Mécanismes neuronaux sous-tendant l'apprentissage perceptif olfactif chez la souris adulte
Texte intégral
(2) .
(3) .
(4) .
(5)
(6)
(7)
(8) .
(9)
(10)
(11)
(12). !"#
(13) $ " %%!&# '%!" %%!&# ((&")'&# *+!")'&# ")'&#.
(14) Remerciements. Je voulais tout d’abord remercier les membres du jury, Mr Carleton, Mme Rampon, Mme Caillé, et Mr Gheusi, pour avoir accepté d’évaluer ma thèse, pour leurs remarques pertinentes et pour avoir permis un débat plus qu’intéressant à l’oral ouvrant sur de nombreuses perspectives de recherche. Merci à Christiane Linster pour son aide et sa collaboration qui ont permis d’aller plus loin dans la compréhension des mécanismes de l’apprentissage perceptif. Je remercie également les membres du laboratoire que j’ai côtoyés ces nombreuses années et plus particulièrement les membres de l’équipe d’Anne Didier qui m’ont initiée à la recherche, soutenue et aidée à de nombreuses reprises au cours de ma formation. Merci à Nathalie, Anne et Joëlle qui m’ont toujours soutenue dans mes choix, encouragée, fait énormément progresser et ont réussi à m’amener jusqu’à la soutenance, ce que je ne croyais pas possible. J’ai partagé avec vous encore plus qu’un merveilleux sujet de recherche mais véritablement 6 ans de bonheur, de fous rires, d’aventures et de défis. Nathalie et Anne, merci pour votre encadrement hors du commun, merci pour toute la confiance que vous m’avez accordée ainsi que pour votre amitié. J’ai eu énormément de chance de découvrir la recherche avec vous, c’est une aventure que je n’oublierai jamais. Joëlle, merci pour ton aide extraordinaire et ton soutien, je n’aurais jamais pu avancer les manips sans toi. Tu es un véritable ange gardien ! Je remercie également ma famille, mon mari et mes amis qui m’ont suivie et appuyée dans cette aventure et plus particulièrement Sylvie pour son aide et son soutien de tous les jours..
(15)
(16) !"#
(17)
(18) "
(19) $%
(20) &
(21)
(22) "'" ( )" % & %+, """+"- #"/0"1"2 /" % & %+, ++"3 "
(23) & " % & % +, +4" ' #"&
(24) "'"2 '" " "
(25)
(26) "
(27) +4 """4"3"" % & %"1"" "&
(28) "'"2
(29) %
(30) " % & +4 4+"#
(31)
(32) "/ #0" " (
(33) 0"'"" % & %+4 44" ' #" "'"2 '"5+ """6"7
(34) "'" ( )" % & %+8 !" & " "
(35) "
(36) " &"29
(37) &"
(38) "5+ """"+": '
(39) "'" " " " "
(40) "
(41) + """"4": '
(42) "'" " " "2
(43) #" & %+ """"6": '
(44) "'" " " "2
(45) #"& %4, """"";%% "'"
(46) " " " & " % & " ""&
(47) "54+ !"<"' " "
(48) ' & "'
(49) " "
(50) # " % & % "544 """"+"3" ( )"&/
(51) #046 """"4"3" ( )"
(52)
(53) #04 """"6"3" ( )"
(54) '
(55) #04 """""7
(56) "' " "&
(57) %# 4 7!"3 "
(58) #
(59) ) " "&/>" " " ' 48 """7"+" &
(60) "'" "
(61) #
(62) ) " 48 7"+"+" %
(63) " "#
(64) "' "
(65) 4 7"+"4"
(66) #
(67) " " "' "
(68) $
(69)
(70) " "
(71) "'"" % & %6, """7"4" #
(72) ) " " & "%
(73) &
(74)
(75) "'"" % & %64 7"4"+" #
(76) ) " "
(77) "
(78) 66 7"4"+"+";%% "'" "
(79) " % & "66 4.
(80) 7"4"+"4";%% "'"2
(81) &/
(82) " % & %66 7"4"4" #
(83) ) " "
(84) # " % & % 6 """7"6" &
(85)
(86) " "
(87) #
(88) ) "56 7"6"+"
(89) '&
(90) 6 7"6"4"32
(91) #"& %6 """7"": '
(92) "'" "
(93) #
(94) ) " " " ( ) "
(95) ' "6? """7""7
(96) " "
(97) #
(98) ) " 68 7!"@& % 6 ;A3A4 - '"+6 % & ("& "
(99)
(100) #"0 " ' "
(101) #
(102) - '"4 &
(103) " %" /"
(104) '
(105) #&" ( "
(106) " ' $
(107) "& " "0'"% " % & ("
(108)
(109) #"
(110) "& - '"6
(111) " %" % & ("& "
(112)
(113) #"
(114) " #
(115) #"&"
(116) '" "& " AAA"B;;3;" !"3 "
(117) 9"
(118)
(119) "'"" % & %"1" "'"C
(120) #"& %? !":&
(121) "#
(122)
(123) "C
(124) #
(125) "' "
(126) 9"
(127)
(128) " "'"C
(129) #+ !"3C%% "'"
(130) " "C
(131) #"& %6 """"+"D "'" "
(132) #
(133) ) " "'&
(134) " % & %"56 """"4"3 "
(135) '
(136)
(137) "1"
(138) " '"'2 &
(139) "'%%
(140) "
(141) "%
(142) &
(143) "'"2E#"5 7!"
(144) &
(145) %
(146) & " # /0 ""8" ;F;?8. 6.
(147)
(148) . :. / $
(149) $6$/(' 9($$ /( > $$
(150) . . (
(151) "
(152) '. D. D" % & %. DG. D
(153) "''"
(154) /&"% & . D. $D $4$' 9('
(155) . DH.
(156) " $/(' 9( . BD. &'"# $
(157) (0. B '!?. +$B &" &'"'& 9( . BG. B
(158) "G &
(159) &"
(160). B3. &/"# . B3. &/"#
(161) . 3. 3 & " . 3.
(162)
(163) "I"
(164) #" . ;.
(165) "'" "
(166) '
(167)
(168) . :. $: /($$ . . JA7. J
(169) " "
(170) & . 4$B. 4$ 9($#& . $H. $H(' 9( (
(171) " "
(172)
(173) . .
(174) ;A:; 3 " " % & % " " " )# " '2
(175) "
(176) #
(177) ) " ' "
(178) " " "
(179) " & " & " ' "
(180) # " % & % " "
(181) " "
(182) " "<"'"& " "
(183)
(184) "'"
(185) 9"
(186)
(187)
(188) "
(189) / "'
(190) "
(191) " ("'C
(192) #" &"0C "C
(193) #"& %" "
(194) #K" ) "& #0" "C
(195) "%) "
(196) " & "
(197)
(198) "'C
(199) "'" C
(200)
(201)
(202)
(203) "
(204) " C "
(205) "
(206) # " ' " ( " & " 0 " " '%
(207) " & "
(208) "
(209) "' "%
(210) & "'"' &
(211)
(212) " "I"
(213) "9
(214) " "'" "
(215) "
(216) " ) " 'K "
(217) "
(218) " " 0 " C
(219) # " & % " #
(220) "
(221)
(222) "
(223) " " "' "
(224)
(225) "#
(226) "
(227) % " "#E&" " & #"'" "
(228) #
(229) ) "
(230) "
(231) " #
(232) " "
(233) " '
(234) & " " & & ) "
(235) '
(236) " ' "
(237) 9 "
(238)
(239) " I C &0
(240) "'"& "
(241) #". " #
(242).
(243) "'" "
(244) #
(245) ) " " "&
(246) 0
(247) &"
(248) " #
(249).
(250) " ' "
(251) " 'C
(252) /
(253) " '
(254) " " " % & % " I " C #
(255) " ' " C
(256) " ' " %
(257) & "'"' &
(258)
(259) " 3 " &
(260) " #
(261)
(262) " C
(263) #
(264) " " " " ' "
(265)
(266) "
(267) % "
(268) '
(269) " C
(270) #"
(271) " ) " "&
(272)
(273) K"
(274) "
(275) "% "C/( /) "'C
(276) "<"'" "
(277) '
(278)
(279) "'
(280) " " &
(281) < " ' " C
(282) #
(283) " ' "
(284) 9 "
(285)
(286) " & " " " @. "
(287) " " "
(288)
(289)
(290) " " " " ) "0"'
(291) " "
(292) # " % & % "
(293) "
(294) "'9)" 'K"
(295) "
(296) " "0" "
(297)
(298) "'& "'" "
(299)
(300) "
(301) '
(302) #0" %%& " #
(303) %&
(304) " " " ' "
(305) 9 "
(306)
(307) " " " %
(308) & " '
(309) " C
(310) #" & %";
(311) "%% K"C #
(312).
(313) "'" "
(314) "'"
(315) '
(316)
(317) "&
(318) ' "I"
(319) "
(320) "' %
(321) & " ' " ' &
(322)
(323) "
(324) " 0CI "
(325) " #
(326).
(327) " ' " " " ' "
(328) $
(329)
(330) "
(331)
(332) K" " & #" ' " & "
(333) '
(334) #0 " )" C
(335) " ' "%
(336) & " % & " "
(337) C #
(338) " " "
(339) #
(340) ) " G
(341)
(342) K "
(343) "
(344) "
(345) " 0 " " % "
(346) '
(347) #0 " " @
(348) " " "
(349) 9"
(350)
(351) "0"
(352) '
(353) "I" "
(354) '
(355)
(356) " "0" "
(357) &"' "
(358) 9"
(359)
(360) " "
(361) '
(362) " "0" "
(363) '
(364)
(365) " " " "%
(366) & "'"' &
(367)
(368) " ;
(369) %
(370) K " '
(371) "
(372) " ) " ' "
(373) "
(374) " "
(375) " I " 2%% " ' "
(376) " " 2
(377) #"& %" "
(378) " "
(379) "'% "'2
(380) #"& %"&/>" " "E#" &"I"
(381) "'
(382)
(383) "'" "
(384) #
(385) ) ";
(386) "
(387) &/K"
(388) "
(389) "
(390) '
(391) #0" " '" "2
(392) #"& %"
(393) " '" "
(394) #
(395) ) " " ##
(396) "29
(397) &"' &
(398) "&
(399) " " 9"
(400) " "'"9"&
(401) '""<"' "
(402) 9"
(403)
(404) "'
(405) ""& " ' K" &
(406) " "
(407) " & " " &
(408) " " #
(409) " " " " 9
(410) " " ' " " &
(411) " .
(412)
(413)
(414)
(415)
(416)
(417)
(418)
(419) "
(420) #
(421) "'"% L
(422) "
(423)
(424) " &"
(425) "
(426)
(427)
(428)
(429) ";
(430) "%% K"' " "
(431) " &
(432) "
(433) "
(434) "1" "
(435) K"2 ' K""# N K"2 O" "" &/"
(436) %
(437) L
(438) "
(439) "
(440) " &
(441) " "
(442) ""
(443) K"C % &
(444) K"
(445) "
(446) "
(447) "
(448) ##K" " ) "
(449) ";
(450) " %% K"" "
(451) " "&
(452) "'"&/&/" "'" "'"
(453) " "#
(454) "I" C
(455) " ' " '
(456) # " & " " 9 " ' "
(457) &
(458) ' " " ' " ' " "
(459) %
(460) " I " " '&
(461) "3 " ' "#'
(462) "
(463) "
(464) "&
(465) " " " "
(466) "
(467) "'" " " " "
(468) ( "& " "'C ' "0"
(469) "&
(470) " " "@ "
(471) "'
(472) "'C
(473) " ( )" % & %"& "'"' & K"'
(474) %" "' &
(475) " " ' "3"
(476) "
(477) "' " ' "
(478) " & "'(
(479) 0" " "
(480) #"'"
(481) "" "0"
(482) & "' "
(483) # " "'
(484) &" "
(485) " @ " ' " &
(486) " ' " & " & " " 9 " '
(487) " " ( ) " % & % "
(488) " % " &) " ' " & " "
(489) #
(490) ) " ' " " 0 " &
(491) "
(492) " C "
(493)
(494) " '"
(495) 9"
(496)
(497) "'
(498) ""&& "
(499)
(500) "9
(501) . ;
(502) "+ ,K""
(503) # " #
(504) "A
(505) # "
(506) "(" @ "R+84"S"+ 6T"
(507) ""9" " " " 9" "2/ #"'"& "
(508) "' "
(509) 9" "02" "% " %$ %"0""& " ' " "
(510) " #
(511) "
(512) & "'" "#
(513) 1"U ! " "
(514) K"I" "
(515) $
(516) &K"
(517) "'
(518) "'C
(519) " &V"'"
(520)
(521) "0"
(522) "
(523) "9"'"
(524) "&
(525)
(526) 9
(527) " ""
(528) 2(" " "'2
(529) "'"
(530) 9"
(531)
(532) "&/>"C ' "" " " ) "#
(533) '"
(534) $ %
(535) &"'" W
(536) "(" @ "
(537) "
(538) &
(539) & K"& "
(540)
(541) &" C# "
(542) "' #"
(543) '
(544) K"' " "
(545)
(546) ",K" & "'"& ' "&
(547) &
(548)
(549) " "
(550) #
(551) ) " "
(552) "%%
(553) ' ";
(554) "%% K" " 'C 'K"
(555) "&/&/" &
(556) K"X /"
(557) K"'& "#E&"I"C
(558) @&
(559) "'" /('
(560) " K" ' "& "
(561) "'
(562) " (
(563) "
(564) " / #"'"
(565)
(566) "'
(567) ""& 9K""" % & %" ""#($ "'
(568) "'" " ' "R
(569) "
(570) '" K"+ T"
(571) " "
(572)
(573) "8," "#E&"I" "& & "&$.
(574) 0K":&/"Y
(575) " "' "'
(576)
(577) "
(578) "'
(579) "&"
(580) "
(581) "
(582)
(583) "' "& "
(584) $
(585) "% " ) "
(586) " 9"& "
(587) "'
(588) ""& "'" " "'" "RY
(589) "
(590) '"H
(591) ' K"+ ??T"G
(592)
(593) ' ". /" "#
(594) "&
(595) "&"' #"
(596) "
(597)
(598) "'" "
(599) #$
(600) ) " ' " "&. "% $&"&/>"C "RB '
(601) "
(602) '". /K"+ 86T"";
(603) "%% K"" "
(604) " '
(605) &"C "'"
(606) 9"
(607)
(608) "'
(609) "
(610) "#
(611) "& "
(612) " " " '&
(613) " '"&/
(614) "RB '
(615) "
(616) '". /K"+ 86T" &"0"& "'& K" " &)
(617) "&/>"" .
(618) %)K "
(619) " "
(620) # " &
(621) ' " " V " V&K "
(622) &
(623) %0 " ) "
(624) %
(625) " I" C 0"R V&K"+ 8TK"" "% ".
(626) '" "
(627)
(628) " ," " " 9"'""D
(629) "(
(630) ' " "A " Z " "&
(631) %"C9
(632) &"'C
(633) "
(634) #
(635) ) ";
(636) "%% K"& "&/&/ "
(637) " "
(638) "'
(639) &" ' "& " &/ "'
(640) ""& " ' "R(
(641) ' "
(642) '"Z K"+ 4T"C "I" "%
(643) "' "
(644)
(645) " ," 0C;> /"B 'K"G'"B #" " ";V
(646) ")
(647) "
(648) " "'" "0"
(649) "
(650) "
(651) 9
(652) "'" "&/&/" "C9
(653) &" " "%
(654) &
(655) "'" "
(656) #
(657) ) " ' "&/>"" %)" ' "RB '"
(658) '"B K"4,,4T . "
(659) #
(660) ) " " "'& "&/>"'"
(661) " )& "& ""
(662) #"R . " " K"+ 6["Y/
(663) " " K"+ ["3 "
(664) '" >$D( K"+ ["
(665) " #" " K"4,,4T" "
(666) & "" " RB ' " " K " + ?K " +. [ " Y
(667) &V "
(668) ' " V&K " +. T " "
(669) %
(670) " &/> " 2/ " '
(671) ". 2/ & "R;V
(672) " " K"+ 8T" "'
(673) """ % & %"R " " K"4,,?["A
(674) " " K"4,,T"
(675) "02
(676) " '"&
(677) " ##)"0C" " "RD#
(678)
(679) " " K"4,+4T"2 "
(680) &/ "
(681) #
(682) 0 "
(683) " " "'
(684) ""& "& " "
(685) "'"& 9" # "' "'"
(686) &" & "RD " " K"4,,[": ( " " K"4,,TK"'
(687) ""& 9"% "RAV"
(688) '" K"+ +TK" 2 (#' "RD
(689) " " K"4,,4TK"2/( / "RH
(690) #"
(691) '"A K"+ 8["Y V " " K"4,,T"
(692) & ""
(693) & 9"R;/
(694)
(695) #"
(696) '"Y
(697)
(698) K"4,,6["B 'K"4,,?["Y
(699) &V"
(700) '" V&K"4,,+T" \ 0C "
(701) " K " "
(702) #
(703) ) " ' " '
(704) " & " & K " 0C "
(705) C " " "
(706) " 0
(707) K" " "I"
(708) " 9" & " "%
(709) K" " "'"& "
(710)
(711) "'"&/&/K" "&
(712) "
(713) &
(714) %0" C && '"I" '"0" "'9" "
(715) & 9"'"
(716) #
(717) ) " ' "
(718) "C/ & " """ % & %"
(719) &
(720)
(721) "C/ & K" "& " &/ " "
(722) "'
(723) " ">
(724) " "#
(725) "'"#( "'
(726) $. "RG#"+T" " "
(727) " "'"& "& " &/ "
(728) "#" "
(729) "]"'" "& &/" #
(730) " "'
(731)
(732) "' "& "#
(733) "
(734) $% "3 "
(735) #
(736) ) " "0
(737) "I"" " "'"& " &/ "0" %)
(738) " "
(739) "'" ">
(740) " $
(741) & "' "
(742) & " $. 9" "& " &/ "
(743) "'
(744)
(745) "' "
(746) "0"#
(747) " "
(748) "
(749) #"'
(750) &"
(751) #"'"%9" "# "@ 0C "" % & %"RG#"+T. 3 "
(752) #
(753) ) " ' K"]"
(754) "'
(755) ""& "'" %)" "%
(756) " 0"'
(757) " " & "'C
(758) #" "'" K" "
(759) "'
(760) "C/ & "RB " " K"4,+4T"0"'
(761) """ % & %"R3 >
(762) "
(763) '"3' K"4,++T";"&
(764) "
(765) " & "'" & " && " '
(766) " ' " & " & " ' " C
(767) # " " ' " " "
(768) " .
(769) " C
(770) #
(771) " ' "
(772)
(773) " '
(774) "
(775) " " 9
(776) " & 9 " :
(777) " " C " " &)
(778) "
(779) " I " "
(780) #
(781) ) " '
(782) " " ( ) " % & % " " I "
(783) " < " '
(784) " C
(785) #" % & %"3 " "'"' &
(786) "
(787) "'
(788) &"C #
(789)
(790) "
(791) $%
(792) &
(793)
(794) " '" ( )" % & %K" " & "
(795) "
(796) " &"C9
(797) &"
(798) " " "
(799) & 9"
(800). ?.
(801) '" ". "&
(802) &
(803)
(804) " "
(805) #
(806) ) " % & " "
(807) "&
(808) "'
(809) " "%
(810) &
(811) " % & " 3 " . "
(812) "
(813) " " " % " ' " 6" & " '
(814) " 4"
(815) " " "
(816) " K " 0" '&
(817) ""<"'" "
(818) #
(819) ) "'
(820) "C
(821) #"& %K"C&
(822) "'" "
(823) '
(824)
(825) " '
(826) " " '
(827) "'" "
(828) #
(829) ) "
(830) & "I"C
(831) #"& %" "
(832) %
(833) "" & "'"
(834) "' "& & "'C
(835) #"& %" "&
(836) "'"& "&
(837) "3C
(838) "' & " . " "
(839) "' & ". . 8. .
(840) !"#$%&'" $!.
(841) ($#)! *" $!!"$+$,-$!'" $!!.//.%&*0*".+.$/-'" 3 "&
(842) " % & " "% " &/ 0
(843) "
(844) " " " "'2 'K"
(845) " '
(846) K &"&/0" K" ".
(847) '" "& "
(848) "#E&"I" "
(849) " " '
(850) " " " "%9" " "& " % & % "
(851) " " "
(852)
(853) "
(854) "'"2 /" "%
(855) '"'" " & "
(856) "3 "&
(857)
(858)
(859) &" '
(860) $& " "
(861) " "
(862) '&
(863) "'" #"&/0"
(864) "
(865) " #"& 0 ;
(866) K" " " 9
(867) "' "
(868)
(869) "
(870) K"" #" % & %" " &/
(871) "@ 02 "" % & %K"" "& "'"C
(872) %
(873) " % & ""
(874) "'"" % & %K" " 9
(875) "' "
(876)
(877) "
(878) "
(879) "% " (
(880) " &" "'
(881) ' "' "
(882)
(883) " K" "& " " C "'
(884) """ % & %"0"C
(885) %
(886) " % & " " "
(887) ""
(888) "
(889) "'C] ".
(890) " 9"&
Documents relatifs
رشع عساتلا نرقلا ةفاقث عوزن تاهجوتلا هذه ززع دقو لىإ و ةيعيبطلا رهاوظلا يرسفت زواتج ناسنلإا ةي يم ايرسفت ت حضاو ليمو ايقيزيفا لىإ ةيعوضولما تايرسفتلا
Multiplier un nombre par une fraction revient parfois à augmenter ce
Tập sách này không đầy đủ, vì ngài google có bao giờ cho đọc trọn vẹn một tác phẩm nào đâu. Hi vọng bà con ta khai thác có hiệu quả tập
Abstract — A Short History of the Last 15 year's Quest for IT Interoperability in the Petroleum E&P Industry — This paper gives a personal view on the drivers of the
experiments which showed that a visual stimulus previously asso- ciated with an odorant is able to activate primary olfactory cortical regions ( Gottfried et al., 2002, 2004 ) in
Using the success rate as another index of learning, we also found that both spaced and massed conditioned groups performed differently from their control groups (group effect;
ﻣﻠﺨﺺ ﺗﻌﺘﺒﺮ ﺻﻨﺎدﯾﻖ اﻻﺳﺘﺜﻤﺎر ﻣﻦ ﺑﯿﻦ أھﻢ اﻟﻮﺳﺎﺋﻞ اﻻﺳﺘﺜﻤﺎرﯾﺔ ﻓﻲ ﻣﺠﺎل ﺗﻤﻮﯾﻞ اﻻﻗﺘﺼﺎد ﺑﺎﻟﻤﻮارد اﻟﻤﺎﻟﯿﺔ اﻟﻼزﻣﺔ ﻹﻋﺎدة ﺑﻌﺚ اﻟﻨﺸﺎط اﻻﻗﺘﺼﺎدي ﺣﯿﺚ ﺗﻌﺘﺒﺮ أداة
For example, a steel sheet with a piece of plastic tape (ε = 0.95) fixed to it have the same measured temperature. If the steel and the plastic are warmed to a higher