❈♦♠♣✉t❛❝✐♦♥❛❧ ✴ ❉❡♥✐s❡ ❆rr✉❞❛ ❋❛❝❝❤✐♥✐❀ ♦r✐❡♥t❛❞♦r Pr♦❢✳ ❉r✳ ❆♥tô♥✐♦ ❈❛r❧♦s ❘♦q✉❡ ❞❛ ❙✐❧✈❛ ❋✐❧❤♦✳ ❘✐❜❡✐rã♦ Pr❡t♦ ✲ ❙P✱ ✷✵✶✺✳

✶✵✶ ❢✳✿✐❧✳

❚❡s❡ ✭❉♦✉t♦r❛❞♦ ✲ Pr♦❣r❛♠❛ ❞❡ Pós✲❣r❛❞✉❛çã♦ ❡♠ ❋ís✐❝❛ ❛♣❧✐❝❛❞❛ à ▼❡❞✐❝✐♥❛ ❡ ❇✐♦❧♦❣✐❛✮ ✲ ❋❛❝✉❧❞❛❞❡ ❞❡ ❋✐❧♦s♦✜❛✱ ❈✐ê♥❝✐❛s ❡ ▲❡tr❛s ❞❡ ❘✐❜❡✐rã♦ Pr❡t♦ ❞❛ ❯♥✐✈❡rs✐❞❛❞❡ ❞❡ ❙ã♦ P❛✉❧♦✱ ✷✵✶✺✳

✶✳ ◆❡✉r♦❝✐ê♥❝✐❛ ❝♦♠♣✉t❛❝✐♦♥❛❧✳ ✷✳ ▼♦❞❡❧♦s ❜❛s❡❛❞♦s ❡♠

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✐✈

◆♦♠❡✿ ❆rr✉❞❛ ❋❛❝❝❤✐♥✐✱ ❉❡♥✐s❡

❚ít✉❧♦✿ ❖ ♣❛♣❡❧ ❞♦s ■♥t❡r♥❡✉rô♥✐♦s ■♥✐❜✐tór✐♦s ❞♦ ❇✉❧❜♦ ❖❧❢❛tór✐♦ ♥♦ Pr♦❝❡ss❛♠❡♥t♦ ❞❡ ❖❞♦r❡s✿ ❯♠ ❊st✉❞♦ ❈♦♠♣✉t❛❝✐♦♥❛❧

❚❡s❡ ❛♣r❡s❡♥t❛❞❛ à ❋❛❝✉❧❞❛❞❡ ❞❡ ❋✐❧♦s♦✜❛✱

❈✐ê♥❝✐❛s ❡ ▲❡tr❛s ❞❡ ❘✐❜❡✐rã♦ Pr❡t♦ ❞❛

❯♥✐✈❡rs✐❞❛❞❡ ❞❡ ❙ã♦ P❛✉❧♦✱ ❝♦♠♦ ♣❛rt❡ ❞❛s ❡①✐❣ê♥❝✐❛s ♣❛r❛ ❛ ♦❜t❡♥çã♦ ❞♦ tít✉❧♦ ❞❡ ❉♦✉t♦r ❡♠ ❈✐ê♥❝✐❛s✳

❆♣r♦✈❛❞♦ ❡♠✿ ✴ ✴ ✳

❇❛♥❝❛ ❊①❛♠✐♥❛❞♦r❛

Pr♦❢✳ ❉r✳ ✿ ■♥st✐t✉✐çã♦✿

❏✉❧❣❛♠❡♥t♦✿ ❆ss✐♥❛t✉r❛✿

Pr♦❢✳ ❉r✳ ✿ ■♥st✐t✉✐çã♦✿

❏✉❧❣❛♠❡♥t♦✿ ❆ss✐♥❛t✉r❛✿

Pr♦❢✳ ❉r✳ ✿ ■♥st✐t✉✐çã♦✿

❏✉❧❣❛♠❡♥t♦✿ ❆ss✐♥❛t✉r❛✿

Pr♦❢✳ ❉r✳ ✿ ■♥st✐t✉✐çã♦✿

❏✉❧❣❛♠❡♥t♦✿ ❆ss✐♥❛t✉r❛✿

Pr♦❢✳ ❉r✳ ✿ ■♥st✐t✉✐çã♦✿

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✈

❆♦s ♠❡✉s q✉❡r✐❞♦s ♣❛✐s✱

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❆❣r❛❞❡❝✐♠❡♥t♦s

❆♦s ♠❡✉s q✉❡r✐❞♦s ♣❛✐s✱ ❈és❛r ❡ ❩✐❧❞❛✱ ♣❡❧♦ ❛♠♦r ❝♦♠ ♦ q✉❛❧ ♠❡ ❡❞✉❝❛r❛♠✱ ♣♦r t❡r❡♠ ✐♥✈❡st✐❞♦ ❡ ❛❝r❡❞✐t❛❞♦ ♥❛ ♠✐♥❤❛ ❡❞✉❝❛çã♦✱ s❡♠♣r❡ ♠❡ ✐♥❝❡♥t✐✈❛♥❞♦ ♥❛ ❜✉s❝❛ ♣❡❧♦ ❝♦♥❤❡❝✐♠❡♥t♦ ❡ ❛ ♠❡ t♦r♥❛r ✉♠❛ ♣❡ss♦❛ ♠❡❧❤♦r✳

➚ ♠✐♥❤❛ ♠ã❡✱ ♠❡✉ ❡①❡♠♣❧♦ ❞❡ ❞❡t❡r♠✐♥❛çã♦ ❡ ❝♦r❛❣❡♠✱ ❛❣r❛❞❡ç♦ ♣❡❧❛ ♣r♦t❡çã♦ ❡ ♣♦r s❡✉s ❡♥s✐♥❛♠❡♥t♦s✱ ♣♦r s❡✉s ❣❡st♦s s♦❧✐❞ár✐♦s ❡ ♣♦r ❝♦♠♣❛rt✐❧❤❛r ❞❡ ♠✉✐t❛s ❞❛s ♠✐♥❤❛s ♣r❡♦❝✉♣❛çõ❡s ❡ ❝♦♥q✉✐st❛s✳

❆♦ ♠❡✉ ♣❛✐✱ ♣❡❧♦s ❛❜r❛ç♦s ❛♣❡rt❛❞♦s✱ ♣♦r ♠❡ ❡♥s✐♥❛r ❛ ♥ã♦ ❞❡s✐st✐r ❞♦s ♠❡✉s s♦♥❤♦s ❡ ♣♦r ❛❝r❡❞✐t❛r ❡♠ ♠✐♠✳ P❡❧❛ ❧✉t❛ ❝♦♥st❛♥t❡✳ ❊❧❡ é ♠❡✉ ❡①❡♠♣❧♦ ❞❡ ❢♦rç❛ ❡ ♣❡rs❡✈❡r❛♥ç❛✳

❆♦ ♠❡✉ ♦r✐❡♥t❛❞♦r ❆♥tô♥✐♦ ❈❛r❧♦s ❘♦q✉❡ ❞❛ ❙✐❧✈❛ ❋✐❧❤♦✱ ❝♦♠ q✉❡♠ ♠✉✐t♦ ❛♣r❡♥❞✐ t♦❞♦s ❡ss❡s ❛♥♦s✳ ❆❣r❛❞❡ç♦ ♣♦r s✉❛ ♣❛❝✐ê♥❝✐❛ ❝♦♠♦ ❡❞✉❝❛❞♦r✱ s✉❛s ❡①♣❧✐❝❛çõ❡s ❡ s✉❣❡stõ❡s ♥♦ ❞❡❝♦rr❡r ❞♦ ♣r♦❥❡t♦ ❡ ♣❡❧❛ ❝♦rr❡çã♦ ❞❡st❛ t❡s❡✳

➚s ♠✐♥❤❛s ✐r♠ãs✱ ❉❛♥✐❡❧❛ ❡ ❉❡②s❡✱ s✐♠♣❧❡s♠❡♥t❡ ♣♦r ❡st❛r❡♠ s❡♠♣r❡ ❛♦ ♠❡✉ ❧❛❞♦✳ ❆❣r❛❞❡ç♦ ♣❡❧❛ ❝✉♠♣❧✐❝✐❞❛❞❡ ❡ ❛♠✐③❛❞❡ q✉❡ t❡♠♦s✳

➚ ♠✐♥❤❛ ❢❛♠í❧✐❛ q✉❡r✐❞❛✱ ♣❡❧♦ ❝❛r✐♥❤♦ ❡ ❛♣♦✐♦✳ ❊s♣❡❝✐❛❧♠❡♥t❡✱ à ♠✐♥❤❛ ❚✐❛ ❙t❡❧❛✱ ♣♦r s✉❛ ✐♥❝❛♥sá✈❡❧ ❞✐s♣♦s✐çã♦ ❡♠ ❛❥✉❞❛r t♦❞♦s à s✉❛ ✈♦❧t❛ ❡✱ às ♠✐♥❤❛s ♣r✐♠❛s ❡ ❛♠✐❣❛s P❛✉❧❛ ❡ ❘❡♥❛t❛✱ ♣❡❧♦ ❝♦♠♣❛♥❤❡✐r✐s♠♦ ❡ ♣♦r s❡♠♣r❡ t♦r❝❡r❡♠ ♣♦r ♠✐♠✳

❆♦s ♠❡✉s ❛♠✐❣♦s ❡ ❝♦❧❡❣❛s ❞♦ ❧❛❜♦r❛tór✐♦ ❞❡ ❙✐st❡♠❛s ◆❡✉r❛✐s ❈és❛r✱ ❘❛❢❛❡❧✱ ❘♦❞r✐❣♦✱ ❘❡♥❛♥ ❡ ❏♦s✐✱ ♣❡❧❛ tr♦❝❛ ❞❡ ✐❞é✐❛s✱ ♣❡❧❛s ❧♦♥❣❛s ❡①♣❧✐❝❛çõ❡s ❡ ♣❡❧❛ ❤♦r❛ ❞♦ ❝❛❢é✳

❆♦s ❛♠✐❣♦s q✉❡ ✜③ ♥♦ ❞❡❝♦rr❡r ❞❛ ♠✐♥❤❛ ♣ós✲❣r❛❞✉❛çã♦✱ ❏✉❧✐❛♥✱ ▲✉❝❛s ❡ ❉✐♦❣♦✱ ♣❡❧♦ r✐❝♦ ❛♣r❡♥❞✐③❛❞♦ ❡ ♣❡❧❛ ❛t❡♥çã♦ ❡♠ ♣r♦♣♦r s♦❧✉çõ❡s ❡ ✐❞é✐❛s ❡♠ ❞✐❢❡r❡♥t❡s ❢❛s❡s ❞❡st❡ ♣r♦❥❡t♦✳

❆♦s ♣r♦❢❡ss♦r❡s ❘✐❝❛r❞♦ ▲❡ã♦ ❡ ❘❡♥❛t♦ ❚✐♥ós q✉❡ ♣❛rt✐❝✐♣❛r❛♠ ❞❛ ♠✐♥❤❛ ❜❛♥❝❛ ❞❡ q✉❛❧✐✜❝❛çã♦ ❛ss✐♠ ❝♦♠♦ ♦ ♣r♦❢❡ss♦r ❘❡②♥❛❧❞♦ P✐♥t♦ q✉❡ ♣❛rt✐❝✐♣♦✉ ❞❛

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✈✐✐

♠✐♥❤❛ ❞❡❢❡s❛ ❞❡ ♠❡str❛❞♦✳ ❆❣r❛❞❡ç♦ ♣♦r ❝♦♥tr✐❜✉ír❡♠ ❝♦♠ ✐♠♣♦rt❛♥t❡s s✉❣❡stõ❡s ❡ ❝rít✐❝❛s ♣❛r❛ ♦ ❞❡s❡♥✈♦❧✈✐♠❡♥t♦ ♣♦st❡r✐♦r ❡ ❝♦♥❝❧✉sã♦ ❞❡st❡ tr❛❜❛❧❤♦✳

➚s ♠✐♥❤❛s ❛♠✐❣❛s ❞❡ t♦❞❛s ❛s ❤♦r❛s ▼❛r✐❡❧❡✱ ❈❛r❧❛✱ ❉❛♥✐❡❧② ❡ ❆♥❣❡❧❛✱ ♣❡❧❛s ♣❛❧❛✈r❛s ❞❡ s✉♣❡r❛çã♦ ❡ ✐♥❝❡♥t✐✈♦✳

❆♦s ♠❡✉s ❛♠✐❣♦s ❞♦ ✈ô❧❡✐✱ ♣❡❧♦s ♠♦♠❡♥t♦s ❞❡ ❞❡s❝♦♥tr❛çã♦ ❡ ❛❧❡❣r✐❛✳

➚ t♦❞♦s ❛q✉✐ ♠❡♥❝✐♦♥❛❞♦s ❡ ❛♦s ♠❡✉s ❛♠✐❣♦s q✉❡ ♥ã♦ ❢♦r❛♠ ❝✐t❛❞♦s ♠❛s q✉❡ ❡st✐✈❡r❛♠ ♣r❡s❡♥t❡s ❡♠ ❞✐❢❡r❡♥t❡s ❢❛s❡s ❞❡st❡ tr❛❜❛❧❤♦✱ ❛❣r❛❞❡ç♦ ♣❡❧♦s ♠♦♠❡♥t♦s ❞❡ ❞✐✈❡rsã♦✱ ❝♦♥✈❡rs❛s ❡ r✐s❛❞❛s✳

❆♦s ♠❡✉s ❛❧✉♥♦s✱ q✉❡ ♠❡ ❡♥s✐♥❛r❛♠ q✉❡ ♦ ♠❡❧❤♦r ♠♦❞♦ ❞❡ ❛♣r❡♥❞❡r é ❡♥s✐♥❛r✳ ➚ t♦❞♦s ♦s ♣r♦❢❡ss♦r❡s ❡ ❢✉♥❝✐♦♥ár✐♦s ❞♦ ❉❡♣❛rt❛♠❡♥t♦ ❞❡ ❋ís✐❝❛✱ ❡♠ ❡s♣❡❝✐❛❧ à s❡❝r❡t❛r✐❛ ◆✐❧③❛✱ ♣❡❧❛ ❛t❡♥çã♦ ❡ ❞✐s♣♦♥✐❜✐❧✐❞❛❞❡✳

➚ ❈♦♦r❞❡♥❛çã♦ ❞❡ ❆♣❡r❢❡✐ç♦❛♠❡♥t♦ ❞❡ P❡ss♦❛❧ ❞❡ ◆í✈❡❧ ❙✉♣❡r✐♦r ✭❈❆P❊❙✮✱ ♣❡❧♦ s✉♣♦rt❡ ✜♥❛♥❝❡✐r♦✳

❊ ❛❝✐♠❛ ❞❡ t✉❞♦✱ ❛❣r❛❞❡ç♦ ❛ ❉❡✉s ♣♦r ❝♦❧♦❝❛r t♦❞❛s ❡ss❛s ♣❡ss♦❛s ♠❛r❛✈✐❧❤♦s❛s ❡♠ ♠✐♥❤❛ ✈✐❞❛✱ ♣❡❧❛ r❡❛❧✐③❛çã♦ ❞❡st❡ tr❛❜❛❧❤♦ ❡ ♣❡❧❛ ❝♦♥q✉✐st❛ ❞❡ ✉♠ s♦♥❤♦✳

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✈✐✐✐

(10)

❘❡s✉♠♦

❆❘❘❯❉❆ ❋❆❈❈❍■◆■✱ ❉❡♥✐s❡ ❖ ♣❛♣❡❧ ❞♦s ■♥t❡r♥❡✉rô♥✐♦s ■♥✐❜✐tór✐♦s

❞♦ ❇✉❧❜♦ ❖❧❢❛tór✐♦ ♥♦ Pr♦❝❡ss❛♠❡♥t♦ ❞❡ ❖❞♦r❡s✿ ❯♠ ❊st✉❞♦

❈♦♠♣✉t❛❝✐♦♥❛❧✳ ✷✵✶✺✳ ✶✵✶ ❢✳ ❚❡s❡ ✭❉♦✉t♦r❛❞♦ ✲ Pr♦❣r❛♠❛ ❞❡ Pós✲❣r❛❞✉❛çã♦ ❡♠ ❋ís✐❝❛ ❛♣❧✐❝❛❞❛ à ▼❡❞✐❝✐♥❛ ❡ ❇✐♦❧♦❣✐❛✮ ✲ ❋❛❝✉❧❞❛❞❡ ❞❡ ❋✐❧♦s♦✜❛✱ ❈✐ê♥❝✐❛s ❡ ▲❡tr❛s ❞❡ ❘✐❜❡✐rã♦ Pr❡t♦✱ ❯♥✐✈❡rs✐❞❛❞❡ ❞❡ ❙ã♦ P❛✉❧♦✱ ❘✐❜❡✐rã♦ Pr❡t♦ ✲ ❙P✱ ✷✵✶✺✳

❖ ❡♥t❡♥❞✐♠❡♥t♦ ❞♦s ♠❡❝❛♥✐s♠♦s ❞❡ r❡♣r❡s❡♥t❛çã♦ ❡ ♣r♦❝❡ss❛♠❡♥t♦ ❞❡ ♦❞♦r❡s ♣❡❧♦ s✐st❡♠❛ ♦❧❢❛tór✐♦ é ✉♠❛ ❞❛s q✉❡stõ❡s ❝❡♥tr❛✐s ❞❛ ♥❡✉r♦❝✐ê♥❝✐❛ ♠♦❞❡r♥❛✳ ❖s ♦❞♦r❡s sã♦ ❝♦❞✐✜❝❛❞♦s ♣❡❧❛ ❝✐r❝✉✐t❛r✐❛ ✐♥t❡r♥❛ ❞♦ ❜✉❧❜♦ ♦❧❢❛tór✐♦ ❡♠ ♣❛❞rõ❡s ❡s♣❛ç♦✲t❡♠♣♦r❛✐s r❡✢❡t✐❞♦s ♣❡❧❛ ❛t✐✈✐❞❛❞❡ ❞❡ s✉❛s ❝é❧✉❧❛s ❞❡ s❛í❞❛✱ ❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❡ t✉❢♦s❛s✱ q✉❡ tr❛♥s♠✐t❡♠ ♦s r❡s✉❧t❛❞♦s ❞❛s ❝♦♠♣✉t❛çõ❡s ❞❡ss❛ ❡str✉t✉r❛ ✐♥✐❝✐❛❧ ❞❡ ♣r♦❝❡ss❛♠❡♥t♦ ❛ r❡❣✐õ❡s ❝♦rt✐❝❛✐s s✉♣❡r✐♦r❡s✳

❆ ❛rq✉✐t❡t✉r❛ ❞❛s ❝♦♥❡①õ❡s ❡①✐st❡♥t❡s ♥♦ ❜✉❧❜♦ ♦❧❢❛tór✐♦ ❛♣r❡s❡♥t❛ ✐♥✐❜✐çã♦ ❧❛t❡r❛❧ ❡♠ ❞✉❛s ❝❛♠❛❞❛s ❞✐❢❡r❡♥t❡s ❞❡ s✉❛ ❡str✉t✉r❛ ❧❛♠✐♥❛r✱ ✐♥t❡r♠❡❞✐❛❞❛s ♣♦r ❞♦✐s t✐♣♦s ❞✐st✐♥t♦s ❞❡ ✐♥t❡r♥❡✉rô♥✐♦s✳ ◆❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r✱ ♠❛✐s ❡①t❡r♥❛✱ ❛ ✐♥✐❜✐çã♦ ❧❛t❡r❛❧ é ♠❡❞✐❛❞❛ ♣❡❧❛s ❝é❧✉❧❛s ♣❡r✐❣❧♦♠❡r✉❧❛r❡s ❡ ♥❛ ❝❛♠❛❞❛ ♣❧❡①✐❢♦r♠❡ ❡①t❡r♥❛✱ ♠❛✐s ✐♥t❡r♥❛✱ ❛ ✐♥✐❜✐çã♦ ❧❛t❡r❛❧ é ♠❡❞✐❛❞❛ ♣❡❧❛s ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s✳ ❖ ♣❛♣❡❧ ❞❡ss❡s ❞♦✐s ♥í✈❡✐s ❞✐st✐♥t♦s ❞❡ ✐♥✐❜✐çã♦ ❧❛t❡r❛❧ ❡ ♦s ♠❡❝❛♥✐s♠♦s s❡❣✉♥❞♦ ♦s q✉❛✐s ❡❧❡s ❛t✉❛♠ ♠♦❧❞❛♥❞♦ ♦s ♣❛❞rõ❡s ❡s♣❛ç♦✲t❡♠♣♦r❛✐s ❞❡ r❡s♣♦st❛ ❞♦ ❜✉❧❜♦ ♦❧❢❛tór✐♦ ❛ ♦❞♦r❡s ❞✐❢❡r❡♥t❡s sã♦ ❛✐♥❞❛ ♣♦✉❝♦ ❝♦♥❤❡❝✐❞♦s✳ ❖ ♦❜❥❡t✐✈♦ ❞❡st❡ tr❛❜❛❧❤♦ ❢♦✐ ❝♦♥str✉✐r ✉♠ ♠♦❞❡❧♦ ❞❡ r❡❞❡ ♥❡✉r❛❧ ❜✐♦❧♦❣✐❝❛♠❡♥t❡ ♣❧❛✉sí✈❡❧ ❞♦ ❜✉❧❜♦ ♦❧❢❛tór✐♦ ♣❛r❛ ✐♥✈❡st✐❣❛r ❝♦♠♦ ❞♦✐s t✐♣♦s ❞✐❢❡r❡♥t❡s ❞❡ ✐♥t❡r♥❡✉rô♥✐♦s✱ ❛t✉❛♥❞♦ ❡♠ ❡stá❣✐♦s ❞✐st✐♥t♦s ❞❡ ♣r♦❝❡ss❛♠❡♥t♦✱ ♣♦❞❡♠ ❝♦♥tr✐❜✉✐r ♣❛r❛ ❛ ❞✐s❝r✐♠✐♥❛çã♦ ❞❡ ♦❞♦r❡s ❡ ❛ ❝♦♦r❞❡♥❛çã♦ ❞♦s ♣❛❞rõ❡s ❞❡ ❞✐s♣❛r♦ ❞❛s ❝é❧✉❧❛s ♠✐tr❛✐s✳

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①

❖ ♠♦❞❡❧♦ ❞❡ r❡❞❡ ❝♦♥str✉í❞♦✱ ❝♦♠ r❡♣r❡s❡♥t❛çã♦ ❞❡ ♦❞♦r❡s ♣❡❧❛ ❛t✐✈✐❞❛❞❡ ❞❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❡ ❜❛s❡❛❞♦ ♥❛s ✐♥t❡r❛çõ❡s r❡❝í♣r♦❝❛s ❡♥tr❡ ❡ss❛s ❝é❧✉❧❛s ❡ ♦s ✐♥t❡r♥❡✉rô♥✐♦s ✐♥✐❜✐tór✐♦s✱ ♠♦str♦✉ q✉❡ ❛ ✐♥✐❜✐çã♦ ❣❡r❛❞❛ ♣❡❧❛s ❝é❧✉❧❛s ♣❡r✐❣❧♦♠❡r✉❧❛r❡s ♣♦❞❡ ♠❡❧❤♦r❛r ♦ ❝♦♥tr❛st❡ ❡♥tr❡ ♦❞♦r❡s s✐♠✐❧❛r❡s✱ ❢❛❝✐❧✐t❛♥❞♦ ❛ ❞✐s❝r✐♠✐♥❛çã♦ ❞❡ ♦❞♦r❡s✱ ❡♥q✉❛♥t♦ q✉❡ ❛ ✐♥✐❜✐çã♦ ❞❛s ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s ❛t✉❛ ♥♦ r❡✜♥❛♠❡♥t♦ ❞❛ r❡s♣♦st❛ ❞❡ s❛í❞❛ ❞❛ ✐♥❢♦r♠❛çã♦ ♦❧❢❛tór✐❛✳

P❛❧❛✈r❛s✲❝❤❛✈❡✿ ✶✳ ◆❡✉r♦❝✐ê♥❝✐❛ ❝♦♠♣✉t❛❝✐♦♥❛❧✳ ✷✳ ▼♦❞❡❧♦s ❜❛s❡❛❞♦s ❡♠

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❆❜str❛❝t

❆❘❘❯❉❆ ❋❆❈❈❍■◆■✱ ❉❡♥✐s❡ ❚❤❡ r♦❧❡ ♦❢ ✐♥❤✐❜✐t♦r② ✐♥t❡r♥❡✉r♦♥s ♦❢ t❤❡ ❖❧❢❛❝t♦r② ❇✉❧❜ ♦♥ ❖❞♦r Pr♦❝❡ss✐♥❣✿ ❆ ❈♦♠♣✉t❛t✐♦♥❛❧ ❙t✉❞②✳ ✷✵✶✺✳ ✶✵✶ ❢✳ ❚❤❡s✐s ✭P❤✳❉✳ ✲ P♦st❣r❛❞✉❛t❡ ♣r♦❣r❛♠ ✐♥ P❤②s✐❝s ❛♣♣❧✐❡❞ t♦ ▼❡❞✐❝✐♥❡ ❛♥❞ ❇✐♦❧♦❣②✮ ✲ ❋❛❝✉❧t② ♦❢ P❤✐❧♦s♦♣❤②✱ ❙❝✐❡♥❝❡s ❛♥❞ ▲✐t❡r❛t✉r❡✱ ❯♥✐✈❡rs✐t② ♦❢ ❙ã♦ P❛✉❧♦✱ ❘✐❜❡✐rã♦ Pr❡t♦ ✲ ❙P✱ ✷✵✶✺✳

❚❤❡ ✉♥❞❡rst❛♥❞✐♥❣ ♦❢ ♦❞♦r r❡♣r❡s❡♥t❛t✐♦♥ ❛♥❞ ♣r♦❝❡ss✐♥❣ ♠❡❝❤❛♥✐s♠s ❜② t❤❡ ♦❧❢❛❝t♦r② s②st❡♠ ✐s ♦♥❡ ♦❢ t❤❡ ❝❡♥tr❛❧ q✉❡st✐♦♥s ♦❢ ♠♦❞❡r♥ ♥❡✉r♦s❝✐❡♥❝❡✳ ❖❞♦rs ❛r❡ ❡♥❝♦❞❡❞ ❜② t❤❡ ♦❧❢❛❝t♦r② ❜✉❧❜ ❝✐r❝✉✐tr② ✐♥ t❡r♠s ♦❢ s♣❛t✐♦t❡♠♣♦r❛❧ s♣✐❦✐♥❣ ♣❛tt❡r♥s✳ ❚❤❡s❡ ❛r❡ r❡✢❡❝t❡❞ ✐♥ t❤❡ ❛❝t✐✈✐t② ♦❢ t❤❡ ♠✐tr❛❧ ❝❡❧❧s✱ ✇❤✐❝❤ ❛r❡ t❤❡ ♦✉t♣✉t ❝❡❧❧s ♦❢ t❤❡ ♦❧❢❛❝t♦r② ❜✉❧❜ t❤❛t tr❛♥s♠✐t t❤❡ ✐♥❢♦r♠❛t✐♦♥ ♣r♦❝❡ss❡❞ ✐♥ t❤✐s ❡❛r❧② str✉❝t✉r❡ t♦ ❤✐❣❤❡r ❝♦rt✐❝❛❧ r❡❣✐♦♥s✳

❚❤❡ ❛r❝❤✐t❡❝t✉r❡ ♦❢ t❤❡ ♦❧❢❛❝t♦r② ❜✉❧❜ ❝♦♥♥❡❝t✐♦♥s ♣r❡s❡♥ts ❧❛t❡r❛❧ ✐♥❤✐❜✐t✐♦♥ ❛t t✇♦ ❞✐✛❡r❡♥t ❧❛②❡rs ♦❢ ✐ts ❧❛♠✐♥❛r str✉❝t✉r❡✱ ♠❡❞✐❛t❡❞ ❜② t✇♦ ❞✐st✐♥❝t t②♣❡s

♦❢ ✐♥t❡r♥❡✉r♦♥s✳ ■♥ t❤❡ ❣❧♦♠❡r✉❧❛r ❧❛②❡r✱ ❧❛t❡r❛❧ ✐♥❤✐❜✐t✐♦♥ ✐s ♠❡❞✐❛t❡❞ ❜②

♣❡r✐❣❧♦♠❡r✉❧❛r ❝❡❧❧s✳ ■♥ t❤❡ ❡①t❡r♥❛❧ ♣❧❡①✐❢♦r♠ ❧❛②❡r✱ ❧❛t❡r❛❧ ✐♥❤✐❜✐t✐♦♥ ✐s ♠❡❞✐❛t❡❞ ❜② ❣r❛♥✉❧❡ ❝❡❧❧s✳ ❚❤❡ r♦❧❡ ♦❢ t❤❡s❡ t✇♦ ❞✐✛❡r❡♥t ❧❛t❡r❛❧ ✐♥❤✐❜✐t✐♦♥ ❧❡✈❡❧s ❛♥❞ t❤❡ ♠❡❝❤❛♥✐s♠s ✇❤❡r❡❜② t❤❡② s❤❛♣❡ t❤❡ s♣❛t✐❛❧ ❛♥❞ t❡♠♣♦r❛❧ ♣❛tt❡r♥s ♦❢ t❤❡ ♦❧❢❛❝t♦r② ❜✉❧❜ r❡s♣♦♥s❡ t♦ ❞✐✛❡r❡♥t ♦❞♦rs ✐s ♥♦t ✇❡❧❧ ❦♥♦✇♥✳ ❚❤❡ ❛✐♠ ♦❢ t❤✐s ✇♦r❦ ✇❛s t♦ ❜✉✐❧❞ ❛ ❜✐♦❧♦❣✐❝❛❧❧② ♣❧❛✉s✐❜❧❡ ♥❡✉r❛❧ ♥❡t✇♦r❦ ♠♦❞❡❧ ♦❢ t❤❡ ♦❧❢❛❝t♦r② ❜✉❧❜ t♦ ✐♥✈❡st✐❣❛t❡ ❤♦✇ t✇♦ ❞✐✛❡r❡♥t t②♣❡s ♦❢ ✐♥t❡r♥❡✉r♦♥s✱ ❛❝t✐♥❣ ❛t ❞✐✛❡r❡♥t ♣r♦❝❡ss✐♥❣ st❛❣❡s✱ ❝♦✉❧❞ ❝♦♥tr✐❜✉t❡ t♦ ♦❞♦r ❞✐s❝r✐♠✐♥❛t✐♦♥ ❛♥❞ t❤❡ ❝♦♦r❞✐♥❛t✐♦♥ ♦❢ t❤❡ ♠✐tr❛❧ ❝❡❧❧s s♣✐❦✐♥❣ ♣❛tt❡r♥s✳

❚❤❡ r❡s✉❧ts ♦❢ s✐♠✉❧❛t✐♦♥s ♦❢ t❤❡ ♥❡t✇♦r❦ ♠♦❞❡❧ s❤♦✇♥ t❤❛t t❤❡ ✐♥❤✐❜✐t✐♦♥

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①✐✐

❣❡♥❡r❛t❡❞ ❜② ♣❡r✐❣❧♦♠❡r✉❧❛r ❝❡❧❧s ❝❛♥ ♣r♦✈✐❞❡ ❝♦♥tr❛st ❡♥❤❛♥❝❡♠❡♥t ❛♥❞ ♦❞♦rs ❞✐s❝r✐♠✐♥❛t✐♦♥✱ ✇❤✐❧❡ t❤❡ ❣r❛♥✉❧❡ ❝❡❧❧ ✐♥❤✐❜✐t✐♦♥ ❝❛♥ r❡✜♥❡ t❤❡ ♦✉t♣✉t r❡s♣♦♥s❡ ♦❢ t❤❡ ♦❧❢❛❝t♦r② ✐♥❢♦r♠❛t✐♦♥✳

❑❡②✲✇♦r❞s✿ ✶✳ ❈♦♠♣✉t❛t✐♦♥❛❧ ◆❡✉r♦s❝✐❡♥❝❡✳ ✷✳ ❈♦♥❞✉❝t❛♥❝❡ ❜❛s❡❞ ♠♦❞❡❧✳

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▲✐st❛ ❞❡ ❋✐❣✉r❛s

✷✳✶ ❉✐✈❡rs✐❞❛❞❡ ♠♦r❢♦❧ó❣✐❝❛ ❞♦s ♥❡✉rô♥✐♦s✳ ❆✲ ❈é❧✉❧❛ ♣✐r❛♠✐❞❛❧ ❞♦ ❝órt❡① ❝❡r❡❜r❛❧✳ ❇✲ ❈é❧✉❧❛ ❞❡ P✉r❦✐♥❥❡ ❞♦ ❝❡r❡❜❡❧♦✳ ❈✲ ❈é❧✉❧❛

❡str❡❧❛❞❛ ❞♦ ❝órt❡① ❝❡r❡❜r❛❧✳ ✭❋✐❣✉r❛s ❛❞❛♣t❛❞❛s ❛ ♣❛rt✐r ❞❡

❞❡s❡♥❤♦s ♦r✐❣✐♥❛✐s ❞❡ ❘❛♠ó♥ ② ❈❛❥❛❧ ❞✐s♣♦♥í✈❡✐s ♥❛ ✐♥t❡r♥❡t ❡♠ ❤tt♣✿✴✴❝✈❝✳❝❡r✈❛♥t❡s✳❡s✴❝✐❡♥❝✐❛✴❝❛❥❛❧✴❞❡❢❛✉❧t✳❤t♠✮ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✼ ✷✳✷ ❈✐r❝✉✐t♦ ❡❧étr✐❝♦ ❡q✉✐✈❛❧❡♥t❡ ♣❛r❛ ✉♠❛ ♠❡♠❜r❛♥❛ ♥❡✉r♦♥❛❧ ❝♦♠

❝♦♥❞✉tâ♥❝✐❛ ♣❛ss✐✈❛✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✾ ✷✳✸ ❈✐r❝✉✐t♦ ❡❧étr✐❝♦ ❡q✉✐✈❛❧❡♥t❡ ❞♦ ♠♦❞❡❧♦ ❞❡ ❍♦❞❣❦✐♥✲❍✉①❧❡②✳ ✳ ✳ ✳ ✳ ✳ ✳ ✶✶

✷✳✹ ❚rês ❢✉♥çõ❡s ❞❡ ♦♥❞❛ ♣❛r❛ ❝♦♥❞✉tâ♥❝✐❛ s✐♥á♣t✐❝❛✿ ✭❛✮ ú♥✐❝♦

❞❡❝❛✐♠❡♥t♦ ❡①♣♦♥❡♥❝✐❛❧ ❝♦♠ τ = 3 ♠s❀ ✭❜✮ ❢✉♥çã♦ ❛❧❢❛ ❝♦♠ τ = 1

♠s❀ ❡ ✭❝✮ ❞✉♣❧❛ ❡①♣♦♥❡♥❝✐❛❧ ❝♦♠ τ1 = 3 ♠s ❡ τ2 = 1 ♠s✳ ❘❡s♣♦st❛

❛ ✉♠ ú♥✐❝♦ ♣♦t❡♥❝✐❛❧ ❞❡ ❛çã♦ ♣ré✲s✐♥á♣t✐❝♦ q✉❡ ❝❤❡❣❛ ♥♦ t❡♠♣♦ ❂ ✶ ♠s✳ ❚♦❞♦s ❛s ❝♦♥❞✉tâ♥❝✐❛s sã♦ ❞✐♠❡♥s✐♦♥❛❞❛s ♣❛r❛ ✉♠ ♠á①✐♠♦ ❞❡ ✶ ✭✉♥✐❞❛❞❡s ❛r❜✐trár✐❛s✮ ❬❙t❡rr❛tt ❡t ❛❧✳ ✷✵✶✶❪✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✶✻

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①✐✈

✷✳✺ ❊sq✉❡♠❛ ❞♦ ♣❛❞rã♦ ❞❡ ❝♦♥❡①õ❡s s✐♥á♣t✐❝❛s ♥♦ ❜✉❧❜♦ ♦❧❢❛tór✐♦✳ ❖s ❞❡♥❞r✐t♦s ♣r✐♠ár✐♦s ❞❛s ❝é❧✉❧❛s ♠✐tr❛✐s ✭▼✮ ❡ t✉❢♦s❛s ✭❚✮ r❡❝❡❜❡♠ s✐♥❛♣s❡s ❡①❝✐t❛tór✐❛s ✭✐♥❞✐❝❛❞❛s ♣♦r s❡t❛s ❜r❛♥❝❛s✮ ❞♦s ❛①ô♥✐♦s ❞♦s ♥❡✉rô♥✐♦s r❡❝❡♣t♦r❡s ♦❧❢❛tór✐♦s ✭◆❘❖s✮ ♥♦s ❣❧♦♠ér✉❧♦s ✭●▲✮✳ ◆❛ ✜❣✉r❛ ❡stã♦ r❡♣r❡s❡♥t❛❞♦s ❞♦✐s t✐♣♦s ❞❡ ✐♥t❡r♥❡✉rô♥✐♦s q✉❡ ❢❛③❡♠ s✐♥❛♣s❡s ❞❡♥❞r♦❞❡♥❞rít✐❝❛s ✐♥✐❜✐tór✐❛s ✭s❡t❛s ❡s❝✉r❛s✮ ♥❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❡ t✉❢♦s❛s ✭♠✴t✮✱ ❛s ❝é❧✉❧❛s ♣❡r✐❣❧♦♠❡r✉❧❛r❡s ✭P●✮ ❡ ❛s ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s ✭●r✮✳ ❆s ♣r✐♠❡✐r❛s ❢❛③❡♠ s✐♥❛♣s❡s ❝♦♠ ♦s ❞❡♥❞r✐t♦s ♣r✐♠ár✐♦s ❞❛s ❝é❧✉❧❛s ♠✴t ❡ ❛s s❡❣✉♥❞❛s ❢❛③❡♠ s✐♥❛♣s❡s ❝♦♠ ♦s ❞❡♥❞r✐t♦s s❡❝✉♥❞ár✐♦s ❞❡ss❛s ❝é❧✉❧❛s✱ q✉❡ s❡ ❡st❡♥❞❡♠ ❤♦r✐③♦♥t❛❧♠❡♥t❡ ♣♦r ❧♦♥❣❛s ❞✐stâ♥❝✐❛s✳ ❆s ❝é❧✉❧❛s ❞❡ s❛í❞❛ ❞♦ ❜✉❧❜♦ sã♦ ❛s ❝é❧✉❧❛s ▼ ❡ ❚ ✭✜❣✉r❛ ❛❞❛♣t❛❞❛ ❞❛ ❋✐❣✉r❛ ✶ ❞❡ ▼♦r✐ ❡t ❛❧ ❬▼♦r✐ ❡ ❨♦s❤✐❤❛r❛ ✶✾✾✾❪ ✶✽

✸✳✶ ❈♦♠♣❛r❛çã♦ ❡♥tr❡ ♦ ♥❡✉rô♥✐♦ r❡❝❡♣t♦r ♦❧❢❛tór✐♦ ✭◆❘❖✮ r❡❛❧ ❬❩✉❢❛❧❧ ❡ ▲❡✐♥❞❡r✲❩✉❢❛❧❧ ✷✵✵✵❪ ✭❡sq✉❡r❞❛✮ ❡ ♦ ♠♦❞❡❧♦ ❝♦♠♣❛rt✐♠❡♥t❛❧ ✭❞✐r❡✐t❛✮✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✸ ✸✳✷ ❊sq✉❡♠❛ ❞❛s ❝♦♥❡①õ❡s ❡♥tr❡ ♦s ♠♦❞❡❧♦s ❞❡ ♥❡✉rô♥✐♦ r❡❝❡♣t♦r ♦❧❢❛tór✐♦

✭◆❘❖✮✱ ❝é❧✉❧❛ ♣❡r✐❣❧♦♠❡r✉❧❛r ✭P●✮✱ ❝é❧✉❧❛ ♠✐tr❛❧ ✭▼✮ ❡ ❝é❧✉❧❛ ❣r❛♥✉❧❛r ✭●r✮✳ ❆s ✢❡❝❤❛s ❛③✉✐s ❡ ✈❡r♠❡❧❤❛s ✐♥❞✐❝❛♠✱ r❡s♣❡❝t✐✈❛♠❡♥t❡✱ s✐♥❛♣s❡s ❡①❝✐t❛tór✐❛s ❡ ✐♥✐❜✐tór✐❛s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✾

✸✳✸ ❈♦♥❡①õ❡s ♥❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r✿◆❡✉rô♥✐♦s r❡❝❡♣t♦r❡s ♦❧❢❛tór✐♦s ✭♣♦♥t♦s

✈❡r❞❡s✮ ❢❛③❡♠ s✐♥❛♣s❡s ❡①❝✐t❛tór✐❛s ❝♦♠ ❝é❧✉❧❛s P●s ✭♣♦♥t♦s ✈❡r♠❡❧❤♦s✮

❡ ❝é❧✉❧❛s ♠✐tr❛✐s ✭♣♦♥t♦s ♣r❡t♦s✮✳ ❈é❧✉❧❛s ♠✐tr❛✐s ❡①❝✐t❛♠ ❝é❧✉❧❛s

P●s ❡ r❡❝❡❜❡♠ ✐♥✐❜✐çõ❡s✱ ❢♦r♠❛♥❞♦ s✐♥❛♣s❡s r❡❝í♣r♦❝❛s✳ ❆s ❝♦♥❡①õ❡s ✐♥t❡r❣❧♦♠❡r✉❧❛r❡s ❡♥tr❡ ♦s ❣❧♦♠ér✉❧♦s ✶ ❡ ✸ ♥ã♦ ❡stã♦ ♠♦str❛❞❛s✱ ♠❛s ❡❧❛s

❡①✐st❡♠✱ ❞❡ ♠❛♥❡✐r❛ q✉❡ ♦ ❝♦♥❥✉♥t♦ ❢♦r♠❛ ✉♠❛ ❡str✉t✉r❛ ❝✐r❝✉❧❛r✳ ✳ ✳ ✳ ✳ ✳ ✸✷

✸✳✹ ❈♦♥❡①õ❡s ♥❛ ❝❛♠❛❞❛ ❣r❛♥✉❧❛r✿ ◆❡✉rô♥✐♦s r❡❝❡♣t♦r❡s ♦❧❢❛tór✐♦s ✭♣♦♥t♦s

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①✈

✸✳✺ ❈♦♥❡①õ❡s ♥♦ ♠♦❞❡❧♦ ❞♦ ❜✉❧❜♦ ♦❧❢❛tór✐♦ ♣❛r❛ ♦ ❡st✉❞♦ ❞♦ ❝❛s♦ ❝♦♠ ❛s

❝❛♠❛❞❛s ❣❧♦♠❡r✉❧❛r ❡ ❣r❛♥✉❧❛r ❥✉♥t❛s✿ ◆❡✉rô♥✐♦s r❡❝❡♣t♦r❡s ♦❧❢❛tór✐♦s ✭♣♦♥t♦s ✈❡r❞❡s✮ ❢❛③❡♠ s✐♥❛♣s❡s ❡①❝✐t❛tór✐❛s ❝♦♠ ❝é❧✉❧❛s P●s ✭♣♦♥t♦s ✈❡r♠❡❧❤♦s✮ ❡ ❝é❧✉❧❛s ♠✐tr❛✐s ✭♣♦♥t♦s ♣r❡t♦s✮✳ ❈é❧✉❧❛s ♠✐tr❛✐s ❡①❝✐t❛♠ ❝é❧✉❧❛s P●s ✭♣♦♥t♦s ✈❡r♠❡❧❤♦s✮ ❡ ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s ✭♣♦♥t♦s ❛③✉✐s✮ ❡ r❡❝❡❜❡♠ ✐♥✐❜✐çõ❡s ❞❡st❛s✱ ❢♦r♠❛♥❞♦ s✐♥❛♣s❡s r❡❝í♣r♦❝❛s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✼

✹✳✶ ❘❡s♣♦st❛s ❞♦s ♠♦❞❡❧♦s ❞❛s ❝é❧✉❧❛s ✐♥❞✐✈✐❞✉❛✐s✳ ❆✿ P♦t❡♥❝✐❛❧ ❞❡ ♠❡♠❜r❛♥❛

❞♦ ♠♦❞❡❧♦ ❞❛ ❝é❧✉❧❛ ♠✐tr❛❧ ♣❛r❛ ✉♠❛ ❝♦rr❡♥t❡ ❞❡s♣♦❧❛r✐③❛♥t❡ ❞❡ 0_.2738

♥❆ ❛♣❧✐❝❛❞❛ ♣♦r 450 ♠s ❝♦♠ ✐♥í❝✐♦ ❡♠ _t = 50 ♠s✳ ❇✿ P♦t❡♥❝✐❛❧ ❞❡

♠❡♠❜r❛♥❛ ❞♦ ♠♦❞❡❧♦ ❞❛ ❝é❧✉❧❛ P● ♣❛r❛ ✉♠❛ ❝♦rr❡♥t❡ ❞❡s♣♦❧❛r✐③❛♥t❡

❞❡ ✵✳✶ ♥❆ ❛♣❧✐❝❛❞❛ ♣♦r ✻✵✵ ♠s ❝♦♠ ✐♥í❝✐♦ ❡♠ t = 200 ♠s✳ ❈✿

P♦t❡♥❝✐❛❧ ❞❡ ♠❡♠❜r❛♥❛ ❞♦ ♠♦❞❡❧♦ ❞❛ ❝é❧✉❧❛ ❣r❛♥✉❧❛r ♣❛r❛ ✉♠❛ ❝♦rr❡♥t❡ ❞❡s♣♦❧❛r✐③❛♥t❡ ❞❡ 6_.25·10−3

♥❆ ❛♣❧✐❝❛❞❛ ♣♦r 2500 ♠s ❡ ✐♥í❝✐♦ ❡♠ _t = 0

♠s✳ Pr♦t♦❝♦❧♦s ❡st❛❜❡❧❡❝✐❞♦s ❞❡ ❛❝♦r❞♦ ❝♦♠ ♦s ❞❛❞♦s ❞♦s ♠♦❞❡❧♦s ♦r✐❣✐♥❛✐s

❬❉❛✈✐s♦♥ ✷✵✵✶✱ ❉❛✈✐s♦♥✱ ❋❡♥❣ ❡ ❇r♦✇♥ ✷✵✵✸✱ ❆rr✉❞❛ ✷✵✶✵❪ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✹✵

✹✳✷ ❋r❡q✉ê♥❝✐❛ ❞❡ ❞✐s♣❛r♦s ❞♦ ◆❘❖ ❡♠ ❢✉♥çã♦ ❞❛ ❝♦rr❡♥t❡ ✐♥❥❡t❛❞❛✳ ❘❡s♣♦st❛ ❞♦ ♠♦❞❡❧♦ ✭✈❡r♠❡❧❤♦✮ ❝♦♠♣❛r❛❞❛ ❝♦♠ ❞❛❞♦s ❡①♣❡r✐♠❡♥t❛✐s ❞❡ r❛t♦s ✭♣r❡t♦✮ ❡ ❝❛♠✉♥❞♦♥❣♦s ✭❛③✉❧✮ ❬▼❛✱ ❈❤❡♥ ❡ ❙❤❡♣❤❡r❞ ✶✾✾✾❪ ✳ ✹✶ ✹✳✸ P♦t❡♥❝✐❛❧ ❞❡ ♠❡♠❜r❛♥❛ ❞♦ ♠♦❞❡❧♦ ❞♦ ◆❘❖ ✭❛❜❛✐①♦✮ ♣❛r❛ ✉♠❛ ✐♥❥❡çã♦

❞❡ ❝♦rr❡♥t❡ ♥✉❧❛ ✭❛❝✐♠❛✮✳ ❖ ♠♦❞❡❧♦ ❡①✐❜❡ ❞✐s♣❛r♦s ❡s♣♦♥tâ♥❡♦s✳ ✳ ✳ ✳ ✹✶ ✹✳✹ P♦t❡♥❝✐❛❧ ❞❡ ♠❡♠❜r❛♥❛ ❞♦ ♠♦❞❡❧♦ ❞♦ ◆❘❖ ✭❛❜❛✐①♦✮ ♣❛r❛ ❛ ❝♦rr❡♥t❡

r❡s♣✐r❛tór✐❛ ♣r♦♣♦r❝✐♦♥❛❧ ❛ ❝♦♥❝❡♥tr❛çã♦ ❞❡ ♦❞♦r ✭❛❝✐♠❛✮✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✹✷ ✹✳✺ ❋r❡q✉ê♥❝✐❛ ❞❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❞♦s ❣❧♦♠ér✉❧♦ ✶✱ ✷ ❡ ✸ ❡♠ r❡s♣♦st❛ à ❝✐♥❝♦

♦❞♦r❡s ❞✐❢❡r❡♥t❡s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✹✸

✹✳✻ ❘❡s♣♦st❛ ❞❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✶✿ ❋r❡q✉ê♥❝✐❛ ❞❡ ❞✐s♣❛r♦s

❞❛s ❞✉❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❞❡ ❝❛❞❛ ❣❧♦♠ér✉❧♦ ❡♠ ❢✉♥çã♦ ❞❛ ❝♦♥❞✉tâ♥❝✐❛ ♠á①✐♠❛ ✭gP G✮✱ ❡♠ ♥❙✱ ❞❛s s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✹✹

✹✳✼ ❘❡s♣♦st❛ ❞❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✷ ♣❛r❛ p= 1✿ ❋r❡q✉ê♥❝✐❛

(17)

①✈✐

✹✳✽ ❘❡s♣♦st❛ ❞❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✷ ♣❛r❛ p <1✿ ❋r❡q✉ê♥❝✐❛ ❞❡ ❞✐s♣❛r♦s ❞❛s ❞✉❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❞❡ ❝❛❞❛ ❣❧♦♠ér✉❧♦ ❡♠ ❢✉♥çã♦ ❞❛ ❝♦♥❞✉tâ♥❝✐❛ ♠á①✐♠❛ ✭gP G✮✱ ❡♠ ♥❙✱ ❞❛s s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✹✻

✹✳✾ ❘❡s♣♦st❛ ❞❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✷ ♣❛r❛ p >1✿ ❋r❡q✉ê♥❝✐❛ ❞❡ ❞✐s♣❛r♦s ❞❛s ❞✉❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❞❡ ❝❛❞❛ ❣❧♦♠ér✉❧♦ ❡♠ ❢✉♥çã♦ ❞❛ ❝♦♥❞✉tâ♥❝✐❛ ♠á①✐♠❛ ✭gP G✮✱ ❡♠ ♥❙✱ ❞❛s s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✹✻

✹✳✶✵ ❘❡s♣♦st❛ ❞❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✷ ♣❛r❛ p = 1 ❡ _τ₂ = 18

♠s✿ ❋r❡q✉ê♥❝✐❛ ❞❡ ❞✐s♣❛r♦s ❞❛s ❞✉❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❞❡ ❝❛❞❛ ❣❧♦♠ér✉❧♦ ❡♠ ❢✉♥çã♦ ❞❛ ❝♦♥❞✉tâ♥❝✐❛ ♠á①✐♠❛ ✭gP G✮✱ ❡♠ ♥❙✱ ❞❛s s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s✳✳ ✳ ✳ ✹✼

✹✳✶✶ ❘❡s♣♦st❛ ❞❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✷ ♣❛r❛ p = 1 ❡ _τ₂ = 50

♠s✿ ❋r❡q✉ê♥❝✐❛ ❞❡ ❞✐s♣❛r♦s ❞❛s ❞✉❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❞❡ ❝❛❞❛ ❣❧♦♠ér✉❧♦ ❡♠ ❢✉♥çã♦ ❞❛ ❝♦♥❞✉tâ♥❝✐❛ ♠á①✐♠❛ ✭gP G✮✱ ❡♠ ♥❙✱ ❞❛s s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s✳✳ ✳ ✳ ✹✽

✹✳✶✷ ❘❡s♣♦st❛ ❞❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✷ ♣❛r❛ p = 1 ❡ _τ₂ = 200

♠s✿ ❋r❡q✉ê♥❝✐❛ ❞❡ ❞✐s♣❛r♦s ❞❛s ❞✉❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❞❡ ❝❛❞❛ ❣❧♦♠ér✉❧♦ ❡♠ ❢✉♥çã♦ ❞❛ ❝♦♥❞✉tâ♥❝✐❛ ♠á①✐♠❛ ✭gP G✮✱ ❡♠ ♥❙✱ ❞❛s s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s✳✳ ✳ ✳ ✹✾

✹✳✶✸ ▼❡❧❤♦r❛♠❡♥t♦ ❞♦ ❝♦♥tr❛st❡ ♥❛ r❡♣r❡s❡♥t❛çã♦ ❣❧♦♠❡r✉❧❛r ❞♦s ♦❞♦r❡s✿ ❈❛❞❛

❣rá✜❝♦ ♠♦str❛ ❛ r❛③ã♦ ❡♥tr❡ ❛ r❡s♣♦st❛ ❞♦ s❡❣✉♥❞♦ ❣❧♦♠ér✉❧♦ ♠❛✐s ❛t✐✈❛❞♦ ♣♦r ✉♠ ♦❞♦r ❡ ♦ ♣r✐♠❡✐r♦ ❣❧♦♠ér✉❧♦ ♠❛✐s ❛t✐✈❛❞♦ ♣♦r ❡❧❡ ✭s❡✉ ❣❧♦♠ér✉❧♦ ♣r❡❢❡r❡♥❝✐❛❧✮ ♣❛r❛ q✉❛tr♦ ✈❛❧♦r❡s ❞✐❢❡r❡♥t❡s ❞♦ t❡♠♣♦ ❞❡ ❞❡❝❛✐♠❡♥t♦ ❞❛s s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s ❢❡✐t❛s ♣❡❧❛s ❝é❧✉❧❛s P●s✳ ❆s r❡s♣♦st❛s ❞♦s ❣❧♦♠ér✉❧♦s ❛♦s ♦❞♦r❡s sã♦ q✉❛♥t✐✜❝❛❞❛s ♣❡❧❛ ❢r❡q✉ê♥❝✐❛ t♦t❛❧ ❞❡ ❞✐s♣❛r♦s ❞❡ s✉❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❛♦ ♦❞♦r✱ ✐st♦ é✱ ❋❳ ✐♥❞✐❝❛ ❛ ❢r❡q✉ê♥❝✐❛ ❞❡ ❞✐s♣❛r♦s ❞♦ ❣❧♦♠ér✉❧♦ ❳ ❛♦ ♦❞♦r✳ ❆s r❛③õ❡s sã♦ ♠♦str❛❞❛s ❡♠ ❢✉♥çã♦ ❞♦ ✈❛❧♦r ❞❛ ❝♦♥❞✉tâ♥❝✐❛ ♠á①✐♠❛ ✭gP G✮ ❞❛ s✐♥❛♣s❡ ✐♥✐❜✐tór✐❛✳✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✵

✹✳✶✹ ❘❡s♣♦st❛s ❞❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✸✿ ❋r❡q✉ê♥❝✐❛s ♠é❞✐❛s

❞♦s ❞✐s♣❛r♦s ❞❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❞♦ ❣❧♦♠ér✉❧♦✱ ❝❛❧❝✉❧❛❞❛s ♣♦r ✶✵ r❡♣❡t✐çõ❡s ❞❛s s✐♠✉❧❛çõ❡s✱ ♣❛r❛ ♦s ✈❛❧♦r❡s ❞❡ ❝♦♥❞✉tâ♥❝✐❛ s✐♥á♣t✐❝❛ ♠á①✐♠❛gP G ❞❛s

(18)

①✈✐✐

✹✳✶✺ ❘❡s♣♦st❛ ❞❛ ❝❛♠❛❞❛ ❣r❛♥✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✶✿ ❋r❡q✉ê♥❝✐❛ ❞❡ ❞✐s♣❛r♦s

❞❛s ❞✉❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❞❡ ❝❛❞❛ ❣❧♦♠ér✉❧♦ ❡♠ ❢✉♥çã♦ ❞❛ ❝♦♥❞✉tâ♥❝✐❛ ♠á①✐♠❛ ✭gGr✮✱ ❡♠ ♥❙✱ ❞❛s s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s ❢❡✐t❛s ♣❡❧❛s ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s✳ ✺✸

✹✳✶✻ ❘❡s♣♦st❛ ❞❛ ❝❛♠❛❞❛ ❣r❛♥✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✶ ❝♦♠ ❛t✐✈❛çã♦ ❞❛ ❝❛♠❛❞❛

♣♦r ❝♦rr❡♥t❡ ❞❡ ✵✱✵✻✷✺ ♥❆✿ ❋r❡q✉ê♥❝✐❛ ❞❡ ❞✐s♣❛r♦s ❞❛s ❞✉❛s ❝é❧✉❧❛s ♠✐tr❛✐s

❞❡ ❝❛❞❛ ❣❧♦♠ér✉❧♦ ❡♠ ❢✉♥çã♦ ❞❛ ❝♦♥❞✉tâ♥❝✐❛ ♠á①✐♠❛ ✭gGr✮✱ ❡♠ ♥❙✱ ❞❛s

s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s ❢❡✐t❛s ♣❡❧❛s ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✹

✹✳✶✼ ❘❡s♣♦st❛ ❞❛ ❝❛♠❛❞❛ ❣r❛♥✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✶ ❝♦♠ ❛t✐✈❛çã♦ ❞❛ ❝❛♠❛❞❛

♣♦r ❝♦rr❡♥t❡ ❞❡ ✵✱✻✷✺ ♥❆✿ ❋r❡q✉ê♥❝✐❛ ❞❡ ❞✐s♣❛r♦s ❞❛s ❞✉❛s ❝é❧✉❧❛s ♠✐tr❛✐s

❞❡ ❝❛❞❛ ❣❧♦♠ér✉❧♦ ❡♠ ❢✉♥çã♦ ❞❛ ❝♦♥❞✉tâ♥❝✐❛ ♠á①✐♠❛ ✭gGr✮✱ ❡♠ ♥❙✱ ❞❛s

s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s ❢❡✐t❛s ♣❡❧❛s ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✺

✹✳✶✽ ❘❡s♣♦st❛s ❞❛ ❝❛♠❛❞❛ ❣r❛♥✉❧❛r ♥❛ ❝♦♥✜❣✉r❛çã♦ ✷✿ ❋r❡q✉ê♥❝✐❛s ♠é❞✐❛s ❞♦s

❞✐s♣❛r♦s ❞❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❞♦ ❣❧♦♠ér✉❧♦✱ ❝❛❧❝✉❧❛❞❛s ♣♦r ✶✵ r❡♣❡t✐çõ❡s ❞❛s s✐♠✉❧❛çõ❡s✱ ♣❛r❛ ♦s ✈❛❧♦r❡s ❞❡ ❝♦♥❞✉tâ♥❝✐❛ s✐♥á♣t✐❝❛ ♠á①✐♠❛ gGr ❞❛s

s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s ❡♥tr❡ ❛s ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s ❡ ♠✐tr❛✐s ✐♥❞✐❝❛❞♦s ❛♦ ❧❛❞♦✳ ❖s ❞❛❞♦s sã♦ ♠♦str❛❞♦s ♣❛r❛ ♦s ♦❞♦r❡s ✶✱ ✹ ❡ ✺✱ t❛♠❜é♠ ✐♥❞✐❝❛❞♦s ❛♦ ❧❛❞♦ ❞♦ ❣rá✜❝♦✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✻

✹✳✶✾ ❘❡s♣♦st❛ ❞♦ ♠♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✶ ❛♦ ♦❞♦r ✶✿ ❋r❡q✉ê♥❝✐❛s ❞❡ ❞✐s♣❛r♦s ❞❛s

s❡✐s ❝é❧✉❧❛s ♠✐tr❛✐s ❞♦ ♠♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✶ ♣❛r❛ ❛s ❝♦♠❜✐♥❛çõ❡s ❞♦s ✈❛❧♦r❡s ❞❡gGr ✭❡✐①♦x✮ ❡gP G✭❡✐①♦y✮✱ ❡♠µ❙✳ ❆s ❢r❡q✉ê♥❝✐❛s ❢♦r❛♠ ❝❛❧❝✉❧❛❞❛s ♣♦r

♠é❞✐❛s s♦❜r❡ ✶✵ s✐♠✉❧❛çõ❡s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✽

✹✳✷✵ ❘❡s♣♦st❛ ❞♦ ♠♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✶ ❛♦ ♦❞♦r ✷✿ ❋r❡q✉ê♥❝✐❛s ❞❡ ❞✐s♣❛r♦s ❞❛s

♠é❞✐❛s s♦❜r❡ ✶✵ s✐♠✉❧❛çõ❡s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✾

✹✳✷✶ ❘❡s♣♦st❛ ❞♦ ♠♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✶ ❛♦ ♦❞♦r ✸✿ ❋r❡q✉ê♥❝✐❛s ❞❡ ❞✐s♣❛r♦s ❞❛s

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①✈✐✐✐

✹✳✷✷ ❘❡s♣♦st❛ ❞♦ ♠♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✶ ❛♦ ♦❞♦r ✹✿ ❋r❡q✉ê♥❝✐❛s ❞❡ ❞✐s♣❛r♦s ❞❛s

♠é❞✐❛s s♦❜r❡ ✶✵ s✐♠✉❧❛çõ❡s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✻✶

✹✳✷✸ ❘❡s♣♦st❛ ❞♦ ♠♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✶ ❛♦ ♦❞♦r ✺✿ ❋r❡q✉ê♥❝✐❛s ❞❡ ❞✐s♣❛r♦s ❞❛s

♠é❞✐❛s s♦❜r❡ ✶✵ s✐♠✉❧❛çõ❡s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✻✷

✹✳✷✹ ❘❡s♣♦st❛s ❞♦ ♠♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✷✿ ❋r❡q✉ê♥❝✐❛s ❞❡ ❞✐s♣❛r♦s ❞❛s ✷✺ ❝é❧✉❧❛s

♠✐tr❛✐s ❞♦ ♠♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✷ ♣❛r❛ ❛s ❝♦♠❜✐♥❛çõ❡s ❞♦s ✈❛❧♦r❡s ❞❡gGr✭❡✐①♦

x✮ ❡ gP G ✭❡✐①♦ y✮✱ ❡♠ µ❙✱ ♣❛r❛ ♦s ♦❞♦r❡s ✶✱ ✹ ❡ ✺✳ ❆s ❢r❡q✉ê♥❝✐❛s ❢♦r❛♠

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▲✐st❛ ❞❡ ❚❛❜❡❧❛s

✷✳✶ ❈♦♥❝❡♥tr❛çõ❡s ✐ô♥✐❝❛s tí♣✐❝❛s ❡ ♣♦t❡♥❝✐❛✐s ❞❡ ◆❡r♥st ❝♦rr❡s♣♦♥❞❡♥t❡s ♣❛r❛ ✉♠ ♥❡✉rô♥✐♦ ❞❡ ♠❛♠í❢❡r♦s✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✾

✸✳✶ ❉✐♠❡♥sõ❡s ❞♦s ❝♦♠♣❛rt✐♠❡♥t♦s ❞♦ ♠♦❞❡❧♦ ❞♦ ♥❡✉rô♥✐♦ r❡❝❡♣t♦r ♦❧❢❛tór✐♦ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✸ ✸✳✷ ❉✐♠❡♥sõ❡s ❞♦s ❝♦♠♣❛rt✐♠❡♥t♦s ❞♦ ♠♦❞❡❧♦ ❞❛ ❝é❧✉❧❛ ♣❡r❣❧♦♠❡r✉❧❛r✳ ✳ ✷✺ ✸✳✸ ❉❡♥s✐❞❛❞❡s ❞❡ ❝♦♥❞✉tâ♥❝✐❛ ❞❛s ❝♦rr❡♥t❡s ✐ô♥✐❝❛s ❛❞✐❝✐♦♥❛❞❛s ❛♦s

❝♦♠♣❛rt✐♠❡♥t♦s ❞♦ ♠♦❞❡❧♦ ❞❛ ❝é❧✉❧❛ P● ❡ s❡✉s r❡s♣❡❝t✐✈♦s ♣♦t❡♥❝✐❛✐s ❞❡ r❡✈❡rsã♦✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✻ ✸✳✹ P❛râ♠❡tr♦s ❞♦ ♠♦❞❡❧♦ ❞❛ ❝é❧✉❧❛ ♠✐tr❛❧ ❬❉❛✈✐s♦♥ ✷✵✵✶❪ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✼ ✸✳✺ P❛râ♠❡tr♦s ❞♦ ♠♦❞❡❧♦ ❞❛ ❝é❧✉❧❛ ❣r❛♥✉❧❛r ❬❉❛✈✐s♦♥ ✷✵✵✶❪ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✽

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❙✉♠ár✐♦

▲✐st❛ ❞❡ ❋✐❣✉r❛s ①✐✐✐

▲✐st❛ ❞❡ ❚❛❜❡❧❛s ①✐①

✶ ■♥tr♦❞✉çã♦ ✶

✷ ❘❡✈✐sã♦ ❚❡ór✐❝❛ ✻

✷✳✶ Pr♦♣r✐❡❞❛❞❡s ❞♦s ◆❡✉rô♥✐♦s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✻ ✷✳✷ ❊q✉❛çã♦ ❞❛ ♠❡♠❜r❛♥❛ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✽ ✷✳✸ ❋♦r♠❛❧✐s♠♦ ❞❡ ❍♦❞❣❦✐♥✲❍✉①❧❡② ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✶✵ ✷✳✹ ❙✐♥❛♣s❡s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✶✺ ✷✳✺ Pr♦❝❡ss❛♠❡♥t♦ ♦❧❢❛tór✐♦✿ ❈❛r❛❝t❡ríst✐❝❛s ●❡r❛✐s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✶✼ ✷✳✺✳✶ ❈♦♥❡①õ❡s ✐♥tr❛❣❧♦♠❡r✉❧❛r❡s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✶✽ ✷✳✺✳✷ ❈♦♥❡①õ❡s ✐♥t❡r❣❧♦♠❡r✉❧❛r❡s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✵ ✷✳✺✳✸ ❈♦♥❡①õ❡s ❧❛t❡r❛✐s ✐♥✐❜✐tór✐❛s ❡♥tr❡ ❝é❧✉❧❛s ♠✴t ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✵

✸ ▼❡t♦❞♦❧♦❣✐❛ ✷✷

✸✳✶ ▼ét♦❞♦s ◆✉♠ér✐❝♦s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✷ ✸✳✷ ◆❡✉rô♥✐♦ ❘❡❝❡♣t♦r ❖❧❢❛tór✐♦ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✸ ✸✳✷✳✶ ❊stí♠✉❧♦ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✹ ✸✳✸ ❈é❧✉❧❛ P❡r✐❣❧♦♠❡r✉❧❛r ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✺ ✸✳✹ ❈é❧✉❧❛ ▼✐tr❛❧ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✼ ✸✳✺ ❈é❧✉❧❛ ●r❛♥✉❧❛r ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✽ ✸✳✻ ❙✐♥❛♣s❡s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✷✾ ✸✳✼ ▼♦❞❡❧♦ ❞♦ ❇✉❧❜♦ ❖❧❢❛tór✐♦ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✶ ✸✳✼✳✶ ❈❛♠❛❞❛ ●❧♦♠❡r✉❧❛r ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✶

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①①✐

✸✳✼✳✶✳✶ ❈❛♠❛❞❛ ●❧♦♠❡r✉❧❛r ✶ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✷ ✸✳✼✳✶✳✷ ❈❛♠❛❞❛ ●❧♦♠❡r✉❧❛r ✷ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✸ ✸✳✼✳✶✳✸ ❈❛♠❛❞❛ ●❧♦♠❡r✉❧❛r ✸ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✹ ✸✳✼✳✷ ❈❛♠❛❞❛ ●r❛♥✉❧❛r ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✺ ✸✳✼✳✷✳✶ ❈❛♠❛❞❛ ●r❛♥✉❧❛r ✶ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✺ ✸✳✼✳✷✳✷ ❈❛♠❛❞❛ ●r❛♥✉❧❛r ✷ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✼ ✸✳✼✳✸ ❈❛♠❛❞❛s ●❧♦♠❡r✉❧❛r ❡ ●r❛♥✉❧❛r ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✼ ✸✳✼✳✸✳✶ ▼♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✶ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✼ ✸✳✼✳✸✳✷ ▼♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✷ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✽

✹ ❘❡s✉❧t❛❞♦s ✸✾

✹✳✶ ❈é❧✉❧❛s ■♥❞✐✈✐❞✉❛✐s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✸✾ ✹✳✶✳✶ ◆❡✉rô♥✐♦ r❡❝❡♣t♦r ♦❧❢❛tór✐♦ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✹✵ ✹✳✷ ❊stí♠✉❧♦ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✹✷ ✹✳✸ ❊❢❡✐t♦ ❞❛ ✐♥✐❜✐çã♦ ✈✐❛ ❝é❧✉❧❛s ♣❡r✐❣❧♦♠❡r✉❧❛r❡s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✹✸ ✹✳✸✳✶ ▼♦❞❡❧♦ ❞❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ✶ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✹✸ ✹✳✸✳✷ ▼♦❞❡❧♦ ❞❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ✷ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✹✹ ✹✳✸✳✸ ▼♦❞❡❧♦ ❞❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ✸ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✶ ✹✳✹ ❊❢❡✐t♦ ❞❛ ✐♥✐❜✐çã♦ ✈✐❛ ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✸ ✹✳✹✳✶ ▼♦❞❡❧♦ ❞❛ ❝❛♠❛❞❛ ❣r❛♥✉❧❛r ✶ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✸ ✹✳✹✳✷ ▼♦❞❡❧♦ ❞❛ ❝❛♠❛❞❛ ❣r❛♥✉❧❛r ✷ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✻ ✹✳✺ ▼♦❞❡❧♦ ❝♦♠ ❛s ❝❛♠❛❞❛s ❣❧♦♠❡r✉❧❛r ❡ ❣r❛♥✉❧❛r ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✼ ✹✳✺✳✶ ▼♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✶ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✺✼ ✹✳✺✳✷ ▼♦❞❡❧♦ ❝♦♠♣❧❡t♦ ✷ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✻✷

✺ ❉✐s❝✉ssã♦ ❡ ❈♦♥❝❧✉sõ❡s ✻✺

✺✳✶ ▼♦❞❡❧♦s ✐♥❞✐✈✐❞✉❛✐s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✻✺ ✺✳✷ ❊❢❡✐t♦ ✐♥✐❜✐tór✐♦ ✈✐❛ ❝é❧✉❧❛s ♣❡r✐❣❧♦♠❡r✉❧❛r❡s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✻✼ ✺✳✸ ❊❢❡✐t♦ ■♥✐❜✐tór✐♦ ✈✐❛ ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✻✾ ✺✳✹ ❊❢❡✐t♦ ✐♥✐❜✐tór✐♦ ❞❛s ❞✉❛s ❝❛♠❛❞❛s ❝♦♥❥✉♥t❛s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✼✵ ✺✳✺ ❈♦♥❝❧✉sõ❡s ❋✐♥❛✐s ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✳ ✼✶

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❈❛♣ít✉❧♦

1

■♥tr♦❞✉çã♦

❊♠ ❛♠❜✐❡♥t❡s ♥❛t✉r❛✐s✱ ♠♦❧é❝✉❧❛s ♦❞♦r❛♥t❡s s❡ ❡♥❝♦♥tr❛♠ ❞✐str✐❜✉í❞❛s ❞❡ ❢♦r♠❛ ✐♠♣r❡✈✐sí✈❡❧ ♥♦ ❡s♣❛ç♦ ❡ ♥♦ t❡♠♣♦ ❝♦♠♦ r❡s✉❧t❛❞♦ ❞❛ ❧✐✈r❡ ♠✐st✉r❛ ❡ ❞❡ ✢✉①♦s t✉r❜✉❧❡♥t♦s ❞❡ ♦❞♦r❛♥t❡s ♣r♦✈❡♥✐❡♥t❡s ❞❡ ✐♥ú♠❡r❛s ❢♦♥t❡s✳ ❖ s✐st❡♠❛ ♦❧❢❛tór✐♦ ❞❡t❡❝t❛ s✐♥❛✐s ❞❡ ✐♥t❡r❡ss❡ ❞❡♥tr♦ ❞❡ss❡s ❛♠❜✐❡♥t❡s q✉✐♠✐❝❛♠❡♥t❡ r✉✐❞♦s♦s ♣❛r❛ ❢♦r♠❛r r❡♣r❡s❡♥t❛çõ❡s ❞❡ ❡stí♠✉❧♦s ♦❞♦r❛♥t❡s✱ ❞✐❢❡r❡♥❝✐❛r ❡stí♠✉❧♦s r❡❧❡✈❛♥t❡s ❞❡ ✐rr❡❧❡✈❛♥t❡s ❡ ❝♦♥❞✉③✐r ❛ ✉♠❛ r❡s♣♦st❛ ❛♣r♦♣r✐❛❞❛ ♣♦r ♣❛rt❡ ❞♦ ♦r❣❛♥✐s♠♦✳ ❖ ❜✉❧❜♦ ♦❧❢❛tór✐♦ é ♦ ♣r✐♠❡✐r♦ ❡stá❣✐♦ ❞❡ ♣r♦❝❡ss❛♠❡♥t♦ ❞❡ ♦❞♦r❛♥t❡s ❡♠ ✈❡rt❡❜r❛❞♦s ❬❙❤❡♣❤❡r❞ ✷✵✵✹❪✳ ❙✉❛ ♣r✐♥❝✐♣❛❧ ❢✉♥çã♦ é r❡❝❡❜❡r ❡ ♣r♦❝❡ss❛r s✐♥❛✐s ♠♦❧❡❝✉❧❛r❡s ❡ ♣r♦❞✉③✐r r❡♣r❡s❡♥t❛çõ❡s ❞❡ss❡s s✐♥❛✐s q✉❡ ❧❡✈❡♠✱ ❞❡ ❛❧❣✉♠❛ ❢♦r♠❛✱ ❛♦ s❡♥t✐❞♦ ❞♦ ❝❤❡✐r♦✳ ❖ ❜✉❧❜♦ ♦❧❢❛tór✐♦ r❡❝❡❜❡ ❡♥tr❛❞❛ s❡♥s♦r✐❛❧ ❞✐r❡t❛ ♣r♦✈❡♥✐❡♥t❡ ❞♦s ♥❡✉rô♥✐♦s r❡❝❡♣t♦r❡s ❞♦ ❡♣✐té❧✐♦ ♦❧❢❛tór✐♦ ❡ ❡♥✈✐❛ ✐♥❢♦r♠❛çã♦ ♣❛r❛ ♦ ❝órt❡① ♦❧❢❛tór✐♦✱ s✐st❡♠❛ ❧í♠❜✐❝♦ ❡ ♦✉tr❛s r❡❣✐õ❡s ❞♦ ❝ér❡❜r♦✳

❈♦♠ ✉♠❛ ❡str✉t✉r❛ ❜❡♠ ❞❡✜♥✐❞❛✱ ❞✐✈✐❞✐❞❛ ❡♠ ♠ó❞✉❧♦s ❢✉♥❝✐♦♥❛❧♠❡♥t❡ ❡ ❛♥❛t♦♠✐❝❛♠❡♥t❡ ❞✐st✐♥t♦s✱ ♦ ❜✉❧❜♦ ♦❧❢❛tór✐♦ ✭❇❖✮ é ❜❛s✐❝❛♠❡♥t❡ ✉♠❛ r❡❞❡ ♥❡✉r❛❧ ❞❡ ❛❧✐♠❡♥t❛çã♦ ❞✐r❡t❛ ❝♦♠ ✐♥t❡r❛çõ❡s ❧❛t❡r❛✐s✱ ♣♦✉❝❛s ❝♦♥❡①õ❡s r❡❝♦rr❡♥t❡s ❡ ❡♥tr❛❞❛ ❜❡♠ ❝❛r❛❝t❡r✐③❛❞❛✳ ■ss♦ ❝♦♥tr❛st❛ ❝♦♠ ♦✉tr❛s ár❡❛s ❞♦ ❝ér❡❜r♦ ❝♦♠♦ ♦ ❤✐♣♦❝❛♠♣♦ ❡ ♦ ❝órt❡①✱ ♣♦r ❡①❡♠♣❧♦✱ ❝✉❥❛s ❡str✉t✉r❛s sã♦ ♠❛✐s ❝♦♠♣❧❡①❛s ❬❙❤❡♣❤❡r❞ ✷✵✵✹❪✳ P♦r ❝❛r❛❝t❡ríst✐❝❛s ❝♦♠♦ ❡st❛✱ ♦ ❇❖ é ❝♦♥s✐❞❡r❛❞♦ ✉♠ s✐st❡♠❛ ♠♦❞❡❧♦ ♣❛r❛ ♦ ❡st✉❞♦ ❞❡ ♠❡❝❛♥✐s♠♦s ♥❡✉r❛✐s ❡ s✐♥á♣t✐❝♦s r❡s♣♦♥sá✈❡✐s ♣❡❧♦ ♣r♦❝❡ss❛♠❡♥t♦ ❞❡ ✐♥❢♦r♠❛çã♦ ♥♦ ❝ér❡❜r♦✳

❆s ❝❛r❛❝t❡ríst✐❝❛s ❞♦s ❡stí♠✉❧♦s ♦❧❢❛tór✐♦s t❛♠❜é♠ ❞ã♦ ❛♦ ❡st✉❞♦ ❞♦ ♣r♦❝❡ss❛♠❡♥t♦ ♦❧❢❛tór✐♦ ♣❡❧♦ ❇❖ ✈❛♥t❛❣❡♥s ❡♠ ❝♦♠♣❛r❛çã♦ ❝♦♠ ♦✉tr♦s s✐st❡♠❛s

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✶ ✲ ■♥tr♦❞✉çã♦ ✷

s❡♥s♦r✐❛✐s✳ Pr✐♠❡✐r♦✱ ♣♦rq✉❡ ❛ ❞✐♥â♠✐❝❛ ❞♦ s✐♥❛❧ ❞❡ ♦❞♦r é ♠❛✐s ❧❡♥t❛ ❡♠

r❡❧❛çã♦ ❛ ♦✉tr♦s ❡stí♠✉❧♦s✱ ✈✐s✉❛✐s ❡ ❛✉❞✐t✐✈♦s ♣♦r ❡①❡♠♣❧♦✳ ❙❡❣✉♥❞♦✱ ♣♦rq✉❡ ❛s ♣r♦♣r✐❡❞❛❞❡s q✉❡ ❞❡✈❡♠ s❡r ✐❞❡♥t✐✜❝❛❞❛s sã♦ ❛ ✐❞❡♥t✐❞❛❞❡ ❡ ❛ ✐♥t❡♥s✐❞❛❞❡ ✭❝♦♥❝❡♥tr❛çã♦✮ ❞♦ ♦❞♦r✳ ❈❛❞❛ s✉❜stâ♥❝✐❛ ♦❞♦r❛♥t❡ ♣r♦❞✉③ ✉♠ ♣❛❞rã♦ ❡s♣❛❝✐❛❧ ❞❡ ❛t✐✈❛çã♦ ❡s♣❡❝í✜❝♦ ✕ ✉♠ ♠❛♣❛ ❞♦ ♦❞♦r❛♥t❡ ✕✱ ♥❛ ♣r✐♠❡✐r❛ ❝❛♠❛❞❛ ❞♦ ❇❖✱ ❛ ❝❛♠❛❞❛ ❣❧♦♠❡r✉❧❛r ❬❯❝❤✐❞❛ ❡t ❛❧✳ ✷✵✵✵✱ ❳✉✱ ●r❡❡r ❡ ❙❤❡♣❤❡r❞ ✷✵✵✵❪✳ ❊ss❡ ♣❛❞rã♦ ❡s♣❛❝✐❛❧ é tr❛♥s❢♦r♠❛❞♦ ♣❡❧♦ ❇❖✱ ❞❡✈✐❞♦ à s✉❛ ❝♦♥❡❝t✐✈✐❞❛❞❡ ✐♥t❡r♥❛✱ ❡♠ ✉♠ ♣❛❞rã♦ ❡s♣❛ç♦✲t❡♠♣♦r❛❧ ❞❡ ❛t✐✈✐❞❛❞❡ ❬▲❛✉r❡♥t ❡t ❛❧✳ ✷✵✵✶✱ ❙♣♦rs ❡ ●r✐♥✈❛❧❞ ✷✵✵✷✱ ❙❝❤♦♣♣❛ ❡ ❯r❜❛♥ ✷✵✵✸✱ ❙t♦♣❢❡r✱ ❏❛②❛r❛♠❛♥ ❡ ▲❛✉r❡♥t ✷✵✵✸✱ ❘✉❜✐♥ ❡ ❈❧❡❧❛♥❞ ✷✵✵✻❪ r❡✢❡t✐❞♦ ♥♦s ♣❛❞rõ❡s ❞❡ ❞✐s♣❛r♦ ❞❛s s✉❛s ❝é❧✉❧❛s ❞❡ s❛í❞❛✱ ❛s ❝é❧✉❧❛s ♠✐tr❛✐s ❡ t✉❢♦s❛s ✭♠✴t✮✱ q✉❡ tr❛♥s♠✐t❡♠ ♦s r❡s✉❧t❛❞♦s ❞❛s ❝♦♠♣✉t❛çõ❡s ❢❡✐t❛s ♣❡❧♦ ❇❖ ❛♦s ❝❡♥tr♦s ♦❧❢❛tór✐♦s s✉♣❡r✐♦r❡s✳ ❖ ❡♥t❡♥❞✐♠❡♥t♦ ❞♦s ♠❡❝❛♥✐s♠♦s ♣❡❧♦s q✉❛✐s ❛ ❝✐r❝✉✐t❛r✐❛ ✐♥t❡r♥❛ ❞♦ ❇❖ ♣r♦❞✉③ ❡ ♠❛♥té♠ ❡ss❡s ♣❛❞rõ❡s ❞❡ ❛t✐✈✐❞❛❞❡✱ ♣r♦♠♦✈❡♥❞♦ ♦ r❡❝♦♥❤❡❝✐♠❡♥t♦ ❡ ❛ ❞✐s❝r✐♠✐♥❛çã♦ ❞❡ ♦❞♦r❡s✱ ❜❡♠ ❝♦♠♦ ♦ ♣❛♣❡❧ q✉❡ t❛✐s ♣❛❞rõ❡s ❡s♣❛ç♦✲t❡♠♣♦r❛✐s t❡r✐❛♠ ♥♦ ♣r♦❝❡ss❛♠❡♥t♦ ❞❛ ✐♥❢♦r♠❛çã♦ ♦❧❢❛tór✐❛✱ é ✉♠❛ ❞❛s q✉❡stõ❡s ❝❡♥tr❛✐s ❞❛ ♥❡✉r♦❝✐ê♥❝✐❛ ♠♦❞❡r♥❛ ❬❍❛❜❡r❧② ✷✵✵✶✱ ❋r✐❡❞r✐❝❤ ❡ ❑♦rs❝❤✐♥❣ ✶✾✾✼✱ ❑❛② ❡ ❙t♦♣❢❡r ✷✵✵✻❪✳

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✶ ✲ ■♥tr♦❞✉çã♦ ✸

♣❡r✐❣❧♦♠❡r✉❧❛r❡s✳ ❉❡ ❢♦r♠❛ s✐♠✐❧❛r✱ ♦s ❞❡♥❞r✐t♦s ❞❛s ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s ❢❛③❡♠ s✐♥❛♣s❡s ✐♥✐❜✐tór✐❛s ♥♦s ❞❡♥❞r✐t♦s s❡❝✉♥❞ár✐♦s ❞❛s ❝é❧✉❧❛s ♠✴t ❡ ❡st❡s ❞❡♥❞r✐t♦s ❢❛③❡♠ s✐♥❛♣s❡s ❡①❝✐t❛tór✐❛s ♥♦s ❞❡♥❞r✐t♦s ❞❛s ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s✳ ❆ ❝✐r❝✉✐t❛r✐❛ r❡❝í♣r♦❝❛ ❢♦r♠❛❞❛ ♣♦r ❡ss❡s ❞♦✐s ✐♥t❡r♥❡✉rô♥✐♦s ❢♦✐ ♣r♦♣♦st❛ ❝♦♠♦ r❡s♣♦♥sá✈❡❧ ♣♦r ❢❡♥ô♠❡♥♦s ❝♦♠♦ ♠❡❧❤♦r❛♠❡♥t♦ ❞❡ ❝♦♥tr❛st❡ ❡ s✐♥❝r♦♥✐③❛çã♦♦ ❞❛ ❛t✐✈✐❞❛❞❡ ❞❡ ❝é❧✉❧❛s ♠✴t ❬❨♦❦♦✐✱ ▼♦r✐ ❡ ◆❛❦❛♥✐s❤✐ ✶✾✾✺✱ ▲❛✉r❡♥t ✶✾✾✾✱ ❙❝❤♦♣♣❛ ❡ ❯r❜❛♥ ✷✵✵✸✱ ❈❧❡❧❛♥❞ ❡ ❙❡t❤✉♣❛t❤② ✷✵✵✻✱ ❋❛♥t❛♥❛✱ ❙♦✉❝② ❡ ▼❡✐st❡r ✷✵✵✽❪✳

❆❧❣✉♥s ❢❡♥ô♠❡♥♦s q✉❡ ♦❝♦rr❡♠ ♥♦ ❇❖ ♦❝♦rr❡♠ t❛♠❜é♠ ❡♠ ♦✉tr❛s r❡❣✐õ❡s ❝❡r❡❜r❛✐s ❡ sã♦ ❞❡ ❣r❛♥❞❡ ✐♥t❡r❡ss❡ ♣❛r❛ ♦ ❡♥t❡♥❞✐♠❡♥t♦ ❞♦ ♣r♦❝❡ss❛♠❡♥t♦ ♥❡✉r❛❧✳ ❯♠ ❡①❡♠♣❧♦ sã♦ ❛s ♦s❝✐❧❛çõ❡s ❡❧étr✐❝❛s✱ q✉❡ ♥♦ ❇❖ ❡①✐❜❡♠ ✉♠❛ ❢❛s❝✐♥❛♥t❡ r❡❣✉❧❛r✐❞❛❞❡ t❡♠♣♦r❛❧ ❡ ❝♦♠♣♦rt❛♠❡♥t❛❧ ❬❑❛② ❡ ❙t♦♣❢❡r ✷✵✵✻❪✳ ❈✐r❝✉✐t♦s ♥❡✉r❛✐s ♣♦ss✉❡♠ ❛ ❤❛❜✐❧✐❞❛❞❡ ❞❡ ♦rq✉❡str❛r ❛t✐✈✐❞❛❞❡s ❡♠ ❞✐❢❡r❡♥t❡s ❡s❝❛❧❛s ❞❡ t❡♠♣♦✳ ❖ ❜✉❧❜♦ ♦❧❢❛tór✐♦✱ ❡♠ ♣❛rt✐❝✉❧❛r✱ é ✉♠ ❡①❡♠♣❧♦ ❝❧áss✐❝♦ ❞❡ ✉♠❛ r❡❣✐ã♦ ♥❛ q✉❛❧ rít♠♦s ❧❡♥t♦s ❡ rá♣✐❞♦s ❝♦❡①✐st❡♠✱ s✉♣♦st❛♠❡♥t❡ ❝♦♥tr♦❧❛❞♦s ♣♦r ✐♥t❡r♥❡✉rô♥✐♦s ✐♥✐❜✐tór✐♦s s❡❣r❡❣❛❞♦s ❡♠ ❝❛♠❛❞❛s ❞✐st✐♥t❛s✱ ♥✐t✐❞❛♠❡♥t❡ s❡♣❛r❛♥❞♦ ✐♥✐❜✐çõ❡s ❧♦❝❛✐s ❡ ❣❧♦❜❛✐s ❬❋✉❦✉♥❛❣❛ ❡t ❛❧✳ ✷✵✶✹❪✳ ◆♦ ❡♥t❛♥t♦ ❛ ♠❛♥❡✐r❛ ♣❡❧❛ q✉❛❧ ❝✐r❝✉✐t♦s ❞✐st✐♥t♦s ✐♥t❡r❛❣❡♠ ♣❛r❛ ❡s❝✉❧♣✐r ❞✐✈❡rs♦s rít♠♦s ❛✐♥❞❛ ♣❡r♠❛♥❡❝❡ ♥ã♦ r❡s♦❧✈✐❞❛✳ ❆ ✐♥✈❡st✐❣❛çã♦ ❡①♣❡r✐♠❡♥t❛❧ ✐♥ ✈✐✈♦ ❞❡ss❛s ❤✐♣ót❡s❡s ❡♥❢r❡♥t❛ ❞❡s❛✜♦s ♣❛rt✐❝✉❧❛r❡s ♣♦rq✉❡ ♦s ✐♥t❡r♥❡✉rô♥✐♦s ❡s♣❡❝í✜❝♦s ❡♥✈♦❧✈✐❞♦s ♥❛ ❣❡r❛çã♦ ❡ ❝♦♦r❞❡♥❛çã♦ ❞♦s ❞✐❢❡r❡♥t❡s rít♠♦s ❡stã♦ ❧♦❝❛❧✐③❛❞♦s ❡♠ r❡❣✐õ❡s ❛♥❛t♦♠✐❝❛♠❡♥t❡ s♦❜r❡♣♦st❛s✱ ✐♥✈✐❛❜✐❧✐③❛♥❞♦ s❡✉ ❛❝❡ss♦✳ ❉✐❛♥t❡ ❞❛ ✐♠♣♦rtâ♥❝✐❛ ❞♦s ❝✐r❝✉✐t♦s ✐♥✐❜✐tór✐♦s ♣❛r❛ ❛ ❝♦♠♣✉t❛çã♦ ♥❡✉r❛❧✱ ✈✐st♦ q✉❡ ❡❧❡s ❞❡s❡♠♣❡♥❤❛♠ ♣❛♣❡❧ ❝r✉❝✐❛❧ ♥❛ ❢♦r♠❛çã♦ ❞❡ ❡str✉t✉r❛s t❡♠♣♦r❛✐s ❡ ♥❛ ❛♠♣❧❛ ❢❛✐①❛ ❞❡ r✐t♠♦s ♦s❝✐❧❛tór✐♦s ♦❜s❡r✈❛❞❛✱ ♥❡st❡ tr❛❜❛❧❤♦ ♣r♦♣õ❡✲s❡ ❛ ❝♦♥str✉çã♦ ❞❡ ✉♠ ♠♦❞❡❧♦ ❜✐♦❧♦❣✐❝❛♠❡♥t❡ ♣❧❛✉sí✈❡❧ ❞♦ ❇❖ ♣❛r❛ ✐♥✈❡st✐❣❛r ♣♦r ♠❡✐♦ ❞❡ s✐♠✉❧❛çõ❡s ❝♦♠♣✉t❛❝✐♦♥❛✐s ♦ ❡❢❡✐t♦ ❞❛s ✐♥t❡r❛çõ❡s r❡❝í♣r♦❝❛s ❡♥tr❡ ♥❡✉rô♥✐♦s ✐♥✐❜✐tór✐♦s ❡ ❡①❝✐t❛tór✐♦s ❞♦ ❇❖✳ ❖ ♠♦❞❡❧♦ ♣r♦♣♦st♦ é ❝♦♥st✐t✉í❞♦ ♣♦r ❝é❧✉❧❛s ♠✐tr❛✐s ❡ ✐♥t❡r♥❡✉rô♥✐♦s ❞♦s ❞♦✐s t✐♣♦s ♠❡♥❝✐♦♥❛❞♦s ❛❝✐♠❛✿ ❝é❧✉❧❛s ♣❡r✐❣❧♦♠❡r✉❧❛r❡s ❡ ❝é❧✉❧❛s ❣r❛♥✉❧❛r❡s✳ ❈♦♠♦ é ✐♠♣♦rt❛♥t❡ ❡st✉❞❛r ❛s ✐♥t❡r❛çõ❡s ❡♥tr❡ ❡ss❡s ♥❡✉rô♥✐♦s ❡♠ t❡r♠♦s ❞❡ s❡✉s ❝♦♥t❛t♦s s✐♥á♣t✐❝♦s r❡❝í♣r♦❝♦s✱ ♦s q✉❛✐s ♦❝♦rr❡♠ ❡♠ ❞❡♥❞r✐t♦s ❞✐❢❡r❡♥t❡s ❞❛ ❝é❧✉❧❛ ♠✐tr❛❧✱ ♦s ♥❡✉rô♥✐♦s sã♦ ❛q✉✐ ♠♦❞❡❧❛❞♦s ❞❡ ♠❛♥❡✐r❛ ❜✐♦❢ís✐❝❛ ❡ ♠♦r❢♦❧♦❣✐❝❛♠❡♥t❡ ❞❡t❛❧❤❛❞❛ ✉s❛♥❞♦ ♦ ❢♦r♠❛❧✐s♠♦ ❞❡ ❍♦❞❣❦✐♥✲❍✉①❧❡② ❡ ❛ té❝♥✐❝❛

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✶ ✲ ■♥tr♦❞✉çã♦ ✹

✐♥❝❧✉✐ t❛♠❜é♠ ✉♠❛ ❝❛♠❛❞❛ ❞❡ ♥❡✉rô♥✐♦s r❡❝❡♣t♦r❡s✱ ✐❣✉❛❧♠❡♥t❡ ♠♦❞❡❧❛❞♦s ❞❡ ❢♦r♠❛ ❜✐♦✜s✐❝❛♠❡♥t❡ ❞❡t❛❧❤❛❞❛✱ ♣❛r❛ ❢♦r♥❡❝❡r ❡♥tr❛❞❛s r❡♣r❡s❡♥t❛♥❞♦ ❡stí♠✉❧♦s ♦❞♦r❛♥t❡s ❞✐st✐♥t♦s ❛♦ ❇❖✳ ❆ ♠❛✐♦r✐❛ ❞♦s ❡st✉❞♦s ❝♦♠♣✉t❛❝✐♦♥❛✐s ❡①✐st❡♥t❡s s♦❜r❡ ♦ ♣r♦❝❡ss❛♠❡♥t♦ ❞❡ ♦❞♦r❡s ♣❡❧♦ ❇❖ ❝♦♥s✐❞❡r❛ ❛♣❡♥❛s ❛s ❝é❧✉❧❛s ♠✐tr❛✐s✱ ❛s ❣r❛♥✉❧❛r❡s ❡ ♥❡✉rô♥✐♦s s❡♥s♦r✐❛✐s✳ P♦✉❝♦s tr❛❜❛❧❤♦s✱ ♥♦ ❝♦♥t❡①t♦ ❞❛ ♠♦❞❡❧❛❣❡♠ ❝♦♠♣✉t❛❝✐♦♥❛❧✱ ✐♥❝♦r♣♦r❛♠ ❛ ❝é❧✉❧❛ ♣❡r✐❣❧♦♠❡r✉❧❛r ❡♠ s❡✉s ♠♦❞❡❧♦s✳ P♦rt❛♥t♦✱ ❡st❡ tr❛❜❛❧❤♦ ♦❢❡r❡❝❡ ✉♠❛ ❝♦♥tr✐❜✉✐çã♦ ♦r✐❣✐♥❛❧ ❛♦ ❡st✉❞♦ ❞♦ ♣r♦❝❡ss❛♠❡♥t♦ ❞❡ ♦❞♦r❛♥t❡s ♣❡❧♦ ❇❖✱ ♣♦✐s ❝♦♥s✐❞❡r❛ ♦ ♣❛♣❡❧ ✐♥t❡❣r❛❞♦ ❞♦s ❞♦✐s ♣r✐♥❝✐♣❛✐s ✐♥t❡r♥❡✉rô♥✐♦s ✐♥✐❜✐tór✐♦s ❞♦ ❇❖ ♥❡st❡ ♣r♦❝❡ss❛♠❡♥t♦✳ ❆♣r♦✈❡✐t❛♥❞♦ ❛s ❢❛❝✐❧✐❞❛❞❡s ❞❡ s❡ ♠❛♥✐♣✉❧❛r ❛ ✧❛♥❛t♦♠✐❛✧❞❡ ✉♠ ♠♦❞❡❧♦ ❝♦♠♣✉t❛❝✐♦♥❛❧✱ ♦ tr❛❜❛❧❤♦ t❛♠❜é♠ ❡st✉❞♦ ♦ ❡❢❡✐t♦ ❞❡ ❝❛❞❛ ✐♥t❡r♥❡✉rô♥✐♦ ✐s♦❧❛❞❛♠❡♥t❡ ✭❡❧✐♠✐♥❛♥❞♦ ♦ ♦✉tr♦ t✐♣♦ ❞❡ ✐♥t❡r♥❡✉rô♥✐♦ ❞♦ ❝✐r❝✉✐t♦✮ ♣❛r❛ ♠❡❧❤♦r ❡♥t❡♥❞❡r ♦ ♣❛♣❡❧ ❞❡ ❝❛❞❛ ✉♠ s♦❜r❡ ♦ ♣r♦❝❡ss❛♠❡♥t♦✳

❈♦♠♣r❡❡♥❞❡r ❛ ♠❛♥❡✐r❛ ♣❡❧❛ q✉❛❧ ❛ ✐♥❢♦r♠❛çã♦ ♦❧❢❛tór✐❛ é ❣❡r❡♥❝✐❛❞❛ ❡ ♣r♦❝❡ss❛❞❛ ♣❡❧❛ ❝✐r❝✉✐t❛r✐❛ ❞♦ ❇❖ r❡q✉❡r ✉♠❛ ✐♥t❡❣r❛çã♦ ❞❡ ❞❛❞♦s ❡♠ ✈ár✐♦s ♥í✈❡✐s ✭s✉❜✲❝❡❧✉❧❛r✱ ♥❡✉rô♥✐♦ ✐s♦❧❛❞♦✱ ❝✐r❝✉✐t♦ ❧♦❝❛❧✱ s✐st❡♠❛✮ ❡ ♠♦❞❛❧✐❞❛❞❡s ❞❡ ✐♥✈❡st✐❣❛çã♦

✭❜✐♦❧♦❣✐❛ ♠♦❧❡❝✉❧❛r✱ ♥❡✉r♦✜s✐♦❧♦❣✐❛✱ ✐♠❛❣❡❛♠❡♥t♦✱ ♣s✐❝♦❧♦❣✐❛✮✳ ❆ ♠♦❞❡❧❛❣❡♠

❝♦♠♣✉t❛❝✐♦♥❛❧ ❜✐♦❧♦❣✐❝❛♠❡♥t❡ ♣❧❛✉sí✈❡❧ ❝♦♥st✐t✉✐ ✉♠❛ ✐♠♣♦rt❛♥t❡ ❢❡rr❛♠❡♥t❛ ♣❛r❛ ♣r♦♠♦✈❡r ✉♠ ❛✈❛♥ç♦ ♥♦ ❡♥t❡♥❞✐♠❡♥t♦ ❞♦ ❡❢❡✐t♦ ✐♥t❡❣r❛❞♦ ❞♦s ❢❡♥ô♠❡♥♦s ❛❝✐♠❛ s♦❜r❡ ♦ ♣r♦❝❡ss❛♠❡♥t♦ ❞❡ ✐♥❢♦r♠❛çã♦ ♣❡❧♦ ❇❖ ❬❉❛✈✐s♦♥✱ ❋❡♥❣ ❡ ❇r♦✇♥ ✷✵✵✸✱ ❙♦✉③❛ ❡ ❘♦q✉❡ ✷✵✵✹✱ ❘✉❜✐♥ ❡ ❈❧❡❧❛♥❞ ✷✵✵✻✱ ▼✐❣❧✐♦r❡✱ ❍✐♥❡s ❡ ❙❤❡♣❤❡r❞ ✷✵✵✺❪✳ ❆ ❝♦♥str✉çã♦ ❞❡ ✉♠❛ r❡❞❡ ✐♥s♣✐r❛❞❛ ♥❛s ❝♦♥❡①õ❡s ❡①✐st❡♥t❡s ♥♦ ❇❖✱ ❧❡✈❛♥❞♦ ❡♠ ❝♦♥t❛ ❞❛❞♦s ❛♥❛tô♠✐❝♦s ❡ ❜✐♦❢ís✐❝♦s ❡ ♦❜s❡r✈❛♥❞♦ ❛ ❛t✐✈✐❞❛❞❡ ❡ s✉❛ ❞✐♥â♠✐❝❛ é ✉♠ ✐♠♣♦rt❛♥t❡ ♠❡✐♦ ❞❡ ♠❡❧❤♦r❛r ❛ ❝♦♠♣r❡❡♥sã♦ s♦❜r❡ ♦ ♣r♦❝❡ss❛♠❡♥t♦ ❞❡ ✐♥❢♦r♠❛çã♦ ♦❧❢❛tór✐❛ ❡ ❛ ♥❛t✉r❡③❛ ❞❡ s❡✉s ♠❡❝❛♥✐s♠♦s✱ s❡❥❛ ♥♦ s❡♥t✐❞♦ ❞❡ ❝♦♠♣❧❡t❛r ❤✐♣ót❡s❡s ❥á ❡st❛❜❛❧❡❝✐❞❛s ❝♦♠♦ ♥♦ ❞❡ ♣r❡✈❡r r❡s✉❧t❛❞♦s q✉❡ ♣♦❞❡rã♦ s❡r t❡st❛❞♦s ❡①♣❡r✐♠❡♥t❛❧♠❡♥t❡✳

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✶ ✲ ■♥tr♦❞✉çã♦ ✺

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❈❛♣ít✉❧♦

2

❘❡✈✐sã♦ ❚❡ór✐❝❛

❊st❡ ❝❛♣ít✉❧♦ ❛❜r❛❣❡ ✉♠❛ r❡✈✐sã♦ ❞♦s ♣r✐♥❝✐♣❛✐s ❝♦♥❝❡✐t♦s ♥❡❝❡ssár✐♦s ♣❛r❛ ♦ ❞❡s❡♥✈♦❧✈✐♠❡♥t♦ ❞❡st❡ tr❛❜❛❧❤♦✳ Pr✐♠❡✐r❛♠❡♥t❡ ❛♣r❡s❡♥t❛♠♦s ❝♦♥❝❡✐t♦s ❜✐♦❢ís✐❝♦s ❢✉♥❞❛♠❡♥t❛✐s ♣❛r❛ ❛ ♠♦❞❡❧❛❣❡♠ ❝♦♠♣✉t❛❝✐♦♥❛❧ ❞❡ ♥❡✉rô♥✐♦s ❜✐♦❧♦❣✐❝❛♠❡♥t❡ ♣❧❛✉sí✈❡✐s ❡✱ ❡♠ s❡❣✉✐❞❛✱ ♣❛ss❛♠♦s ♣❛r❛ ✉♠❛ r❡✈✐sã♦ ❞♦ ♣r♦❝❡ss❛♠❡♥t♦ ♦❧❢❛tór✐♦ ❡ ❞❛s ❝❛r❛❝t❡ríst✐❝❛s ❣❡r❛✐s ❞♦ ❜✉❧❜♦ ♦❧❢❛tór✐♦✳

✷✳✶ Pr♦♣r✐❡❞❛❞❡s ❞♦s ◆❡✉rô♥✐♦s

❖ ♥❡✉rô♥✐♦ é ❛ ❝é❧✉❧❛ ❞♦ s✐st❡♠❛ ♥❡r✈♦s♦ ❡s♣❡❝✐❛❧✐③❛❞❛ ❡♠ ♣r♦♣❛❣❛r s✐♥❛✐s ❡❧étr✐❝♦s ❝❛r❛❝t❡ríst✐❝♦s✱ ❝❤❛♠❛❞♦s ♣♦t❡♥❝✐❛✐s ❞❡ ❛çã♦✱ ♦✉ s✐♠♣❧❡s♠❡♥t❡ ❞✐s♣❛r♦s ✭❞♦ ✐♥❣❧ês s♣✐❦❡s✮✳ ❖s ♥❡✉rô♥✐♦s tê♠ ❝❛♣❛❝✐❞❛❞❡ ❞❡ r❡♣r❡s❡♥t❛r ❡ tr❛♥s♠✐t✐r ✐♥❢♦r♠❛çõ❡s ❡♠✐t✐♥❞♦ s❡q✉ê♥❝✐❛s ❞❡ ❞✐s♣❛r♦s ❝♦♠ ✈ár✐♦s ♣❛❞rõ❡s t❡♠♣♦r❛✐s ❬❉❛②❛♥ ❡ ❆❜❜♦tt ✷✵✵✷❪✳ ❊♠ ❣❡r❛❧✱ ❛ ♠♦r❢♦❧♦❣✐❛ ♥❡✉r♦♥❛❧ ❝♦♥s✐st❡ ❡♠ três ♣❛rt❡s ♣r✐♥❝✐♣❛✐s✿ s♦♠❛✱ ❞❡♥❞r✐t♦ ❡ ❛①ô♥✐♦✳ ❖s ❞❡♥❞r✐t♦s r❡❝❡❜❡♠ s✐♥❛✐s ❞❡ ❡♥tr❛❞❛ ❞❡ ♦✉tr♦s ♥❡✉rô♥✐♦s ❡ ♦ ❛①ô♥✐♦ ♣r♦♣❛❣❛ ♦ s✐♥❛❧ ✐♥t❡❣r❛❞♦ ♣❡❧♦ s♦♠❛ ♣❛r❛ ♦✉tr❛s ❝é❧✉❧❛s ❬P✉r✈❡s ❡t ❛❧✳ ✷✵✵✹❪✳

❆ ✜❣✉r❛ ✷✳✶ ✐❧✉str❛ ❛ ❞✐✈❡rs✐❞❛❞❡ ❞❡ ❢♦r♠❛s ❞❡ ♥❡✉rô♥✐♦s ❡♥❝♦♥tr❛❞❛ ♥♦ ❝ér❡❜r♦✳ ❊ss❛ ❞✐✈❡rs✐❞❛❞❡ é ❞❡✈✐❞❛ ♣r✐♥❝✐♣❛❧♠❡♥t❡ à ❡str✉t✉r❛ ❞❛ ár✈♦r❡ ❞❡♥❞rít✐❝❛ ❞♦s ♥❡✉rô♥✐♦s✳ ❆ r❛♠✐✜❝❛çã♦ ❡❧❛❜♦r❛❞❛ ❞❡ss❛s ár✈♦r❡s ♣❡r♠✐t❡ ❛ ✉♠ ♥❡✉rô♥✐♦ r❡❝❡❜❡r s✐♥❛✐s ❞❡ ♠✉✐t♦s ♦✉tr♦s ♥❡✉rô♥✐♦s ❛tr❛✈és ❞❡ ❝♦♥❡①õ❡s s✐♥á♣t✐❝❛s✳ ❖ ♥❡✉rô♥✐♦ ♣✐r❛♠✐❞❛❧ ❝♦rt✐❝❛❧ ❞❛ ✜❣✉r❛ ✷✳✶❆ ❡ ♦ ✐♥t❡r♥❡✉rô♥✐♦ ❝♦rt✐❝❛❧ ❞❛ ✜❣✉r❛ ✷✳✶❈ r❡❝❡❜❡♠ ♠✐❧❤❛r❡s ❞❡ ❡♥tr❛❞❛s s✐♥á♣t✐❝❛s✳ P❛r❛ ❛ ❝é❧✉❧❛ ❞❡ P✉r❦✐♥❥❡ ❞♦ ❝❡r❡❜❡❧♦ ❞❛ ✜❣✉r❛ ✷✳✶❇ ♦ ♥ú♠❡r♦ ❞❡ ❡♥tr❛❞❛s s✐♥á♣t✐❝❛s é s✉♣❡r✐♦r ❛ ✶✵✵✳✵✵✵✳ ❆ ❋✐❣✉r❛ ✷✳✶ ♥ã♦ ♠♦str❛

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✷✳✶ ✲ Pr♦♣r✐❡❞❛❞❡s ❞♦s ◆❡✉rô♥✐♦s ✼

A B

C

soma dendrito

apical

dendrito basal soma

axônio colateral

axônio dendrito

axônio

soma dendrito

❋✐❣✉r❛ ✷✳✶✿ ❉✐✈❡rs✐❞❛❞❡ ♠♦r❢♦❧ó❣✐❝❛ ❞♦s ♥❡✉rô♥✐♦s✳ ❆✲ ❈é❧✉❧❛ ♣✐r❛♠✐❞❛❧ ❞♦ ❝órt❡① ❝❡r❡❜r❛❧✳ ❇✲ ❈é❧✉❧❛ ❞❡ P✉r❦✐♥❥❡ ❞♦ ❝❡r❡❜❡❧♦✳ ❈✲ ❈é❧✉❧❛ ❡str❡❧❛❞❛ ❞♦ ❝órt❡① ❝❡r❡❜r❛❧✳ ✭❋✐❣✉r❛s ❛❞❛♣t❛❞❛s ❛ ♣❛rt✐r ❞❡ ❞❡s❡♥❤♦s ♦r✐❣✐♥❛✐s ❞❡ ❘❛♠ó♥ ② ❈❛❥❛❧ ❞✐s♣♦♥í✈❡✐s ♥❛ ✐♥t❡r♥❡t ❡♠ ❤tt♣✿✴✴❝✈❝✳❝❡r✈❛♥t❡s✳❡s✴❝✐❡♥❝✐❛✴❝❛❥❛❧✴❞❡❢❛✉❧t✳❤t♠✮

t♦❞❛ ❛ ❡①t❡♥sã♦ ❞♦s ❞❡♥❞r✐t♦s ❡ ❛①ô♥✐♦s ❞❡ss❡s ♥❡✉rô♥✐♦s✳ ❆①ô♥✐♦s ❞❡ ❞❡t❡r♠✐♥❛❞♦s ♥❡✉rô♥✐♦s ♣♦❞❡♠ ❡①t❡♥❞❡r✲s❡ ♣♦r ❣r❛♥❞❡s r❡❣✐õ❡s ❞♦ ❝ér❡❜r♦ ♦✉✱ ❡♠ ❛❧❣✉♥s ❝❛s♦s✱ ♣♦r t♦❞♦ ♦ ❝♦r♣♦✳

❆❧é♠ ❞❛s ❝❛r❛❝t❡ríst✐❝❛s ♠♦r❢♦❧ó❣✐❝❛s✱ ♦s ♥❡✉rô♥✐♦s tê♠ ❡s♣❡❝✐❛❧✐③❛çõ❡s ✜s✐♦❧ó❣✐❝❛s ❝♦♠ ❞❡st❛q✉❡ ♣❛r❛ ❛ ❣r❛♥❞❡ ✈❛r✐❡❞❛❞❡ ❞❡ ❝❛♥❛✐s ✐ô♥✐❝♦s ♥❛ ♠❡♠❜r❛♥❛ ♥❡✉r♦♥❛❧✱ ❛ q✉❛❧ é ❢♦r♠❛❞❛ ♣♦r ✉♠❛ ❞✉♣❧❛ ❝❛♠❛❞❛ ❧✐♣í❞✐❝❛ ❬P✉r✈❡s ❡t ❛❧✳ ✷✵✵✹❪✳ ❖s ❝❛♥❛✐s ✐ô♥✐❝♦s sã♦ ❢♦r♠❛❞♦s ♣♦r ♣r♦t❡í♥❛s q✉❡ ❛tr❛✈❡ss❛♠ ❛ ♠❡♠❜r❛♥❛ ❝❡❧✉❧❛r ♣❡r♠✐t✐♥❞♦ q✉❡ í♦♥s✱ ♣r❡❞♦♠✐♥❛♥t❡♠❡♥t❡ ❞❡ só❞✐♦ ✭◆❛+_{✮✱ ♣♦táss✐♦ ✭❑}+_{✮✱ ❝á❧❝✐♦} ✭❈❛+

2✮ ❡ ❝❧♦r❡t♦ ✭❈❧ −