Дрёмов С.В., Сёмин И.Р. - Измененные состояния сознания

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.-8A1/4.+ 2/D+.-8-0, ,5<+1+4./2 8/6/,/.+25, 8-.9+6/2, /3-*5./*/2 + 42;CG258+4A 4 2/C- D-04.2+-, /D1//7*5C1G: *5CD*5I+.-8-0 15 C*+.-8A1G0, 48@:/2/0, 2-4.+7@8;*1G0 5158+C5./*G 248-D4.2+- /=*51+9-1+; 3/D2+I1/4.+ ( /*7/2, -7-D-2, 1975; -7-D-2, 1978). -,5 3*+4@.- 4.2/2585 3/D43@D1/ 2 +C@9-1++ "1-/7G91G: 34+:+9-46+: 4/4./;1+0", "4+1D*/,5 34+:/E,/F+/158A- 1/=/ 153*;I-1+;" @ 3/8;*1+6/2, 8-.9+6/2, 6/4,/152./2, *57/.5H?+: 2 E64.*-,58A1G: @48/2+;:

4@?-4.2/251+; ( @C1-F/2, -7-D-2, 1972; -7-D-2, 1979, 1980, 1989; /*/8-16/, 1978). 5958/ E643-*+,-1.58A1G: +448-D/251+0 4-14/*1/0 +C/8;F++, D52<+: C159+.-8A1G0 ,5.-*+58 6 +C@9-- 1+H , 2 /.-9-4.2-11/0 15@6- D/8I1/ 7G.A /.1-4-1/ 6 1578HD-1+;, . . 528/25 15D I+2/.- 1G,+ 2 "75<1- ,/8951+;". 3/48-D@H?-, 3*+ *57/.- 4 8HDA,+ 3*+,-1;8+4A *5C1//7*5C1G- ,--

./D+9-46+- 3*+-,G 78/6+*/251+; /*=51/2 9@24.2 /. *5CD*5I+.-8-0 238/.A D/ 3/81/=/ +468H9-- 1+; 5BB-*-1.1G: 4.+,@8/2 21-<1-0 4*-DG. 448-D/258/4A 28+;1+- 15 34+:+9-46@H D-;.-8A- 1/4.A 656 4-14/*1/0, .56 + 4/F+58A1/0 +C/8;F++ ( 51?+6/2, ./8;*/2, 1966; @C1-F/2, -7-D-2, 1972; -7-D-2, 1978). 9-1A 3/D*/71/ .-,5 /42-?5854A 2 5158+C- 34+:+9-46+: 15*@<-1+0 3/48- 3*+,-1-1+; ,-4658+15, 5,B-.5,+15, -25 + D*@=+: 34+:/56.+21G: 4*-D4.2 ( /19-246+0, 1941; +/:+,+; 34+:/C/2, 1963; Bercel at al., 1956; !+1/=-1/25, 1970; +8A<.-01, 3+256, 1971; Hermle et al., 1992; + D*.).

85=/D5*; @65C511G, *57/.5, .-,5 3/3/81+854A 48-D@H?+,+ D511G,+. 5 /41/2- /.9-./2 +43G.@-,G: + 45,/1578HD-1+0 2 /3G.5: 3/ 4-14/*1/0 +C/8;F++, 2 B5*,56/8/=+9-46+ 2GC2511G: 34+:/C5: 7G85 4/4.528-15 +49-*3G25H?5; B-1/,-1/8/=+9-465; 65*.+15 +C,-1-1+0 4/C151+;.

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

11

.-8A1/4.A... /I1/ 465C5.A, 9./ 3*+ 158+9++ 3*52+8A1/=/ /.1/<-1+; 6 *5CG=*G25H?+,4; +C,-1-- 1+;, 3*/+C/<8/ +C,-1-1+- 1/*,58A1/=/ 4//.1/<-1+; 34+:+C,/2, @.*5.+854A +: 48+.1/4.A 2 -D+- 1/0 56.+21/0 @4.51/2615 21-<1+0 ,+*, 42/04.2-115; 1/*,58A1/,@ 4/4./;1+H 4/C151+;. /E./- ,@ ,1- 3/65C58/4A 3*52+8A1G, 3*+,-1-11/- 2G*5I-1+- "E643-*+,-1.58A1/- *54?-38-1+- 8+9-

1/4.+"" ( /19-246+0, 1941, 4.59-61).

3+4G25; /D+1 +C 25*+51./2 +C,-1-1+; 4/C151+;, . . /19-246+0 @65C58 15+7/8-- 25I1G- B-- 1/,-1/8/=+9-46+- 9-*.G, 42/04.2-11G- 7/8A<+14.2@ D*@=+: : +C,-1-1+- 9@24.25 56.+21/4.+, 15*@<-1+- 2/43*+;.+; 4/74.2-11/=/ -D+14.25, 3*+4@.4.2+- 5BB-6.+21G: 15*@<-1+0, +1.*/2-*-

.+*/25115; @4.51/265 4/C151+; + @D58-1+- /. /6*@I5H?-=/ ,+*5 4 +C,-1-1+-, /.1/<-1+; 6 1-,@.

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

1)E,/F+/158A15; 153*;I-11/4.A 3/D=/./2+.-8A1/=/ E.535;

2)3/8+,/*B1G-, 1- +43G.511G- *51-- 3-*-I+251+; 4 <+*/6+, 5BB-6.+21G, D+535C/1/, /. E0- B/*++ D/ 4/4./;1+; @I545 15 E.53- /4.*G: 34+:+9-46+: *-56F+0, 8/,65 4.5*/=/ + @4.51/28-1+- 1/2/=/ D+15,+9-46/=/ 4.-*-/.+35;

3)*-D@6F+; 34+:/35./8/=+9-46+: B-1/,-1/2 15 E.53- /.1/4+.-8A1/0 34+:+9-46/0 5D53.5F++, /4@?-4.28-1+- 34+:+9-46/0 D-;.-8A1/4.+ 15 @*/21- B@16F+/158A1/=/ /3.+,@,5;

4)3-*-8/,1G0 E.53 34+:+9-46/0 153*;I-11/4.+ 4 B-1/,-15,+, 4/4.528;H?+,+ +1D+2+D@58A1@H *-56F+H 15 +43G.G25-,G- 3-*-I+251+;;

5)3/815; *-5D53.5F+; 8+7/ 3*/.+2/3/8/I-11/- ;28-1+- - B/*,+*/251+- =8@7/6+: 34+:+9-46+: +C,-1-1+0 ( -7-D-2, 1980; @C1-F/2, -7-D-2, 1972).

*-D8/I-115; 68544+B+65F+; 2-4A,5 153/,+15-. D*@=@H, 4/4.528-11@H 3/ 1578HD-1+;, C5 , 2GC2511G, 34+:/./,+,-.+65,+. -56F++ 4/ 4./*/1G E,/F+/158A1/0 4B-*G + 3/2-D-1+; /3+4G- 25H.4; 2 .56/0 3/48-D/25.-8A1/4.+:

1./ -: 8H7G- +C,-1-1+; +4:/D1/=/ 34+:+9-46/=/ 4/4./;1+;.

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

1545, 3*/F-44 +C,-1-1+; 4/C151+;, ,G 23*52- =/2/*+.A / -=/ 5D53.+21/0 153*528-11/4.+.

511G- / *5C1//7*5C++ +C,-1-11G: 4/4./;1+0 4/C151+; + / ,1/I-4.2- 3*+9+1, 2GCG25H?+: E.+ 4/4./;1+;, @65CG25H. 15 ./, 9./ +C,-1-1+- 4/C151+; -4.A -4.-4.2-115; 5D53.+215; *-56F+;.

12

9+48- 3*+9+1 15CG258+4A "15*@<-1+- 4@./91/=/ *+.,5, B+C+9-46/- + @,4.2-11/- 3-*-15- 3*;I-1+-, C5.*@D1-115; 5D53.5F+;, D-C5-D53.5F+; + 3-*-5D53.5F+;, 56F-1.@5F+; 8+91/4.+ + ..3., 1-*-D6/ 4/9-.5H?+-4; 4 =8@7/6+,+ 34+:+9-46+,+ 3-*-I+251+;,+, +1B/*,5F+/11/0 1-/3*-D-- 8-11/4.AH, 4/F+58A1/-34+:/8/=+9-46+,+ /=*51+9-1+;,+, 4@7R-6.+21/ 1-*-58+C/2511G,+ @4.*-,- 8-1+;,+ + D*." ( 3+256, 1988, 4.140). */,- ./=/, 2/C1+61/2-1+- 7G8/ 42;C51/ 4 "5@./=-11/0

.*-1+*/26/0, .2/*9-46/0 *57/./0, 2G3/81-1+-, *+.@58/2. 1+, ,/I-. 3*+2/D+.A 2/CD-04.2+- 3*/,G<8-11G: + 7G./2G: :+,+9-46+ 2*-D1G: 2-?-4.2, 5 .56I- 3*+-, .56+: ,-D+65,-1./2, 656 3-3.+DG, 1//.*/3G, 4.+,@8;./*G, 15*6/.+6+, .*5162+8+C5./*G, 47/*G 1-6/./*G: .*52 + ..D." (.5, I-). 56 3+458 . @*=+1A/1 (Bourguignon, 1979, *.286), 5,-*+65146+, +1D-0F5, +C2-4.1/ 7/8-- 80 2+D/2 *54.-1+0, 2GCG25H?+: . *-D+ *+.@58A1/ 3*56.+6@-,G: 15CG258+4A , 2GC2511G- *+.@58A1G,+ .51F5,+, 3*+1;.+-, /3*-D-8-11G: 3/C, *-=@8;F+-0 DG:51+;, 3/4./;11/ 3/2./*;H- ?+,+4; ,/8+.25,+, 546-.+C,/, + ..3.

+535C/1 4+.@5F+0, 3*+2/D;?+: 6 , 2-4A,5 <+*/6. ./ ,/=@. 7G.A 4F-1G /7GD-11/0 I+C1+, "+C2-4.1G- 3/ D/45D1G, /74./;.-8A4.25,, 6/=D5 "=/2/*;. 3/D *@6@" ( /*7/2, -7-D-2, 1975, 4.88), 78+C6+- 6 1+, 4+.@5F++ /D1/2*-,-11/=/ 2G3/81-1+; D2@: 4:/D1G: 3/ :5*56.-*@ 2+D/2 D-;.-8A1/- 4.+ (9.-1+- 8-6F++ + /D1/2*-,-11/- C5=8;DG251+- 2 .-64., D-685,5F+; */8+ + 2G48@<+251+- 3/D465C/6 4@B8-*5). ./ + 4+.@5F++ +1.+,1/0 78+C/4.+ (Swartz, 1994), + 3*/48@<+251+- 34+:/D-- 8+9-46/0 ,@CG6+ 4/ C159+.-8A1G, *+.,+9-46+, 6/,3/1-1./, (Biasutti, 1990; Lyttle, Montagn, 1992), + C51;.+; 5E*/7+6/0 (Estivill, 1995). ./ .56I- @28-9-1+;, 3*+1+,5H?+- :5*56.-* 5DD+6./2 [5].*/1+9-46+- + /4.*G- 4/,5.+9-46+- C57/8-251+; 4 +C,-1-1+-, +1.*/F-3F++ + @4+8-11G, 2/4- 3*+;.+-, 2+4F-*58A1G: 4+=158/2 .56I- 1-+C,-11/ 4/3*/2/ID5H.4; (Kokoszka, 1992). H7G- /74./;.-8A4.25, 42;C511G- 4 @=*/C/0 D8; I+C1+, 656-./: 3*GI6+ 4 35*5<H./, (Cancio, 1991) + ,1/- =+- 4+.@5F++ E64.*-,58A1/=/ .+35 2 3*/B-44+/158A1/0 D-;.-8A1/4.+ 8-.9+6/2, ,/*;6/2, 3/8;*1+- 6/2, 6/4,/152./2, 65465D-*/2 + D*., ,/=@. *544,5.*+25.A4; 2 659-4.2- 4.+,@8/2 .

*+9+1G, 2GCG25H?+- , /7R-D+1;-. ,-:51+C, +: *-58+C5F++. / 24-: 48@95;: +,--. ,-4./ 56.+25F+; 3*52/=/ 3/8@<5*+; (Bick, 1989; Brauchli at al., 1995) + +C,-1-1+; 2 2/43*+;.++ 4-14/*- 1G: 4+=158/2. 56+- +C,-1-1+; 154.@35H. 8+7/ 248-D4.2+- 2+D/+C,-1-1+; (@4+8-1+;, 3/81/=/ +8+ 954.+91/=/ 78/6+*/251+;) 21@.*-11+: + 21-<1+: 4.+,@8/2, 8+7/ /3/4*-D/2511/ C5 49-. 28+;- 1+; 15 1-0*/,-D+5./*G, 1-0*/158A1G- 4+1534G 4 @46/*-1+-, +8+ C5,-D8-1+-, 3-*-D59+ 4+=15- 8/2. *-C@8A.5.- @.*59+25-.4; 43/4/71/4.A 6 5158+C@ +C,-1-11G: 4-14/*1G: 4+=158/2. 48+ I- =/2/*+.A 1- / 1-0*/B+C+/8/=+9-46+: 3*-D3/4G865: 533-*F-3F++, 5 / 4/C151++, ./ +C,-1-1+- -=/ +4:/D1/=/ 4/4./;1+; 2/C1+65-. 2/ 24-: 48@95;: 15*@<-1+; 43/4/71/4.+ 9-8/2-65 *-=@8+*/25.A 2C5+,/D-04.2+- 4 21-<1+, ,+*/, 3*-I1+, 43/4/7/,.

*-D4.528-1+- / *543*/4.*51-11/4.+ 4+.@5F+0, +1+F++*@H?+: , 3/C2/8;-. 465C5.A, 9./ B56-

.+9-46+ 24; I+C1A -4.A 8+<A 15 2*-,; 3*-*G25H?5;4; D+15,+65 3/D/71G: 4+.@5F+0, + 1/*,58A1/ B@16F+/1+*@H?-- 4/C151+- - E./ 4/C151+-, 43/4/71/- +C,-1;.A4;. 56 C5,-.+8 . -08 (Well, 1972, *.17), "I-851+- 3-*+/D+9-46+ +C,-1;.A 4/4./;1+- 4/C151+; ;28;-.4; 1/*,58A1/0 2*/ID-1- 1/0 468/11/4.AH, 5158/=+91/0 =/8/D@ +8+ 4-64@58A1/,@ 28-9-1+H". 56 +6 + 4/52./*G (McPeake et al., 1991)/3+4G25H. +C,-1-1+; 4/C151+; 656 /41/21/0 9-8/2-9-46+0 ,/.+2. *+1+,5; E.+ 2G- 465CG251+;, ,G ,/I-, C568H9+.A, 9./ 1-. 656/=/-./ *5C + 1524-=D5 B+64+*/2511/=/ /3.+,58A1/=/ 4/4./;1+;, 5 +,--.4; F-8G0 157/* 4/4./;1+0 4/C151+;, 5D-625.1G: D8; *-<-1+; *5C1G: C5D59.*-D4.528-1+- / ,1/I-4.2- 4/4./;1+0 4/C151+;, /7-43-9+25H?+: 2GI+25-,/4.A 9-8/2-65 2 *5C1G: @48/2+;: 4@?-4.2/251+;, ;28;-.4; @D/28-.2/*+.-8A1G, /7R;41-1+-, 2G4/6/0 857+8A1/4.+ 9-8/2-9-46+: 5D53.5F+/11G: 2/C,/I1/4.-0 + 4//.1/4+.4; 4 C5,-951+-, 51=8+046/=/ 51.*/3/8/=5

13

.!/@8+: " /, 9./ ,/I1/ 15C25.A @1+658A1/4.AH 9-8/2-65, -4.A *-C@8A.5. 1- 656/0-./ /D1/0 5D53-

.5F++, 5 6/,7+15F+; F-8/=/ *;D5 5D53.5F++" (!/@8+, 1990, 4.122).

56+, /7*5C/,, ,G D/8I1G C5,-.+.A, 9./ +C,-1-1+- 4/C151+; 3*-D4.528;-. 4/7/0 6/14.+.@.+2- 1@H :5*56.-*+4.+6@ 4/C151+;. 85=/D5*; +C,-1-1+;, 4/C151+; /4@?-4.28;H.4; -=/ /41/21G- B@16F++: /*+-1.+*/265 + 3*+43/4/78-1+-. /7G91G: @48/2+;: 4/:*51-1+- *521/2-4+; 4/ 4*-D/0 1- .*-7@-. +C,-1-1+; 4/C151+;, +7/ /4@?-4.28;-.4; 2 *5,65: +4:/D1/=/ 4/4./;1+;. /=D5 I- 9-- 8/2-6 D/8I-1 4/:*51+.A 2C5+,/D-04.2+- 4 /6*@I-1+-, 2 1/2G: + /4/7-11/ 2 1-785=/3*+;.1G: @4- 8/2+;:, 154.@35-. +C,-1-1+- 4/C151+;.

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

1.3.

G*5I-1+-, "4/2*-,-11G0 2C=8;D" /7/C159-1/ 5D-625.1/- 3*/78-,- 4.*-,8-1+- /3*-D-- 8+.A 3/C+F+H D8; F-8/4.1/=/ 34+:/8/=/-34+:+5.*+9-46/=/ + B+8/4/B46/=/ *544,/.*-1+; 3/D1+- ,5-,G: 2/3*/4/2. 5;265 15 F-8/4.1/4.A 3/D:/D5 6 3*/78-,- 3*-D3/85=5-. +C@9-1+- -?- /D1/=/ 543-6.5 5D53.+21/4.+ , 5 +,-11/, */8+ +C,-1-1+0 4/C151+; 2 4/:*51-1++ 21@.*-11-=/ *521/- 2-4+; +1D+2+D@@,5.

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

./ -4.A /3*-D-8-115; 4+4.-,5 8-64+6/-=*5,,5.+9-46+: 4*-D4.2. 15; E.+ :5*56.-*1G- 3*+C156+ ;CG6/2/0 @3/*;D/9-11/4.+ 4//.2-.4.2@H?-=/ @*/21;, +43/8AC@; 43-F+B+9-46+0 ";CG6", 2/C,/I1/ @4.51/2+.A 3*/D@6.+21/- *-9-2/- /7?-1+- 4 9-8/2-6/, 2 .

14

*+,-1-1+- 8+1=2+4.+9-46/=/ .-4.5 2 :/D- 3/48-D/25.-8A1/0 D+44/8HF++ 4/C151+; (41+I-1+; @*/21;, 3/ I-64/1@) 3/C2/8+8/ /7R-6.+21/ *-6/14.*@+*/25.A E.53G, 3*/0D-11G- 2 *5C2+.++ *@4- 46/=/ ;CG65 C5 3/48-D1+- 4./8-.+;. GD-8-11G- 2 5*:5+91G- 543-6.G ;CG6/2/0 4.*@6.@*G 3/C2/8+8+ 4/3/4.52+.A ;CG6 +C,-1-11G: 4/4./;1+0 4/C151+; 4 7*511/0 *-9AH, 4 *-8+6./2G,+ 45- 6*58A1G,+ ;CG65,+ ( 3+256, 1986, 4.14,37).

/74.2-11G- /*+=+158A1G- +448-D/251+; . . 3+256 3*-D25*;-. /7?-0 351/*5,/0 +448-D/25-

.-8A46+: 3*/=*5,, 3/ 3*/78-,- . " C@9-1+- , - 3+<-. . . 3+256 (1986, 4.9), - 3*/2/D+- ,/- 7/8A<+, 6/8+9-4.2/, 15@91G: 6/88-6.+2/2, 4.58/ 2 3/48-D1-- 2*-,; 3*+/*+.-.1G, 153*52- 8-1+-, 2 +448-D/251++ ,G<8-1+;". 1973=. +CD5-.4; ,-ID@15*/D1G0 ")@*158 +C,-1-11G: 4/- 4./;1+0 4/C151+;", 7+78+/=*5B+9-46+- 3-*-91+ 1549+.G25H. .G4;9+ .*@D/2 3/ E./0 .-,-, 3/8@- 9+2<-0 /4/7/- *5C2+.+- 3/48- ,-ID@15*/D1/=/ 4+,3/C+@,5 3/ +C,-1-11G, 4/4./;1+;, 4/C151+;

( /*/1./, 1978).

/2*-,-11G0 @*/2-1A *5C*57/.6+ 3*/78-,G 3*-D4.528-1 .*-,; 15+7/8-- +C2-4.1G,+ .-/- *+;,+. '2-0F5*46+0 34+:/8/= + 34+:+5.* . +..*+: (1986), 2G7*52 45,@H 1-3*/.+2/*-9+2@H ,/- D-8A 4/C151+; - 6*@=, 248-D C5 . @1D./, *543/8/I+8 2-4A 43-6.* 4/C151+; .56+, /7*5C/,, 9./ 2 F-1.*- 3/,-?5-.4; 1/*,58A1/- 7/D*4.2@H?-- 4/C151+-, 5 3/ /6*@I1/4.+ - 7-44/C15.-8A1/-, 4/- 3/4.52+,/- 4 2G68H9-1+-, 4/C151+;. -ID@ 1+,+ 15 *5C1G: *5D+@45: *543/8/I-1G 3-*-:/D1G- +C,-1-11G- 4/4./;1+; 4/C151+;, 4/3/4.52+,G- ,-ID@ 4/7/0 3*+ @48/2++ *521/0 @D58-11/4.+ /. F-1.*5 - .56 15CG25-,G- 4,-I1G- 4/4./;1+; 4/C151+;. ./ D5-. /41/251+- =/2/*+.A, 9./

+,-H. /7?+- 4.57+8A1G- :5*56.-*+4.+6+, 1-C52+4+,G- /. 2GCG25H?-0 3*+9+1G (Dittrich etal., 1986). 5115; ,/D-8A, +8+ .-/*+; 4,-I1G: 4/4./;1+0 4/C151+;, /785D5-. /3*-D-8-11/0 *5C*-- <5H?-0 43/4/71/4.AH, 3*/;41;; B-1/,-1/8/=+9-46/- 4:/D4.2/ 4,-I1G: 4/4./;1+0 4/C151+;, 7@DA ./ B5*,56/8/=+9-46+ 2GC2511G-, 2/C1+6<+- 2 @48/2+;: 4-14/*1/0 (4/F+58A1/0) +C/8;F++ +8+ 2 34+:/C-.

+15,+65 + 153*528-11/4.A +C,-1-1+0 4/C151+; 15:/D;. 2G*5I-1+- 2 .-/*++ 1-3*-*G21G: 4/4./;1+0 4/C151+; . 5*.+1D-085 (Martindail, 1981). 3+4G25; /41/21/- +C,-*-1+-, 2D/8A 6/./- */=/ +C,-1;-.4; 4/C151+-, /1 /.4.5+25-. .*5D+F+/11@H D8; 5,-*+65146/0 15@6+ "/D1/,-*1@H"

4:-,@ 4/C151+;. 15 C5D5-.4; 8+1+-0 *-=*-44++, 2D/8A 6/./*/0 3*/+4:/D+. D+44/8HF+; 4/C151+; 3/ 153*528-1+H 6 -=/ 3-*2+91G, D*-21+, 4/4./;1+;,. *@=+,+ 48/25,+, +,--.4; 6/1.+1@@, 1-- 3*-*G21G: 4/4./;1+0 4/C151+;, 3*-D4.52+,G0 2 2+D- /4+, 3*/1+CG25H?-0 4.*@6.@*@ 4/C151+;. 1+I1-0 954.+ E./0 /4+ 15:/D;.4; 34+:/CG, 2G<- ,/=@. +D.+ 41G, -?- 2G<- - @*/2-1A, D/4.+=5-- ,G0 34+:/B5*,56/8/=+-0, D58A<- - 1-2*/CG + ..D. /I58@0, /41/21/-, 9./ D5-. 3*-D8/I-115;5*.+1D-08/, 4:-,5, 2/ ,1/=/, 3-*-68+65H?5;4; 4 34+:/5158+.+9-46+,+ 6/14.*@6.5,+, .-/*+-0 -D+1/=/ 34+:/C5, - E./ 45,/- /7?-- 3*-D4.528-1+- /7 +C,-1-1+;: 4/C151+; 2 6/1.-64.- /3*-D-- 8-11/0 4.*5.+B+65F++. ,-4.- 4 .-, 574/8H.1/- /./ID-4.28-1+- *5C1G: 3/ 3*/+4:/ID-1+H 4/- 4./;1+0 4/C151+; 2 4//.2-.4.2++ 4 -D+14.2-11G, 6*+.-*+-, - 3/8/I-1+-, 15 /4+ *-=*-44++ - 2*;D 8+ 2/C,/I1/. *-D4.528-1+; 5*.+1D-085 3/ E./,@ 2/3*/4@ 2/ ,1/=/, D/3/81;H.4; + 6/**-6.+- *@H.4; D*@=+, 2G<-@3/,;1@.G, 5,-*+65146+, +448-D/25.-8-, - . 5*./,, 6/1F-3F+; 6/./*/=/ 1@ID5-.4; 2 /.D-8A1/, /74@ID-1++.

7C/* 1/2G: +448-D/25.-8A46+: 3/D:/D/2 6 7G8 7G 1-3/8/1 7-C /7*5?-1+; 6 .*5143-*4/- 158A1/,@ 153*528-1+H 2 34+:/8/=++, 2/7*52<-,@ 2 4-7; +D-+ 68544+9-46/=/ 34+:/5158+C5, H1=+- 5146/0 5158+.+9-46/0 34+:/8/=++, *50:51+514.25 + E6C+4.-1F+58A1/0 34+:/8/=++. *5143-*4/15- 8+4.465; 3*/=*5,,5 +C@9-1+; +C,-1-11G: 4/4./;1+0 4/C151+; 2/ ,1/=/, 58A.-*15.+215 7/8A- <+14.2@ 3/8/I-1+0 .*5D+F+/11/0 34+:+5.*++ + 565D-,+9-46/0 34+:/8/=++. 24- I- E./ 153*52-

15

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

/2/*; / .*5143-*4/158A1/, 153*528-1++ 34+:/8/=++ + 34+:+5.*++ 2 42;C+ 4 3*/78-,/0 , 3*-ID- 24-=/ 48-D@-. /.,-.+.A *5C*57/.511G- CD-4A 25*+51.G 4.*5.+B+65F++ 4/C151+;. 56/2G, 2 954.1/4.+, "65*./=*5B+; 4/C151+;" . */B5, <658G . 5*.5, "43-6.* 4/C151+;" . +87-*5. *-D- 8/I-11G- .*5143-*4/158+4.5,+ "68544+B+65F++", 152-*1/-, ,/I1/ 49+.5.A @./91-1+-, 2C=8;D/2.&1=5 15 4.*/-1+- +1D+2+D@58A1/=/ 4/C151+;. /.; 3/D/71G- 4:-,G D5H. 8+<A 4.*@6.@*1/-, 4.5.+9-46/- 3*-D4.528-1+- / 4/C151++, /1+ /.*5I5H. =8521G- 8+1++ + *5C8+9+;, 3/C2/8;H. C5 /=- */,1G, ,1/I-4.2/, 4/D-*I51+0 @2+D-.A F-8/4.1@H 65*.+1@ 4/4./;1+0 4/C151+;. 7?-- + F-1- 1/- 2 1+: - E./ @65C51+; 15 3*+4@.4.2+- 2 4/C151++ 1-;21G:, .*514+1D+2+D@58A1G: @*/21-0, 3*/- ;28-1+; 6/./*G: /B+F+58A15; 34+:+5.*+; 2 7/8A<+14.2- 48@95-2 *54F-1+25-. 656 34+:/35./8/- =+H.

.D-8A1G0 +1.-*-4 3*-D4.528;H. 2/CC*-1+; 15 +-*5*:+9-46@H 4.*@6.@*@ 4/C151+; . 5*.5. 4/- 7-11/4.A -=/ .-/*++ 2 ./,, 9./ 2GD-8-1+- <658, @3/*;D/9+25H?+: 4/4./;1+; 4/C151+;, /1 3*/2/- D+. 3/ 564+/8/=+9-46/,@ 3*+1F+3@. . 5*. (1994, 4.210-211) /.,-95-., 9./ /3*-D-8-1+; "7/8-- 2G4/6/=/ + 7/8-- 1+C6/=/" @*/21-0 4/C151+; C52+4;. /. ,+*/2/CC*-1+; 9-8/2-65 + F-11/4.1/0 /*+-1.5F++ 6@8A.@*G 2 F-8/,. 56, 153*+,-*, 2 C535D1/0 F-11/4.1/0 <6582G4<+, 3*+1;./ 49+-

.5.A 4/4./;1+- 3/81/0 *5F+/158A1/4.+. 4//.2-.4.2++ 4 E./0 <658/0 7/8-- 1+C6+,+ ;28;H.4;

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

. 5*. (.5, I-) 3+<-., 9./ *5F+/158A1/4.A, 3*-2/C1/4+,5; 2 -2*/3-046/0 <658- F-11/4.-0, ;2- 8;-.4; 1-D/4.+I+,G, ,+*5I/,. *+ E./, "/7G91/- 4/4./;1+- ;28;-.4; /665C+/158A1/ 1-2*/.+- 9-46+, 2 ./, 4,G48-, 9./ 2 1-, *5F+/158A1/4.A /65CG25-.4; 8+<A *5F+/158+C5F+-0 .-: 3*/F-4- 4/2, 6/./*G- /41/251G 15 7-44/C15.-8A1G: 3/7@ID-1+;: + E,/F+;:". */,- ./=/, "*5F+/158A1/4.A 2*;D 8+ 3*+<854A 3/ D@<- 7/8A<+14.2@ +C 154 - 2-DA 9-8/2-6 4 .56+, 4/C151+-, 7G8 7G 9-,-./ 2*/D- 6/,3AH.-*5".

65C51+-, 1-46/8A6+: F-11/4.1G: <658 . 5*. /7*5?5-. 21+,51+- 15 /.1/4+.-8A1/4.A 15@91G: 3*-D4.528-1+0, 3*+C1511G: 574/8H.1G,+. +D+,/, 3/85=51+- +4.+11G, /7R-6.+21/=/ 3*+ 1-D/4-

.5./91/, @9-.- 4@7R-6.+21/=/ F-11/4.1/-E.+9-46/=/ 543-6.5 -?- 1- =5*51.+*@-. 4.+1G. /4/-

16

7-11/4.+ E./ 56.@58A1/ 2 D+4F+38+15:, C51;.G: /7R-6.+25F+-0 4@7R-6.+21/=/, 2 15@65:, =D- /7R- -6.+21/4.A @.2-*ID5-.4; 8+<A 2 6/1.-64.- /3*-D-8-11/0 35*5D+=,G, ,-./D/8/=+9-46+- /41/25- 1+; 6/./*/0 24-=D5 4@7R-6.+21G.

+4./*++ 15@6+ +C2-4.1G 48@95+, 6/=D5 *5F+/158A1/4.A, 2G*5I5H?5; /7R-6.+21/4.A, 4,-<+25- 854A 4 *5F+/158+C5F+;,+. 4+:+5.*+;, 2 954.1/4.+, +C/7+8@-. 3*+,-*5,+, 6/=D5 *5F+/158+C5F++ @9-1/=/, 3*+C1511G- 15+7/8-- @D591G,+ 15 /3*-D-8-11/, E.53- 3/C151+; 2 6/16*-.1G0 +4./*+- 9-46+0 /.*-C/6, 3*-2*5?5854A E3+=/15,+ 2 +D-58 *5F+/158A1/4.+, D/=,5.+C+*/258+4A + 3/9+.5- 8+4A +4.+1/0 2 3/48-D1-0 +14.51F++.

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17

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18

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19

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20