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´ÄÞÎ²Î²Ü Ð²Ø²Èê²ð²Ü
ºðºì²Ü 2009
´áí³Ý¹³ÏáõÃÛáõÝ
1. Ü»ñ³ÍáõÃÛáõÝ 4
2. ¶ñ³Ï³Ý ³ÏݳñÏ 10
32
å³ÛÙ³ÝÝ»ñáõÙ
41
ë³Ï³í³ß³ñÅáõÃÛ³Ý å³ÛÙ³ÝÝ»ñáõÙ
41
ë³Ï³í³ß³ñÅáõÃÛ³Ý å³ÛÙ³ÝÝ»ñáõÙ
50
ë³Ï³í³ß³ñÅáõÃÛ³Ý å³ÛÙ³ÝÝ»ñáõÙ 53
ë³Ï³í³ß³ñÅáõÃÛ³Ý å³ÛÙ³ÝÝ»ñáõÙ 62
ë³Ï³í³ß³ñÅáõÃÛ³Ý å³ÛÙ³ÝÝ»ñáõÙ
74
8. øÝݳñÏáõÙ ¨ ³Ù÷á÷áõÙ
81
9. º½ñ³Ï³óáõÃÛáõÝÝ»ñ
92
¶ñ³Ï³ÝáõÃÛ³Ý ó³ÝÏ
94
2
3
ÀݹáõÝí³Í ѳå³íáõÙÝ»ñÇ ó³ÝÏ
¶²Î ¶³Ù³-³ÙÇݳϳñ³·³ÃÃáõ
AMPA α-amimo-3-hydroxy-5-methylisoxazole-4-propionate
NMDA N-methyl-d-aspartate
Øä Ù»ï³Õ³åñáï»ÇÝÝ»ñ
êú¸ ëáõå»ñûùëǹ¹ÇëÙáõï³½
زú ÙáÝá³ÙÇÝûùëǹ³½
4
1. ܺð²ÌàõÂÚàõÜ
²ß˳ï³ÝùÇ ³ñ¹Ç³Ï³ÝáõÃÛáõÝÁ
í³ñùÁ §÷³ËáõëïǦ ϳ٠§å³Ûù³ñǦ Ó¨áí, í×éáñáß ¹»ñ áõÝÇ ëÃñ»ëáñ ³½¹³ÏÇ
ٻ˳ÝǽÙÝ»ñáõÙ:
5
ëÃñ»ëáñ »Ý ¹³éÝáõÙ: ²ëí³ÍÇó Ñ»ï¨áõÙ ¿, áñ ë³Ï³í³ß³ñÅáõÃÛ³Ý å³ÛÙ³ÝÝ»ñáõÙ
(Schmahmann J.D., 1997, Martin L.A. et al, 2003, Vogel M.W. et al., 2007):
Ýí³½áõÙ (Сорохтин Г.Н., Могендович М.Р., 1972, Mitsushima et al., 1998, Langlet
et al., 1999, Ерицян Н.Б., 2005), áñÁ ÙϳݳÛÇÝ ³ÏïÇíáõÃÛ³Ý ë³Ñٳݳ÷³ÏÙ³Ý ³ÝÙÇç³Ï³Ý
6
ѻ勉ÝùÝ»ñÇó ¿: ØϳݳÛÇÝ åñáåñÇáó»åódzÛÇó ͳ·áÕ Ï»ÝïñáÝ³Ù»ï ³½¹³Ýß³ÝáõÙÁ
¿³Ï³Ý ¹»ñ áõÝÇ ÎÜÐ-Ç ³é³Ýó µ³ó³éáõÃÛ³Ý µáÉáñ µ³ÅÇÝÝ»ñÇ Ñ³Ù³ñÅ»ù ïáÝáõëÇ
׳ݳãáճϳÝ) ÇÙ³ëïÝ»ñáí:
7
³å³Ñ³ñÙ³ñíáճϳÝáõÃÛ³Ý ë³ÑÙ³ÝÝ»ñÁ: Üßí³Í ѳñó³¹ñáõÙÝ»ñÇ ¨ ÏÝ×ÇéÝ»ñÇ
ï»Õ³ß³ñÅ»ñÁ:
¹Ç³É¹»ÑÇ¹Ç ï»Õ³ß³ñÅ»ñÁ:
8
ÝÛ³ñ¹³ÛÇÝ ·áñÍáõÝ»áõÃÛ³Ý áñáß ï»Õ³ß³ñÅ»ñÇ ½³ñ·³óÙ³Ý Ù»Ë³ÝǽÙÝ»ñÇ
÷³Ã»ÃáõÙ: سëݳíáñ³å»ë.
ѳٳå³ï³ë˳Ý:
áõëáõóÙ³Ý ·áñÍÁÝóóÁ:
ûñ·³ÝǽٳÛÇÝ Ù³Ï³ñ¹³Ïáí:
Ý»ñ¹ñÙ³ÝÁ:
9
2. êï³óí³Í ïíÛ³ÉÝ»ñÁ ÏÉñ³óÝ»Ý ¨ ϳٵáÕç³Ï³Ý³óÝ»Ý í³ñù³ÛÇÝ µ³ñ¹
10
2. ¶ð²Î²Ü ²ÎܲðÎ
³ÏïÇíáõÃ³Ý ÏïñáõÏ Çç»óáõÙ ¿ ¹Çïí»É 20-ñ¹ ¹³ñÇ 60-³Ï³Ý Ãí³Ï³ÝÝ»ñÇÝ (Коваленко Е.А.,
11
ß³ñáõݳÏáõÙ ¿ ÙÝ³É µ³½áõÙ áõëáõÙݳëÇñáõÃÛáõÝÝ»ñÇ ³é³ñϳ: ì»ñçÇÝÇë ѳٳñ
12
·áÛ³óáõÃÛ³Ý ¨ ÑÇåáóɳÙá-ϻը³ÛÇÝ Ñ³Ù³Ï³ñ·Ç µÝ³Ï³Ý ËóÝÇãÝ ¿, áñÁ
å³Ñå³ÝáõÙ ¿ ÝÛ³ñ¹³ÛÇÝ Ñ³Ù³Ï³ñ·Ç ѳٳñÅ»ù ýáõÝÏóÇáÝ³É íÇ׳ÏÁ (Сорохтин Н.Н.,
1969, Canu M.H. et al., 2003): àõÕ»ÕÇÏÇ Ï³ñ·³íÇ׳ÏÝ ³Ûë ÇÙ³ëïáí ѳïáõÏ
Ñ»ï³ùñùñáõÃÛáõÝ ¿ Ý»ñϳ۳óÝáõÙ: ¸»é ³ÝóÛ³É ¹³ñÇ »ñ»ëáõݳϳÝÝ»ñÇÝ Ï³ï³ñ³Í
µ³½Ù³Í³í³É ѻﳽáïáõÃÛáõÝÝ»ñáõÙ È.². úñµ»ÉÇÝ óáõÛó ¿ ïí»É í»·»ï³ïÇí
ÝÛ³ñ¹³ÛÇÝ Ñ³Ù³Ï³ñ·Ç ϳåÁ áõÕ»ÕÇÏÇ Ñ»ï: àõÕ»ÕÇÏÁ å³ïÏ»ñ³óíáõÙ ¿ áñå»ë
ÑÇÙÝ³Ï³Ý é»ýÉ»Ïïáñ ³Õ»ÕÝ»ñÇ ³¹³åïÇí-ïñáýÇÏ µ³ñÓñ³·áõÛÝ Ï»ÝïñáÝ (Орбели Л.А.,
1940): àõÕ»ÕÇÏáõÙ ï»ÕÇ áõÝ»óáÕ ·áñͳéáõÛÃÝ»ñÇ Ñ»ï³·³
áõëáõÙݳëÇñáõÃÛáõÝÝ»ñÁ ѳëï³ïáõÙ ¨ Éñ³óÝáõÙ »Ý ³Ûë ѳñ³óáõÛóÁ:
àõÕ»ÕÇÏÇ í»ñ³µ»ñÛ³É ³í³Ý¹³Ï³Ý å³ïÏ»ñ³óáõÙÝ»ñÇ Ñ³Ù³Ó³ÛÝ ³ÛÝ »½³ÏÇ ¹»ñ
Glickstein M., 1992, Brodal P., Bjaalie J.G., 1997): ¸ñ³ Ù³ëÇÝ »Ý íϳÛáõÙ ³ÝóÛ³É
(Holmes G., 1932, Орбели Л.А., 1940, Albus J.S., 1971, Marr D., 1969, Eccles J.C., 1977, Fine
ß³ñÅáÕ³Ï³Ý ÑëÏáÕáõÃÛáõÝÇó (Ivry R.B., Baldo J.V., 1992, Kim S.G. et al., 1994,
Fiez J.A. et al., 1996, Allen G. et al., 1997, Timmann D, Daum I., 2007, Baldaçara
¹Çï³ñÏ»ÉÇë (Desmond J.E. et al., 1998, Schmahmann J.D., 2004, Chen S.H.,
13
áõëáõÙݳëÇñáõÃÛáõÝÝ»ñÁ, áñáÝù ÑÇÙù »Ý ï³ÉÇë »Ýó¹ñ»Éáõ ³Ûë ϳéáõÛóÝ»ñÇ
2005):
³ëáódzïÇí ϻը³ÛÇÝ ßñç³ÝÝ»ñÇ Ñ»ï (Dum and Strick, 2003; Middleton F.A. and
(Courchesne et al., 1994, Konczak J., Timmann D., 2007): àõÕ»ÕÇÏÇ ÏÇ볷ݹ»ñÇ
Strick P.L., 1994, 2001; Schmahmann J.D., 2004): àõÕ»ÕÇÏÇ áñ¹Ç Ñ»ïÇÝ µ³ÅÇÝÁ
áõÕ»ÕÇÏÇ áõñáõÛÝ ¹»ñÁ ¨ ï»ÕÁ ÎÜÐ ï³ñµ»ñ ·áñͳéáõÛÃÝ»ñáõÙ: Ü³Ë ¨ ³é³ç ËáëùÁ
Ýñ³ Ù³ëÇÝ ¿, áñ áõÕ»ÕÇÏÁ Ù»Í ÏÇ볷ݹ»ñÇ ÝÙ³Ý áõÝÇ Çñ ë»÷³Ï³Ý ϻըÁ (·áñß
14
(áõÕ»ÕÇÏÇ ëåÇï³Ï ÝÛáõÃÇ ËáñùáõÙ ï»Õ³¹ñí³Í »Ýóϻը³ÛÇÝ Ý»ÛñáݳÉ
Ï»Õ¨Ç Ñ»ï (Bloedel J.R., 1992): ì»ñç³å»ë, Ù³ñ¹áõ áõÕ»ÕÇÏÁ å³ñáõݳÏáõÙ ¿ ßáõñç
Ï»Õ¨Ç áÕç ÁÝóóùáí (Bloedel J.R., 1992), áñÁ ÑÇÙù ¿ ï³ÉÇë »Ýó¹ñ»Éáõ
15
Ïááñ¹ÇݳódzÛÇ, í³ñù³ÛÇÝ ³ÏïÇíáõÃÛ³Ý ¨ ÇÙ³ó³Ï³Ý-Ïá·ÝÇïÇí ·áñͳéáõÛÃÝ»ñáõÙ
Middleton F.A., Strick P.L. et al. 1998, 2000, Thach W.T., 1998, 2004, Mandolesi
L. et al., 2003, Zhang W., Linden D.J., 2006): ºÉÝ»Éáí ٻͳù³Ý³Ï áõ ɳÛÝ ÝÛ³ñ¹³-
(Middleton F.A., Strick P.L., 2000, Odeh F. et al., 2005)` ³é³ç ¿ ù³ßí»É ³ÛÝ Ã»½Á,
(Gandhi C.C. et al., 2000, Joyal et al., 2001): ì»ñçÇÝÇë ѳٳñ ³ÝÑñ³Å»ßï ¿
ßïÏáõÙ ÙïóÝ»Éáõ ϳñáճϳÝáõÃÛ³Ý ßÝáñÑÇí (Ito M., 2000, Martin L.A. et al.,
Leggio M.G. et al., 1999, Mandolesi L. et al., 2001, Rondi-Reig L. et al., 2002):
1998, Caston J. et al., 1998, Colombel C. et al., 2002, Linden D.J., 2003,
N. et al., 1989, Caston J. et al., 1995, Joyal C.C. et al., 1996, Colombel C. et al.,
16
ýáõÝÏódzÛÇ Ë³Ý·³ñáõÙÝ»ñÇ: àõÕ»ÕÇϳÛÇÝ Ë³Ý·³ñáõÙÝ»ñáí µÝ³Ï³Ý ¨ ÇݹáõÏóí³Í
³Ûë µ³ÅÝÇ ¹»ñÁ ß³ñÅáÕ³Ï³Ý Ñ³Ù³Ï³ñ·Ù³Ý ¨ áõëáõóÙ³Ý Ù»ç (Steinmetz J.E. et al.,
1993, Aiba A. et al., 1994, Chen C. et al., 1995, Hilber P., Caston J., 2001,
ùÝݳñÏÙ³Ý ³é³ñϳ: àñáß Ñ»ÕÇݳÏÝ»ñÇ Ï³ñÍÇùáí (Sacchetti B. et al., 2002, Song Y.-
ÁÝÏ×áõÙÁ (Ito M., 2000, Salin P.A. e al., 1996, Aizenman C.D. et al., 1996,
Zhang W., Linden D.J., 2006, Steinberg J.P., et al., 2006): ²Ûë µáÉáñ
(Marr D., 1969, Albus J.S., 1971, Eccles J.C., 1977): àõÕ»ÕÇÏÇ áõëáõóáõÙÝ
áõëáõóÙ³Ý Ù»Ë³ÝǽÙÁ (Ito M., 2000, 2001, 2006), áñÇ ÑÇÙùáõÙ ÁÝÏ³Í ¿
17
ѳÕáñ¹íáõÙ ¿ Ù³Ùé³ÝÙ³Ý ÝÛ³ñ¹³Ã»É»ñáí, ÇëÏ Ù³·ÉóáÕ Ã»É»ñÇ ÙÇçáóáí
ѻﳷ³ÛáõÙ ëï³ó³í µ³½áõÙ ³å³óáõÛóÝ»ñ (Ichise T. et al., 2000, King D.A. et al.,
2001, Koekkoek S.K.E. et al., 2003, Zhang W., Linden D.J., 2006, Steinberg J.P.,
³×Ç ·áñÍáÝÁ (Wang Y.T. et al., 2000, Matsuda S., et al., 2000, Ito M., 2000): ²Ûë
ÁÝÏ×Ù³Ý:
18
áõÝ»ó»É »Ý áõÕ»ÕÇÏÇ Ï»Õ¨áõÙ ºÄÀ ½³ñ·³óÙ³Ý ³Ýµ³í³ñ³ñáõÃÛáõÝ, áñáí ¨
í³ÛñÇ ÑëÏÇã ËÙµÇ ÙÏÝ»ñÁ (Chen C. et al., 1995): ÜßÛ³É ³ñ¹ÛáõÝùÝ»ñÁ ѳÙá½Çã
áõëáõó³ÝáÕ áõÕÇÝ ¿ (Thompson R.F. and Kim J.J., 1996, Thompson R.F. et al.,
1998):
¶üÂä-Ý ÉÇáíÇÝ µ³ó³Ï³ÛáõÙ ¿: ²ÛÝ Ý߳ݳϳÉÇ ù³Ý³Ïáí ³éϳ ¿ ´»ñ·Ù³ÝÇ ·ÉdzÛáõÙ, áñÁ
19
Ýϳñ³·ñí³Í §áõÕ»ÕÇϳÛÇÝ ÑÇßáÕáõÃÛ³ÝÝ ³éÝãíáÕ ·»Ý-1¦-Á (Cavallaro S. et al.,
2001): òáõÛó ¿ ïñí»É ³Ûë ·»ÝÇ ¿ùëåñ»ëdzÛÇ ³ÏïÇí³óáõÙÁ Ù»Í ÏÇ볷ݹ»ñÇ Ï»Õ¨áõÙ
ï³ñ³Í³Ï³Ý é»ýÉ»ùëÝ»ñÇ áõëáõóÙ³ÝÁ (Supple W.F., Jr, Leaton R.N., 1990a,b, Martin
20
·áñͳéáõÛÃÝ»ñ, ÇÝãåÇëÇù »Ý. áõÕÕ³ÓÇ· ¹ÇñùÇ å³Ñå³ÝáõÙÁ, ù³ÛÉùÁ, å³éϳÍ
µÝáõÛà ¿ ÏñáõÙ (Bock O., 1998): ì»ñçÇÝë ¿É Ñ³×³Ë å³ï×³é ¿ ¹³éÝáõÙ ß³ñÅáճϳÝ
2001, Hakopian V.P., 2002, Камскова Ю.Г. и др., 2003, Навасардян Г.А. и др., 1998,
2005, Манукян А.А., 2004, Карапетян В.А., 2005, Соцкий О.П. и др., 2005, Ерицян
21
ë³Ï³í³ß³ñÅáõÃÛ³Ý å³ÛÙ³ÝÝ»ñáõÙ ½³ñ·³óáÕ ï»Õ³ß³ñÅ»ñÇ ß³ñùáõÙ Ù»Í
áã ÙdzÛÝ Ï³ñ¨áñ ÃÇñ³Ë ¿ ûùëǹ³ïÇí ëÃñ»ëÇ Ñ³Ù³ñ (MacEvilly C.J., Muller D.P.,
1996), ³ÛÉ Ý³¨ ³ÛÝ, áñ ³Û¹ ûùëǹ³ïÇí íݳëáõÙÁ ëï³ÝáõÙ ¿ Çñ»Ý µÝáñáß
22
âÝ³Û³Í ëÃñ»ëÇ å³ÛÙ³ÝÝ»ñáõÙ (³Û¹ ÃíáõÙ ¨ ë³Ï³í³ß³ñųÛÇÝ) ÎÜÐ
(Shaheen A.A. et al., 1993, Kovacs P. et al., 1996, Liu J., Mori A., 1999, Olivenza
R. et al., 2000, Камскова Ю.Г. и др., 2003): ØÛáõë ÏáÕÙÇó, ѳÛïÝÇ ¿, áñ ûùëǹ³ïÇí
ëÃñ»ëÇ ³é³í»É ½·³ÛáõÝ ÃÇñ³Ë ¿ ·ÉËáõÕ»ÕÁ (MacEvilly C.J., Muller D.P., 1996):
(McIntosh L.J. et al., 1998, Камскова Ю.Г. и др., 2003): ²Ûë µáÉáñÁ Ýå³ëïáõÙ »Ý
23
ì»ñÁ Ýßí³Í ·Éáõï³Ù³ï³ÛÇÝ ¿ùë³ÛïáïáùëÇÏáõÃÛ³Ý Ù»Ë³ÝǽÙÁ ϳñ¨áñ ѳïáõÏ
Ïñ»É, ù³Ý Ù»Í ÏÇ볷ݹ»ñáõÙ, áñÇ ÑÇÙ³Ý íñ³ µ»ñí³Í ³ß˳ï³ÝùáõÙ Ñ»ÕÇݳÏÝ»ñÁ
Çç»óáõÙ, ÇëÏ ³í»ÉÇ áõß Å³ÙÏ»ïÝ»ñáõÙ (15-30 ûñ) µ³ñÓñ³óáõÙ: лï³ùñùÇñ »Ý ݳ¨
24
¶²ÎÂ-Ç ù³Ý³ÏáõÃÛ³Ý ÷á÷áËáõÃÛáõÝÝ»ñÁ ÝáõÛÝ ·Çï³÷áñÓáõÙ: Àëï ³Ûë
¿ ÝϳïíáõÙ, ÇëÏ 30-ñ¹ ûñÝ ³ÛÝ í»ñ³¹³éÝáõÙ ¿ ÑëÏÇã ٳϳñ¹³ÏÇ: àõñÇß Ñ»ÕÇݳÏÝ»ñÇ
ѻﳽáïáõÃÛáõÝ»ñáõÙ (Манукян А.А., 2003, Соцкий и соавт., 2005) óáõÛó ¿ ïñí³Í ¶²ÎÂ-
³éáõÙáí ³ñÛ³Ý Ñ³Ù³Ï³ñ·áõÙ ¹ÇïíáÕ ï»Õ³ß³ñÅ»ñÁ (Акопян В.П. и др., 2001, Манукян
óÇïáùñáÙ b5-Ç) ٳϳñ¹³ÏÝ»ñÇ ³×, ³å³ áõß Å³ÙÏ»ïÝ»ñáõÙ (15 ûñ) ¹ñ³Ýù Ýí³½áõÙ »Ý,
25
ϳñÙÇñ ³ñÛ³Ý Ñ³Ù³Ï³ñ·Ç ûùëǹ³ïÇí íݳëÙ³Ý Ù³ëÇÝ: ´»ñí³Í ³ß˳ï³ÝùÝ»ñáõÙ ëï³óí³Í
ï»Õ³¹ñíáõÙ ¿ åÉ»ùëá·É³ë» Ý»Õ ßßÇ Ù»ç, áõñ ³ÝÑݳñ ¿ ¹³éÝáõ٠ϻݹ³Ýáõ Çñ³ÝÇ
Ùá¹»É:
³½¹»óáõÃÛ³Ùµ. ÙÇ ÏáÕÙÇó ³ÛÝ Ñ³Ý¹»ë ¿ ·³ÉÇë áñå»ë ³½³ï é³¹ÇϳÉ, áñÁ ٳϳÍáõÙ
(å»ñûùëÇÝÇïñÇï, ³å³ ÑǹñûùëÇÉ é³¹ÇϳÉ) (Liu P.K., 2003, Schaller B., 2005):
26
»ñ¨áõÛÃÇÝ, ³½áïÇ ûùëÇ¹Ç ³ñ¹Ûáõݳñ³ñ Ý»ñ·áñÍáõÃÛáõÝÁ ϳËí³Í ¿ Ýñ³ ³ñï³¹ñÙ³Ý
ÑáëùÇ ÑÇÙÝ³Ï³Ý Ï³ñ·³íáñÇãÝ»ñÇó ¿ (Yang G. et al., 1999), ÇÝãå»ë ݳ¨ ³ÛÝ Ï³ñ¨áñ
2000):
íñ³ ÏñÏÝíáÕ ëÃñ»ëǪ ÇÝãå»ë ËóÝáÕ (Luine V. et al., 1996, Ohl F., Fuchs E.,
1999), ³ÛÝå»ë ¿É ÁÝÏ×áÕ (Krugers H.J., et al. 1997) ³½¹»óáõÃÛ³Ý Ù³ëÇÝ: Àëï
Ý»ÛñáÝÝ»ñÇ ³ïñáýdzÛáí:
1993):
Fonseca D.B. et al., 2005): àñáß ¹»åù»ñáõÙ í»ñçÇÝë ûùëǹ³ïÇí ëÃñ»ëÇ ³é³í»É
27
³Ý¹³ñÓ»ÉÇ íݳëÙ³Ý »ÝóñÏí³Í ëåÇï³ÏáõóÝ»ñÇ áõ ÉÇåǹݻñÇ ÏáÙåÉ»ùë, áñÝ Çñ
(Radu R.A. et al., 2003): ´³óÇ ¹ñ³ÝÇó, ÉÇåáýáõëóÇÝÇ Ïáõï³ÏáõÙÁ ³é³í»É ϳñ¨áñ
1996, Brunk U.T., Terman A., 2002): γï³ñí³Í ѻﳽáïáõÃÛáõÝÝ»ñÁ íϳÛáõÙ »Ý, áñ
íݳëáõÙÁ, áñáÝù µ»ñáõÙ »Ý Ïá·ÝÇïÇí ˳óñáõÙÝ»ñÇ (Macey P.M. et al., 2002): ²Û¹
28
ٳϳñ¹³ÏÇ ³×: Àݹ áñáõÙ ûùëǹ³ïÇí ëÃñ»ëÇ å³ïÏ»ñÁ ³í»ÉÇ ³ñï³Ñ³Ûïí³Í ¿ »Õ»É
»Ý:
D.P., 1996): ì»ñçÇÝ »ñÏáõëÁ Ñ»ïÇÝ áõÕ»ÕÇ Ñ»ï Ù»Ïï»Õ ѳٳñíáõÙ »Ý ݳ¨
al., 1996):
29
³ÝÏßé»ÉÇáõÃÛ³Ý Ùá¹»ÉÁ) áõÕ»ÕÇ ¨ áõÕ»ÕÇÏÇ Ï»Õ¨Ç Ý»ÛñáóÇïÝ»ñáõÙ ¹ÇïíáõÙ
30
³ÝáóÛÇÝ ÑÛáõë³ÏÝ»ñÇ ·»ñ³ñÛáõݳɻóٳٵ: лÕÇݳÏÇ ïíÛ³ÉÝ»ñáí ½³ñÏ»ñ³ÏÝ»ñÇ
2002):
ï»Õ³ÛÇÝ ³ñÛ³Ý ÑáëùÇ Ýí³½áõÙ, áñÁ ³Ù»Ý³µ³ñÓñ ¹ñë¨áñáõÙÝ áõÝÇ ¿ 30-ñ¹ ûñÁ,
ï»Õ³ß³ñÅ»ñÇ ¹»ñÇ Ù³ëÇÝ (Акопян В.П., 2003, Мелконян К.В. и соавт., 2003, Melkonyan
31
Éǽáýáëý³ïǹݻñÇ ¨ ³½³ï ׳ñå³ÃÃáõÝ»ñÇ Ó»ñµ³½³ïÙ³ÝÁ: üáëýáÉÇå³½ C-Ç
2004):
·ÉËáõÕ»ÕÁ ϳ٠Ýñ³ ³é³ÝÓÇÝ ³ÛÉ Ï³éáõÛóÝ»ñ: ê³Ï³ÛÝ, ѳßíÇ ³éÝ»Éáí áõÕ»ÕÇÏÇ ÙÇ
32
ëË³É³Ï³Ý ÉÇÝ»É: ØÇ³Å³Ù³Ý³Ï í»ñáÝßÛ³É ·ñ³Ï³Ý ïíÛ³ÉÝ»ñÁ ³å³óáõóáõÙ »Ý áõÕ»ÕÇÏÇ
33
3. кàîàôÂÚ²Ü ÜÚàôºðÜ àõ غÂà¸ÜºðÀ
3.1 λݹ³ÝÇÝ»ñÁ
³éáÕç, 6-9 ³Ùë³Ï³Ý Ù³Ûñ-³éÝ»ïÝ»ñÇ »ñÏñáñ¹ ϳ٠»ññáñ¹ ë»ñݹÇó: 1-1.5 ³Ùë³Ï³Ý
5-6-Ç:
34
ß³ñáõÝ³Ï³Ï³Ý ³Ýß³ñÅáõÃÛ³Ùµ, ³ÛÉ ß³ñÅáÕ³Ï³Ý ³ÏïÇíáõÃÛ³Ý ¨ ë³Ï³í³ß³ñÅáõÃÛ³Ý
Ñݳñ³íáñáõÃÛáõÝ ¿ ï³ÉÇë ï»ëÝ»É ¨ ÇÝã-áñ ã³÷áí ½·³É ÙÇÙÛ³Ýó, ÇÝãÁ áñáß ã³÷áí
íñ³ ³ñí»É »Ý ³Ýóù»ñ, áñáÝó ·áõÙ³ñ³ÛÇÝ Ù³Ï»ñ»ëÁ (21 ëÙ2) ³å³Ñáí»É ¿ Ýñ³
Коваленко Е.А., Гуровский Н.Н., 1980, Жвания М.Г., Костенко Н.А., 1997, Ерицян Н.Б.,
2005):
35
ºÉÝ»Éáí í»ñáÝßÛ³ÉÇó, ë³Ï³í³ß³ñÅáõÃÛ³Ý Ù»ñ û·ï³·áñÍ³Í Ùá¹»ÉáõÙ
ѳٳËï³ÝÇßÇ Ó¨³íáñÙ³Ý í³Õ (45 ûñ) ¨ áõß (70 ûñ) ßñç³ÝÝ»ñáõÙ: ²ÛÝ ¹»åù»ñáõÙ,
ë³Ï³í³ß³ñÅáõÃÛ³Ý ÁÝóóùáõÙ:
í³ñù³ÛÇÝ Ã»ëï»ñÇ Ù³ñïÏáóÇ ÙÇçáóáí (Ýϳñ 3.1), áñÁ Ý»ñ³é»É ¿ §ù³ÛÉù ϳÙñçÇ
ûñ»ñÁ) ßñç³ÝáõÙ ¹ÇïíáÕ í³ñù³ÛÇÝ ï»Õ³ß³ñÅ»ñÁ: ²Û¹ ÇëÏ å³ï׳éáí ³Û¹ ßñç³ÝÁ
36
³é³í»É ѳݷ³Ù³Ý³ÉÇó áõëáõÙݳëÇñí»É ¿ ³é³ÝÓÇÝ (ï»ë 3.4 ß³ñÅáճϳÝ
áõëáõóáõÙÁ):
ݳ˳ӻéÝáõÃÛáõÝ
öáëÇϳíáñ ï³Ëï³Ï 9 41 66
ø³ÛÉù ϳÙñçÇ íñ³ 10,11,12 42,43,44 67,68,69
äïï³ÓáÕ 13 45 70
§ø³ÛÉù ϳÙñçÇ íñ³¦ ûëïáõÙ (Forster M.J. et al., 1996) ³éÝ»ïÁ ï»Õ³¹ñíáõÙ ¿
37
ÁÝï»É³óáõÙÁ ϳÝË»Éáõ ¨ ï»Õ³ß³ñÅÙ³Ý ÙáïÇí³óÇ³Ý ËóݻÉáõ ѳٳñ û·ï³·áñÍí»É ¿
ºä´Ð, 2005), áñáõ٠û÷³Í³ÍÏ Ñ³ï³ÏÇ ÷á˳ñ»Ý û·ï³·áñÍí»É ¿ çñáí Éóí³Í ÉáÕ³í³½³ÝÁ:
í³ÛñÏÛ³Ý), áñÇ ÁÝóóùáõÙ ³éÝ»ïÁ ¹áõñë Ï·³ 10 ëÙ ïñ³Ù³·Íáí ßñç³Ý³ÏÇó (Ýϳñ 3.1. ¶):
ïíÛ³ÉÝ»ñÇ ÙÇçÇÝÁ:
§äïï³ÓáÕǦ ûëïáõÙ (Jeljeli M. et al., 1999, Lalonde R., Strazielle C., 2003)
38
ãÁÝÏÝ»Éáõ ѳٳñ ϻݹ³ÝÇÝ å»ïù ¿ ß³ñÅíÇ ¹»åÇ ³é³ç (Üϳñ 3.1. ´): â³÷í»É ¿ í³Ûñ
Å³Ù³Ý³Ï ³éÝ»ïÁ ³Ùáõñ ϳéã»É ¿ åïï³ÓáÕÇÝ ¨ å³ëÇí åïïí»É Ýñ³ Ñ»ï Ù»Ïï»Õ: ²Ûë
³ñ¹ÛáõÝù:
39
÷áñÓ³ßñç³ÝÇ ÁÝóóùáõ٠ٻͳóí»É ¿ª ³éÝ»ïÝ»ñÇ ³¹³åï³óÇ³Ý ³ñï³óáÉáÕ ß³ñÅáճϳÝ
áõëáõóÙ³Ý ÷á÷áËáõÃÛáõÝÝ»ñÁ ³é³í»É³·áõÛÝë ѳëï³ï³·ñ»Éáõ Ýå³ï³Ïáí:
ê³Ï³í³ß³ñÅáõÃÛ³Ý 1 3 5 7 9 11 13 15
ûñÁ
äïï³ÓáÕÇ åïïÙ³Ý 30 30 30 30 45 50 60 60
³ñ³·áõÃÛáõÝÁ åï/ñ åï/ñ åï/ñ åï/ñ åï/ñ åï/ñ åï/ñ åï/ñ
·ÉËáõÕ»ÕÁ:
40
ëáõå»ñûùëǹ é³¹ÇϳÉÝ»ñáí ÜîÎ-Ç í»ñ³Ï³Ý·ÝÙ³Ý ³ñ¹ÛáõÝùáõÙ ³é³ç³óáÕ ýáñÙ³½³ÝÇ
Ù³ùëÇÙ³É ûåïÇÏ³Ï³Ý ÏɳÝÙ³Ý ËïáõÃÛ³Ý Ýí³½Ù³Ý %-Á (Симонян Г.М. и др., 1999):
àñå»ë êú¸ ³ÏïÇíáõÃÛ³Ý Ùdzíáñ ÁݹáõÝí»É ¿ ýñ³ÏódzÛÇ ³ÛÝ ù³Ý³ÏÁ, áñÝ áõÝ³Ï ¿
áñáßí»É ¿ ûåïÇÏ³Ï³Ý ëå»Ïïñ³É Ù»Ãá¹áí (Владимиров Ю.А., Арчаков А.И., 1972): ظ²-Ç
24 ¨ K-70 (¶»ñÙ³Ýdz):
41
ËÙµÇ ÙÇçÇÝ óáõó³ÝÇßÁ:
42
4. Þ³ñÅáõÙÝ»ñÇ Ñ³Ù³Ï³ñ·Ù³Ý ¨ ß³ñÅáÕ³Ï³Ý áõëáõóÙ³Ý
í³ñùÇ ÷á÷áËáõÃÛáõÝÝ»ñÁ
ø³ ÛÉù Ï ³ ÙñçÇ í ñ³
120,0
110,0 *
100,0
êÞ
90,0
êï áõ· Çã
80,0
70,0
60,0
ܳ Ë ³ ûëï ì³ Õ ØÇçÇÝ àõß
ê³ Ï ³ í ³ ß³ ñÅáõÃÛ³ Ý Å³ ÙÏ »ï
áõÕÕí³ÍáõÃÛáõÝ áõÝ»Ý: ºÃ» ëïáõ·Çã ËÙµÇ ³éÝ»ïÝ»ñÇ Ùáï í³Ûñ¿çùÇ ·³ÕïÝÇ ßñç³ÝÁ
43
107.7±6.4 í.), ³å³ ë³Ï³í³ß³ñųÛÇÝ ËÙµáõÙ í³Õ ßñç³ÝÝ»ñáõÙ ³ÛÝ Ýϳï»ÉÇáñ»Ý
»É³Ï»ï³ÛÇÝÇó ó³Íñ): гñÏ ¿ Ýß»É, áñ ÙÇçÇÝ Å³ÙÏ»ïÝ»ñáõÙ (42ñ¹ - 44ñ¹ ûñ»ñ) ³Ûë
Çñ»Ýó óáõó³ÝÇßÝ»ñÁ:
44
ųÙÏ»ïáõÙ, ÇëÏ ë³Ï³í³ß³ñųÛÇÝ ËÙµÇ Ñ³Ù³ñ ³í»ÉÇ »ñϳñ Å³Ù³Ý³Ï ¿ å³Ñ³ÝçíáõÙ
ä ï ï ³ ÓáÕ
60
*
50
40 êï áõ· Çã
30 êÞ
20
10
0 ê³Ï³í³ß³ñÅáõÃÛ³Ý Å³ÙÏ»ï
ݳ Ë ³ ûëï í³ Õ ÙÇçÇÝ áõß
û·ï³·áñÍ»É ³ÏïÇí í³½ùÁ (ÙÇçÇÝ Ñ³ßíáí 80%): Àݹ áñáõÙ, §í³½ùÇ Å³Ù³Ý³Ï/ÁݹѳÝáõñ
45
Üϳñ 4.3. ê³Ï³í³ß³ñÅáõÃÛ³Ý ï³ñµ»ñ ųÙÏ»ïÝ»ñáõÙ åïï³ÓáÕÇ Ã»ëïáõÙ
³éÝ»ïÝ»ñÇ ³ÏïÇí ѳٳϳñ·í³Í í³½ùÇ Ù³ëݳµ³ÅÇÝÁ (n=12, M±m)
100%
80%
60%
40%
êï áõ· Çã
20% êÞ
0%
0 13 45 70
ê³ Ï ³ í ³ ß³ ñÅáõÃÛ³ Ý Å³ ÙÏ »ï Á (ûñ»ñáí )
46
Üϳñ 4.4. Þ³ñÅÙ³Ý Ý³Ë³Ó»éÝáõÃÛ³Ý Ã»ëïáõÙ ³éÝ»ïÝ»ñÇ ßñç³Ý³ÏÇó ¹áõñë ·³Éáõ
·³ÕïÝÇ ßñç³ÝÇ ÷á÷áËáõÃÛáõÝÝ»ñÁ (í³ÛñÏÛ³ÝÝ»ñáí) ë³Ï³í³ß³ñÅáõÃÛ³Ý ï³ñµ»ñ
ųÙÏ»ïÝ»ñáõÙ (n=12, M±m):
Þ³ ñÅÙ³ Ý Ý³ Ë ³ Ó»éÝáõÃÛáõÝ
60
50
40 êï áõ· Çã
30 êÞ
20
10
0
ݳ Ë ³ ûëï í³Õ ÙÇçÇÝ áõß
˳ݷ³ñáõÙ:
47
Üϳñ 4.5. öáëÇϳíáñ ï³Ëï³ÏÇ Ã»ëïáõÙ ³éÝ»ïÝ»ñÇ ß³ñÅáõÙÝ»ñÇ Ïááñ¹ÇݳódzÝ
³ñï³óáÉáÕ K ·áñͳÏóÇ (ë³ÛóùáõÙÝ»ñÇ Ñ³ñ³µ»ñáõÃÛáõÝÁ ³ÏïÇí ï»Õ³ß³ñÅÙ³Ý
ųٳݳÏÇÝ) ÷á÷áËáõÃÛáõÝÝ»ñÁ ë³Ï³í³ß³ñÅáõÃÛ³Ý ï³ñµ»ñ ųÙÏ»ïÝ»ñáõÙ: (n=12,
M±m, * - p<0.05)
ö áëÇÏ ³ í áñ ï ³ Ë ï ³ Ï
15,00
10,00
* êï áõ· Çã
êÞ
5,00
0,00
ݳ Ë ³ ûëï í³ Õ ÙÇçÇÝ áõß
ê³Ï³í³ß³ñÅáõÃÛ³Ý Å³ÙÏ»ï
˳óñáõÙ:
48
Þ³ñÅáÕ³Ï³Ý áõëáõóÙ³Ý ¹³ë³Ï³Ýª §åïï³ÓáÕǦ ûëïáõÙ, ÝáõÛÝ ³Ûë ųÙÏ»ïáõÙ
49
·áñÍÁÝóóÁ Ù³ëݳÏÇáñ»Ý í»ñ³Ï³Ý·ÝíáõÙ ¿, ë³Ï³ÛÝ, ß³ñÅáÕ³Ï³Ý Ñ³Ù³Ï³ñ·Ù³Ý ÙÛáõëª
al., 2004):
å³ïÏ»ñáõÙ:
ÊáëùÁ ѳïϳå»ë ³ÛÝ ÙϳݳËÙµ»ñÇ Ù³ëÇÝ ¿, áñáÝù ³å³ÑáíáõÙ »Ý ëï³ïÇÏ ¹ÇñùÁ: ²Ûë
ϳéáõÛóÝ»ñÇ íñ³ áõݻݳÉáí áñáß³ÏÇ ïñáýÇÏ ³½¹»óáõÃÛáõÝ (Орбели Л.А., 1940, Canu
ѳÛïÝíáõÙ Ñ»Ýó ³ÛÝ ÙÇÏñá½áݳݻñÁ (Apps R., Garwicz M., 2005), áñáÝù
50
ë³Ï³í³ß³ñÅáõÃÛ³Ý å³ÛÙ³ÝÝ»ñáõ٠ϳï³ñáõÙ ¿ÇÝ ³ÏïÇí ·áñͳéáõÛÃÝ»ñ: ²ÛÝ
51
4.2 15-ûñÛ³ ë³Ï³í³ß³ñÅáõÃÛ³Ý ÁÝóóùáõÙ ³éÝ»ïÝ»ñÇ ß³ñÅáճϳÝ
Ýñ³Ýù ¹»é¨ë ½ÇçáõÙ »Ý ëïáõ·Çã ËÙµÇÝ: 9-ñ¹ ¨ 11-ñ¹ ûñÁ åïï³ÓáÕÇ åïïÙ³Ý
Áëï »ñ¨áõÛÃÇÝ, ÝáõÛÝù³Ý ¹Åí³ñ ËݹÇñ ¿ »Õ»É »ñÏáõ ËÙµ»ñÇ Ñ³Ù³ñ ¿É, áñáÝù
52
Üϳñ 4.6: 15-ûñÛ³ ë³Ï³í³ß³ñÅáõÃÛ³Ý ÁÝóóùáõÙ åïï³ÓáÕÇ Ã»ëïáõÙ
³éÝ»ïÝ»ñÇ í³Ûñ¿çùÇ ·³ÕïÝÇ ßñç³ÝÇ ÷á÷áËáõÃÛáõÝÝ»ñÁ (n=12, M±m, * -
p<0.05)
* * *
120 *
100
í ³ ÛñÏ Û³ ÝÝ»ñ
80 ëï áõ· Çã
60 êÞ
40
20
0
30 30 30 30 45 50 60 60 åïïÙ³Ý ³ñ³·áõÃÛáõÝÁ
(åï/ñáå»)
1 3 5 7 9 11 13 15 êÞ ûñÁ
ËÙµáõÙ åïï³ÓáÕÇ íñ³ ÙݳÉáõ ųٳݳÏǪ É³í³·áõÛÝ ¹»åùáõÙ 80%-Ý ¿ »Õ»É ³ÏïÇí
120.0%
100.0%
* * * *
80.0% ëï áõ· Çã
60.0%
êÞ
40.0%
20.0%
0.0%
å ï ï Ù³ Ý ³ ñ³ · áõÃÛáõÝÁ
30 30 30 30 45 50 60 60 (å ï /ñáå »)
1 3 5 7 9 11 13 15 êÞ ûñÁ
53
²ÛëåÇëáí, ³éÝ»ïÝ»ñÇ 15-ûñÛ³ ë³Ï³í³ß³ñÅáõÃÛ³Ý ¹ÇݳÙÇϳÛÇ
ûñ»ñÇÝ:
54
5. ÈÇåǹ³ÛÇÝ ·»ñûùëǹ³óáõÙÁ ³éÝ»ïÝ»ñÇ áõÕ»ÕÇÏáõÙ
²ÕÛáõë³Ï 5.1
àõÕ»ÕÇÏÇ ÑÛáõëí³ÍùáõÙ Ù»ï³Õ³åñáï»ÇÝÝ»ñÇ ¨ Ù³ÉáݳÛÇÝ ¹Ç³É¹»ÑÇ¹Ç Ù³Ï³ñ¹³Ï-
Ý»ñÇ µ³ó³ñÓ³Ï ¨ ѳñ³µ»ñ³Ï³Ý ÷á÷áËáõÃÛáõÝÝ»ñÁ (%) ëïáõ·Çã ËÙµÇ óáõó³ÝÇß-
Ý»ñÇ (100%) ѳٻٳïáõÃÛ³Ùµ (M±m, n=14)
ê³Ï³í³ß³ñÅáõÃÛ³Ý Å³ÙÏ»ïÁ
êïáõ·Çã
òáõó³ÝÇß Ødzí. 2 ûñ 7 ûñ 15 ûñ 45 ûñ 70 ûñ
ËáõÙµ
²Ø/ · 130.0±5.8 96,07±4.03 ** 119.86±0.39 * 132±1.43 † 119.47±1.43* 231.4±7.8 **
êú¸ §
% -26.1±3.1 -7.8±0.3 +2.1±1.1 -8.1±1.1 +78.0±6.0
²Ø/ · 90.2±3.8 98.50±0.90 * 84.70±0.72 * 58.4±3.97 ** 93.36±0.36 † 104.9±1.98 *
ϳï³É³½
% +9.2±1.0 -6.1±0.8 -35.3±4.4 +3.5±0.4 +16.4±2.2
äØ 0.060±0.0 0.037±0.002* 0.029±0.001* 0.027±0.003* 0.065±0.001* 0.061±0.001 †
óÇïáùñáÙ c 1 * *
%
-38.1±3.3 -51.3±2.2 -55.8±4.3 +8.0±1.6 +1.4±1.6
ÙÏÙáÉ/ 264.0±12. 297.0±3.43 ** 289.6±3.7* 170.8±6.34** 269.5±6.07 † 260.8±3.70 †
ظ² · 0
% +12.5±1.3 +9.7±1.4 -35.3±2.4 +2.1±2.3 -1.2 ±1.4
̳ÝáÃáõÃÛáõÝ.
²Ø – ³ÏïÇíáõÃÛ³Ý Ùdzíáñ, äØ – å³ÛÙ³Ý³Ï³Ý Ùdzíáñ, * p<0.05; ** p<0.01; † p>0.05; § –
CuZn- ¨ Mn2+-êú¸ ·áõÙ³ñ³ÛÇÝ ³ÏïÇíáõÃÛáõÝ
55
ÆÝãå»ë »ñ¨áõÙ ¿ Ý»ñϳ۳óí³Í ïíÛ³ÉÝ»ñÇó, ѳϳûùëǹ³Ýï³ÛÇÝ ý»ñÙ»ÝïÝ»ñÇ ¨
ѻﳷ³ÛáõÙ Ýñ³ ù³Ý³ÏÁ ëÏëáõÙ ¿ ٻͳݳÉ, 15ñ¹ ûñÁ ѳëÝ»Éáí ëïáõ·Çã ËÙµÇ
óáõó³ÝÇßÝ»ñÇÝ: 45ñ¹ ûñÁ Ýñ³ ù³Ý³ÏÁ ÙÝáõÙ ¿ ëïáõ·Çã ËÙµÇó ó³Íñ, µ³Ûó 75ñ¹
áõÕÕí³Í ï»Õ³ß³ñÅ»ñÁ ë³Ï³í³ß³ñÅáõÃÛ³Ý 7ñ¹ ûñÁ (Çç»óáõÙ), ¨ áõß (70ñ¹ ûñ) ųÙ-
ñÇÝ, ½·³ÉÇ Çç»óáõÙ 15ñ¹ ûñÁ, ³å³ í»ñ³¹³ñÓ ëïáõ·Çã ٳϳñ¹³ÏǪ áõß Å³ÙÏ»ïÝ»ñáõÙ
(45ñ¹ ¨ 70ñ¹ ûñ»ñ): òÇïáùñáÙ c-Ç Ù³Ï³ñ¹³ÏÁ í³Õ ųÙÏ»ïÝ»ñáõ٠ϳÛáõÝ Ýí³½áõÙ ¿
(2ñ¹, 7ñ¹, 15ñ¹ ûñ»ñ), ÇëÏ áõß Å³ÙÏ»ïÝ»ñáõÙ í»ñ³¹³éÝáõÙ ¿ ëïáõ·Çã óáõó³ÝÇßÝ»ñÇÝ:
5.2):
²ÕÛáõë³Ï 5.2
²ÙµáÕç³Ï³Ý ·ÉËáõÕ»Õ³ÛÇÝ ÑÛáõëí³ÍùáõÙ (³é³Ýó áõÕ»ÕÇÏÇ) Ù»ï³Õ³åñáï»ÇÝÝ»ñÇ
¨ Ù³ÉáݳÛÇÝ ¹Ç³É¹»ÑÇ¹Ç Ù³Ï³ñ¹³ÏÝ»ñÇ µ³ó³ñÓ³Ï ¨ ѳñ³µ»ñ³Ï³Ý ÷á÷áËáõ-
ÃÛáõÝÝ»ñÁ (%) ëïáõ·Çã ËÙµÇ óáõó³ÝÇßÝ»ñÇ (100%) ѳٻٳïáõÃÛ³Ùµ (M±m,
n=14)
ê³Ï³í³ß³ñÅáõÃÛ³Ý Å³ÙÏ»ïÁ
êïáõ·Çã
òáõó³ÝÇß Ødzí. 2 ûñ 7 ûñ 15 ûñ 45 ûñ 70 ûñ
ËáõÙµ
²Ø/ · 174.3±14.1 232.0±4.8 ** 200.1±4.0 * 214.0±2.1**
218.0±3.1** 296.5±8.9 **
êú¸ §
% +33.1±2.8 +14.8±2.3 +23.2±1.2 +25.1±1.8 +70.1±5.1
²Ø/ · 127.5±6.3 155.7±2.2 * 142.8±2.4 * 139.1±1.4 * 146.7±1.8 * 154.8±1.4 *
ϳï³É³½
% +22.1±1.7 +12.0±1.9 +9.1±1.1 +15.1±1.4 +21.4±1.1
äØ 0.048±0.003 0.073±0.002 ** 0.065±0.002 ** 0.059±0.001 * 0.0481±0.001 † 0.055±0.001 *
óÇïáùñáÙ c
% +51.3±4.1 +35.2±3.2 +22.1±1.9 +0.6±0.9 +15.0±1.1
ÙÏÙáÉ/ 580.0±26.5 596.8±1.2 † 392.7±8.7 ** 404.3±12.2 ** 609.0±2.32 † 284.2±19.72 **
ظ² ·
% +2.9±0.2 -32.3±1.5 -30.3±2.1 +5.0±0.4 -51.0±3.4
56
سëݳíáñ³å»ë, ѳñÏ ¿ Ýß»É, áñ û’ êú¸-Ç, ¨ û’ ϳï³É³½Ç ٳϳñ¹³ÏÝ»ñÁ
гٳå³ï³ë˳ݳµ³ñ, ظ² ù³Ý³ÏÁ µ³í³Ï³Ý ó³Íñ ¿ ÙÝáõÙ (µ³ó³éáõÃÛ³Ùµ 45ñ¹ ûñí³, »ñµ
1980, Навасардян Г.А., Ерицян Н.Б., 2003, Григорян А.С. и др., 2006), áõÕ»ÕÇÏáõÙ
ٻ˳ÝǽÙÝ»ñÇó Ù»ÏÁ:
57
ѳݹ»ë ¿ ·³ÉÇë »ñÏñáñ¹³ÛÇÝ Ù»ë»Ýç»ñÇ Ï³ñ·³íÇ׳ÏáõÙ` ³ÏïÇí³óÝ»Éáí áõÝÇí»ñë³É
ýáëý³ï³½Ý»ñÁ (Finkel T., 1998, Rhee S.G., 1999, Dalton T.P. et al., 1999,
Thannickal V.J., 2000, Surh Y.J. et al., 2004): Ø»ñ ϳñÍÇùáí, ³Ëï³ýǽÇáÉá·Ç³Ï³Ý
áõÕ»ÕÇÏÁ ·ÉËáõÕ»ÕÇ Ï»Õ¨Ç ¨ ½áɳíáñ Ù³ñÙÝÇ Ñ»ï Ù»Ïï»Õ ѳݹ»ë ¿ ·³ÉÇë áñå»ë
ûùëǹ³ïÇí ëÃñ»ëÇ ³é³í»É ½·³ÛáõÝ ÃÇñ³Ë (MacEvilly C.J., 1996): ê³Ï³ÛÝ, in vivo
1. êÃñ»ë 黳Ïódz:
·áñÍÁÝóóÝ»ñÇ íñ³ (Kovacs P. et al.,1996, Madrigal J.L. et al., 2001, Gluck M.R. et
al., 2001, Sahin E., Gumuslu S., 2004): Àݹ áñáõ٠ϳñ¨áñ ¹»ñ »Ý Ó»éù µ»ñáõÙ
58
óÇïáåɳ½Ù³ÛÇÝ Cu-Zn êú¸-Ç ¨ ·Éáõï³ÃÇáÝ é»¹áõÏï³½Ç ³ÏïÇíáõÃÛáõÝÝ»ñÁ,
ٻͳóáõÙÁ (Камскова Ю.Г. и др., 2003): ì»ñçÇÝë ÑÇÙù ¿ ï³ÉÇë »Ýó¹ñ»Éáõ, áñ Ù»ñ
ÙïóÝáõÙ ÁÝóóùÇ Ù»ç ·ïíáÕ ß³ñÅÙ³Ý Íñ³·ñáõÙ, ³ÛÉ Ý³¨ áñáß³ÏÇ Ý߳ݳÏáõÃÛáõÝ
59
ºÄÀ ٻ˳ÝǽÙáí ë³Ñٳݳ÷³Ï»Éáí Ca2+ ÙáõïùÁ µçÇç, ³Ûë ٻ˳ÝǽÙÁ ϳñáÕ ¿
å³Ñå³ÝÙ³Ý Ñ³Ù³ñ:
Ýí³½áõÙÁ (Акопян В.П., 1999, Акопян В.П., Кочарян А.Г., 1999, Hakopian V.P.,
Fonnum F., Lock E.A., 2000, Welsh J.P. et al., 2002): ²Ûë ٻ˳ÝǽÙÇ ·áñͳñÏÙ³Ý
÷á÷áËáõÃÛáõÝÝ»ñÁ:
60
·áñÍÁÝóóÇ áñáß³ÏÇ ³ëïÇ׳ÝÇ Ñ³ëÝ»Éáõ ¹»åùáõÙ ³ÛÝ ³Ýç³ïíáõÙ ¿
ѳϳÑÇåûùë³Ýï:
³½³ï é³¹ÇϳÉÝ»ñÇ ³í»ÉÇ Ù»Í ù³Ý³Ïáí ·áÛ³óÙ³Ý, áñÝ ¿É ѳݷ»óÝáõÙ ¿ ظ²-Ç Ù³Ï³ñ¹³ÏÇ
(Skulachev V.P., 1999, Chernyak B.V. et al., 2004), áñÁ ¹Çï³ñÏíáõÙ ¿ áñå»ë
61
³¹³åïÇí ï»Õ³ß³ñÅ` áõÕÕí³Í ³½³ï é³¹ÇϳÉÝ»ñ ·áÛ³óÝáÕ íݳëí³Í ÙÇïáùáݹñÇáõÙÝ»ñÇ
å³Ï³ë ϳÛáõÝ ¿ ¨ Ëáó»ÉÇ, ѳïϳå»ë íݳëáÕ Ù»Ï ³ÛÉ ·áñÍáÝÇ Ñ³í»ÉÙ³Ý ¹»åùáõÙ:
áõÕ»ÕÇÏáõ٠ظ²-Ç, êú¸-Ç, ϳï³É³½Ç, óÇïáùñáÙ c-Ç Ñ»ï ³éÝãíáÕ Ù»ñ ëï³ó³Í
³é³í»É ɳÛÝ ÇÙ³ëïáí, ³ÛÝ ¿. µççáõÙ ÃÃí³ÍÝÇ ³ÏïÇí Ó¨»ñÇ ù³Ý³ÏÇ Ù»Í³óÙ³Ý Ñ»ï¨³Ýùáí
62
ѳݷ³Ù³ÝùÁ, áñ ³½³ï é³¹ÇϳÉÝ»ñÁ (ÃÃí³ÍÝÇ ¨ ³½áïÇ ³ÏïÇí Ó¨»ñÁ)ª ù³Ý³ÏáõÃÛáõÝÇó,
ë³Ï³í³ß³ñÅáõÃÛ³Ý Å³ÙÏ»ïÝ»ñÇó:
63
6. ²éÝ»ïÝ»ñÇ áõÕ»ÕÇÏáõÙ µçç³-³ÝáóÛÇÝ ï»Õ³ß³ñÅ»ñÁ
ï»Õ³ß³ñÅ»ñÁ:
áñáÝù ·ïÝíáõÙ »Ý Ñ»Ýó ³ÛÝ Ñ³ïí³ÍáõÙ, áñï»Õ å»ïù ¿ ÉÇÝ»ñ ÝáñÙ³É äáõñÏÇÝÛ»Ç
64
Üϳñ 6.1
ÐëÏÇã ËáõÙµ,
(ѻٳïáùëÇÉÇÝ-
¿á½ÇÝ, Ëáßáñ³-
óáõÙ x 90)
²ÝáÃÝ»ñÁ ¨
äáõñÏÇÝÛ»Ç
µçÇçÝ»ñÁ ÇÝï³Ïï
»Ý:
Üϳñ 6.2
7 ûñÛ³ ë³Ï³í³-
ß³ñÅáõÃÛ³Ý
ËáõÙµ, (ѻٳ-
ïáùëÇÉÇÝ-¿á½ÇÝ,
Ëáßáñ³óáõÙ x90)
ºñ¨áõÙ ¿
¿Ý¹áûÉÇá-
óÇïÝ»ñÇ ³Ûïáõó-
í³ÍáõÃÛáõÝ, ³ñÛ³Ý
ÑáëùÇ ¹³Ý¹³ÕáõÙ:
²ÝáÃÇÝ Ñ³ñáÕ
ѳïí³ÍáõÙ
äáõñÏÇÝÛ»Ç µçÇç-
Ý»ñÇ íݳëáõÙ` Ýñ³
ï»ÕáõÙ »ñ¨áõÙ ¿
³åáåïáïÇÏ Ù³ñÙÝÇÏ
ÑÇß»óÝáÕ ·áïÇ
(óáõÛó ¿ ïñí³Í
ëɳùáí):
65
Üϳñ 6.3
7 ûñÛ³ ë³Ï³í³-
ß³ñÅáõÃÛ³Ý
ËáõÙµ, Ý»ñÏáõÙ
(Ý»ñÏáõÙ Áëï
ÜÇëÉÇ Ëáßáñ³-
óáõÙ x 64)
êɳùáí óáõÛó ¿
ïñí³Í óÇïá-
åɳ½Ù³ÛÇ
Ù·³óáõÙáí
äáõñÏÇÝÛ»Ç
µçÇç
(óÇïáåÇÏÝá½`
¹»Ý¹ñáé»ùëÇëáí)
ýǵñÇݳÛÇÝ ÃñáÙµ»ñáí,
¿Ý¹áûÉÇáóÇïÝ»ñÇ §µ³ñÓñ³óáõÙ¦,
·) áñáß Ù³½³ÝáÃÝ»ñ ËÇëï ɳÛݳó³Í »Ý, µ³í³Ï³Ý Ñ³×³Ë Ñ³Ý¹ÇåáõÙ »Ý ëï³½Ç ¨ ëɳçÇ
ï»Õ³Ù³ë»ñ,
áñï»Õ 1 ÙÏÙ-Ç íñ³ 5-6 µçÇç ¿ »ñ¨áõÙ, ë³Ï³í³ß³ñųÛÇÝ ËÙµáõ٠ѳßííáõÙ »Ý 2-3Á
(ÝÏ. 6.5): ê³Ï³ÛÝ å»ïù ¿ Ýß»É, áñ ù³Ý³Ï³Ï³Ý ³Ûë ï»Õ³ß³ñÅÁ áã µáÉáñ ¹³ßï»ñáõÙ ¿ñ
66
¹ÇïíáõÙ: ä³Ñå³Ýí³Í äáõñÏÇÝÛ»Ç µçÇçÝ»ñÇ ÙÇ Ù³ëáõÙ ¹ÇïíáõÙ »Ý Ïáñǽ³ÝÛáõÃÇ
67
ØáñýáÉá·Ç³Ï³Ý ï»Õ³ß³ñÅ»ñÁ áõÕ»ÕÇÏáõÙ 70 ûñÛ³ ë³Ï³í³ß³ñÅáõÃÛ³Ý
å³ÛÙ³ÝÝ»ñáõÙ
ѳñÏ ¿ ¨ë Ù»Ï ³Ý·³Ù Áݹ·Í»É, áñ ë³Ï³í³ß³ñÅáõÃÛ³Ý Ã»° í³Õ, ¨ û° áõß Å³ÙÏ»ïÝ»ñáõÙ
68
ï³ñµ»ñáõÃÛáõÝÝ»ñÁ (Hawkes, R., Herrup, K., 1995, Hawkes, R., Eisenman, L.M.,
1997, Welsh J.P. et al., 2002) ѳݷ»óÝáõÙ »Ý íݳëáÕ ·áñÍáÝÇ Ñ³Ý¹»å áõñáõÛÝ
et al., 2002): ²Ûëå»ë, 10 ñáå» ï¨áÕáõÃÛ³Ùµ ëñïÇ Ï³Ý·Çó 1 ß³µ³Ã ³Ýó ¹ÇïíáõÙ ¿
·»ñ³ÏßéÙ³Ý Ù³ëÇÝ:
69
ëɳùáí): ²Ûë »ñ¨áõÛÃÇ ¿áõÃÛáõÝÁ ѳëϳݳÉáõ ѳٳñ ѳñÏ ¿ ÑÇß»É, áñ
¿ùë³ÛïáïáùëÇÏáõÃÛáõÝÁ (Olney, J.W., Ishimaru, M.J., 1999, Dallas M. et al., 2007, Hertz L.,
³Ûïáõó³ÛÇÝ Ý»Ïñá½:
70
Àݹ áñáõÙ, ³é³çÇÝ áõÕÇÝ ÑÇÙݳϳÝáõÙ ÙÇçÝáñ¹íáõÙ ¿ NMDA, ÇëÏ
AMPA ÁÝϳÉÇãÝ»ñáí (Farrant M. and Cull-Candy G. 1991, Slemmer J.E. et al., 2005,
Launey T., 2007, Kovacs A.D., Pearce D.A., 2008): ¸ñ³Ý ѳٳå³ï³ëË³Ý ³Ûë
·áñÍáÝÝ»ñáí:
1. üáõÝÏóÇáÝ³É ³ÏïÇíáõÃÛáõÝ:
³å³ÑáíÙ³Ý ÙÇç¨ ³éϳ ϳåÁ ѳïϳå»ë ë»ñï ¿ äáõñÏÇÝÛ»Ç µçÇçÝ»ñÇ ¹»åùáõÙ (Cesa
71
R, Strata P., 2007): ê³Ï³ÛÝ ³Ûë ¿ÙåÇñÇÏ Ùáï»óáõÙÁ å³Ñ³ÝçáõÙ ¿ áñáß³ÏÇ
al., 1998, Manda K. et al., 2007): ÆѳñÏ», ºÄÀ Ý³Ë ¨ ³é³ç áõÕ»ÕÇÏÇÝ µÝáñáß
2007, Jin Y. et al, 2007, Ogasawara H. et al., 2008), ë³Ï³ÛÝ ³Û¹ ٻ˳ÝǽÙÇ
2. ¶ÉÇ³É ·áñÍáÝ:
û¨ ûñ·³ÝǽÙÇ Ñ³Ù³ñ ¿ë»ÝóÇ³É ã»Ý, ë³Ï³ÛÝ Ýñ³Ýó ëÇÝû½Ç ѳٳñ ³ÝÑñ³Å»ßï 3-
72
Ýß»É Ý³¨, áñ áõÕ»ÕÇÏÇ Ï»Õ¨áõÙ ³ëïñá·ÉdzÛÇ µçÇçÝ»ñÁ ¨ë ³Ýѳí³ë³ñ³ã³÷ »Ý
µ³ßËí³Í:
(Welsh J.P. et al., 2002): ì»ñçÇÝÝ»ñë ϳñ¨áñ ¹»ñ áõÝ»Ý ÑÇåûùëÇÏ å³ÛÙ³ÝÝ»ñáõÙ
Н.Б., 2005):
ï»Õ³ß³ñÅ»ñÁ ³ÝÝß³Ý »Ý: ê³Ï³ÛÝ ³Ûë µçÇçÝ»ñÇ ¨ äáõñÏÇÝÛ»Ç µçÇçÝ»ñÇ ÙÇç¨ ³éϳ
¿ ë»ñï ÙáñýáýáõÝÏóÇáÝ³É Ï³å: ²Û¹ ÇëÏ å³ï׳éáí Ýßí³Í ß»ñïáõÙ ï»Õ ·ï³Í ï»Õ³ß³ñÅ»ñÇ
·Ý³Ñ³ïÙ³Ý Ñ³Ù³ñ ã³÷í»É ¿ ß»ñïÇ Ñ³ëïáõÃÛáõÝÁ (Ýϳñ 6.8): êï³óí³Í ïíÛ³ÉÝ»ñÁ óáõÛó
73
Üϳñ 6.8 àõÕ»ÕÇÏÇ Ï»Õ¨Ç ·ñ³ÝáõÉÛ³ñ ß»ñïÇ É³ÛÝáõÃÛáõÝÁ ë³Ï³í³ß³ñÅáõÃÛ³Ý
å³ÛÙ³ÝÝ»ñáõÙ (n=40, M±m):
áõÝÁ, ÙÏ Ù
25.0
³ ñ ß»ñï Ç Ñ³ ëï áõÃÛ
20.0
15.0
10.0
¶ñ³ ÝáõÉÛ
5.0
0.0
7 ûñ 70 ûñ ÐëÏ Çã
ê³ Ï ³ í ³ ß³ ñÅáõÃÛ
³ Ý Å³ ÙÏ »ï Á
àõß³·ñ³í ¿ ݳ¨ ³ÛÝ Ñ³Ý·³Ù³ÝùÁ, áñ ÇÝï³Ïï ³éÝ»ïÝ»ñÇ Ùáï ³Ûë ß»ñïÇ É³ÛÝáõÃÛáõÝÁ
14
áõÝ
1.0
г ׳ Ë ³ Ï ³ ÝáõÃÛ
12
0.75 10
8
0.5
6
4
0.25
2
0.0 0
7 ûñ 70 ûñ ÐëÏ Çã 18.0 20.0 22.0 24.0 26.0 28.0 18.0 20.0 22.0 24.0 26.0 28.0 18.0 20.0 22.0 24.0 26.0 28.0
74
ÙÇç¨: ê³ Ñ³ïϳå»ë Ýϳï»ÉÇ ¿ ¹³éÝáõÙ áõß Å³ÙÏ»ïáõÙ: Üßí³Í ï»Õ³ß³ñÅÁ Ù»Ïݳµ³Ý»Éáõ
ëïáõ·³µ³ÝáõÙÁ:
75
ÏáÕÙÇó, í»ñçÇÝ ïíÛ³ÉÝ»ñÇ Ñ³Ù³Ó³ÛÝ, ÑÇëï³ÙÇÝÁ áõÕ»ÕÇÏáõ٠ϳï³ñáõÙ ¿ ݳ¨
áñáÝó Ù»Í Ù³ëÁ ¹³ñÓ»ÉÇ µÝáõÛà ¿ ÏñáõÙ: ê³Ï³í³ß³ñÅáõÃÛ³Ý áõß Å³ÙÏ»ïáõÙ ³ñÛ³Ý
³Ýѳٳã³÷áõÃÛáõÝ:
76
7. ²éÝ»ïÝ»ñÇ ½³Ý·í³ÍÇ, áõÕ»ÕÇÏÇ ¨ ³½¹ñÇ ù³é³·ÉáõË ÙϳÝÇ
Ýí³½áõÙ (Коваленко Е.А., Гуровский Н.Н., 1980, Оганов В.С., 1998, Ерицян Н.Б.,
2005):
§û·ï³·áñÍíáõÙ¦:
77
Üϳñ 7.1
150
100
50
0
ÐëÏ Çã 7 ûñ 15 ûñ 45 ûñ 70 ûñ
ê³ Ï ³ í ³ ß³ ñÅáõÃÛ³ Ý ï ¨ áÕáõÃÛáõÝÁ
__
Üϳñ 7.2
1500
1000
500
0
ÐëÏ Çã 7 ûñ 15 ûñ 45 ûñ 70 ûñ
ê³ Ï ³ í ³ ß³ ñÅáõÃÛ³ Ý ï ¨ áÕáõÃÛáõÝÁ
__
78
Üϳñ 7.3
250
200
150
100
50
0
ÐëÏ Çã 7 ûñ 15 ûñ 45 ûñ 70 ûñ
ê³ Ï ³ í ³ ß³ ñÅáõÃÛ³ Ý ï ¨ áÕáõÃÛáõÝÁ
Üϳñ 7.4
0.30
0.20
0.10
0.00
ÐëÏ Çã 7 ûñ 15 ûñ 45 ûñ 70 ûñ
ê³ Ï ³ í ³ ß³ ñÅáõÃÛ³ Ý ï ¨ áÕáõÃÛáõÝÁ
79
ÆÝãå»ë óáõÛó ¿ ïñí»É ë³Ï³í³ß³ñÅáõÃÛ³Ý ³ÛÉ Ùá¹»ÉÝ»ñáõÙ, Ù»ñ ·Çï³÷áñÓÇ
ù³ßÁ Ýí³½áõÙ ¿ Ùáï 16%, ÇëÏ ù³é³·ÉáõË ÙϳÝÇ ù³ßÁ Ýí³½áõÙ ¿ Ùáï 58%: Üßí³Í
p<0.0001):
ٻͳóÙ³ÝÁ ½áõ·ÁÝóó áõÕ»ÕÇÏÇ ù³ßÇ Ýí³½Ù³Ý ÙÇïáõÙ (ÝÏ. 7.3): êï³óí³Í ïíÛ³ÉÝ»ñÇ
Üϳñ 7.5
²éÝ»ïÝ»ñÇ áõÕ»ÕÇÏÇ ¨
³½¹ñÇ ù³é³·ÉáõË ÙϳÝÇ
½³Ý·í³ÍÝ»ñÇ Ñ³ñ³µ»-
ñ³ÏóáõÃÛ³Ý ëϳï»ñ³ÛÇÝ
å³ïÏ»ñ:
80
ï¨áÕáõÃÛ³Ý ³í»É³óÙ³ÝÁ ½áõ·ÁÝóó ÝßÛ³É Ñ³ñ³µ»ñ³ÏóáõÃÛáõÝÁ ûñÇݳã³÷áñ»Ý
81
áëÏñ³ÛÇÝ ÑÛáõëí³ÍùáõÙ ¨ë ·»ñÇßËáõÙ »Ý ϳﳵáÉÇÏ åñáó»ëÝ»ñ (Коваленко Е.А.,
Гуровский Н.Н., 1980, Doty, S. B. et al., 1990, Оганов В.С. 1998): ØdzųٳݳÏ,
ïíÛ³ÉÝ»ñÁ (Krebs J.M., et al., 1990, Acheson K.J. et al. 1995, Gretebeck R.J., et
å³Ï³ë ß»ßï³ÏÇ ¿ ï»ÕÇ áõÝ»ÝáõÙ: ê³Ï³ÛÝ, Ù³ñÙÝÇ ¨ ³½¹ñÇ ù³é³·ÉáõË ÙϳÝÇ ù³ß»ñÇ
÷áËϳå³Ïóí³ÍáõÃÛ³Ý Ù³ëÇÝ:
82
ÆÝãå»ë Ýßí»ó, áõÕ»ÕÇÏÇ ù³ßÁ ë³Ï³í³ß³ñÅáõÃÛ³Ý ÁÝóóùáõÙ ¨ë Ýí³½Ù³Ý
p=0.145) óáõÛó ïí»ó, áñ ³Û¹ óáõó³ÝÇßÝ»ñÇ ÙÇç¨ ³éϳ ÃáõÛÉ Ïáé»ÉÛ³óÇáÝ Ï³åÁ
³ÛÝ Ñ³Ý·³Ù³ÝùÁ, áñ ³Ûë óáõó³ÝÇßÁ ß³ï ùÇã ¿ ï³ï³Ýí»É ÑëÏÇã ËÙµÇ Ï»Ý¹³ÝÇÝ»ñÇ
Ùáï (ÝÏ. 7.4), ϳñ»ÉÇ ¿ »Ýó¹ñ»É, áñ ûñ·³ÝǽÙÇ ÏáÕÙÇó àõøÐ Ýßí³Í §í»ñ³Ý³ÛáõÙÁ¦
ï»Õ³ß³ñÅ»ñÇ Ñ³Ù³ËÙµáõÙ:
83
8. øÝݳñÏáõÙ ¨ ³Ù÷á÷áõÙ
ųÙÏ»ï-ϳËÛ³É ³ëå»Ïïáí:
Е.А., Гуровский Н.Н., 1980, Федоров И.В., 1982, Navasardyan G.A. et al., 2004, Ерицян Н.А.,
2004): Ø»ñ ÏáÕÙÇó ëï³óí³Í ïíÛ³ÉÝ»ñÁ ÃáõÛÉ »Ý ï³ÉÇë §ëáõñ ëÃñ»ë³ÛÇݦ ѳٳñ»É
84
²ñÛ³Ý ßñç³Ý³éáõÃÛ³Ý Ýßí³Í ÷á÷áËáõÃÛáõÝÝ»ñÁ ѳݷ»óÝáõÙ »Ý ݳ¨
·áÛ³óáõÃÛáõÝÝ»ñ:
85
ØáÉ»ÏáõÉÛ³ñ ٳϳñ¹³Ïáí ÁÝóóáÕ ³Ûë ï»Õ³ß³ñÅ»ñÝ áõÝ»ó»É »Ý ѳٳå³ï³ë˳Ý
¹»ýÇóÇïÁ:
´áÉáñáíÇÝ í»ñç»ñë óáõÛó ¿ ïñí»É, áñ ºÄÀ ³é³ç³óÙ³Ý Ñ³Ù³ñ ³ÝÑñ³Å»ßï ¿ Ù»Ï ³ÛÉ
ȶú, ³½³ï 鳹dzϳɳÛÇÝ ·áñͳéáõÛÃÝ»ñÇ ¨ áõëáõóÙ³Ý ÙÇç¨ ³éϳ ¿ ë»ñï ϳå, áñÁ,
86
å³Ñ³ÝçíáõÙ: ²Ûë ï»Õ³ß³ñÅ»ñÁ ѳïϳå»ë ³ñï³Ñ³Ûïí³Í »Ý ë³Ï³í³ß³ñÅáõÃÛ³Ý 3-Çó 9-ñ¹
ûñ»ñÇÝ:
(Finkel T., 1998, Rhee S.G., 1999, Dalton T.P. et al., 1999, Thannickal V.J., 2000,
Surh Y.J. et al., 2004): Ø»ñ ϳñÍÇùáí, Ñ»Ýó ³Ûë ٻ˳ÝǽÙÝ ¿ ÑÇÙù»ñ ëï»ÕÍáõÙ
³ÏïÇí³óÙ³Ý å³ïÏ»ñÇÝ:
87
§åïï³ÓáÕÇ Ã»ëï»ñáõÙ¦ ³Ûë ųÙÏ»ïÝ»ñáõÙ ë³Ï³í³ß³ñÅáõÃÛ³Ý »ÝóñÏí³Í ϻݹ³ÝÇÝ»ñÇ
ûùëǹ³ïÇí íݳëÙ³Ý Ñ³Ý¹»å ³é³í»É ½·³ÛáõÝ µçÇçÝ»ñÝ ³ñ¹»Ý ٳѳó»É »Ý, ÇëÏ
Àëï »ñ¨áõÛÃÇÝ, ¹ñ³ Ñ»ï ¿ ϳåí³Í ݳ¨ ѳïÇϳíáñ ß»ñïÇ Ñ³ëïáõÃÛ³Ý ßáõñç 17%
88
í»ñ³Ï³Ý·ÝíáõÙ »Ý: ê³Ï³ÛÝ, ß³ñÅáÕáõÃÛ³Ý Ñ³Ù³Ï³ñ·Ù³Ý ³¹³åïÇí
å³ïÏ»ñáõÙ:
ÙÇïáõÙÁ:
˳Ûï³µÕ»ï å³ïÏ»ñÁ:
89
̳ÝáÃáõÃÛáõÝ`
´»ñí³Í äáõñÏÇÝÛ»Ç µççÇ ÝϳñÁ í»ñóí³Í ¿ http://www.brains-minds-media.org/archive/228
¿çÇó
1. ÐÇåá/¹»³ý»ñ»Ýï³ódz
90
ÆÝãå»ë Ýßí»É ¿, ë³Ï³í³ß³ñÅáõÃÛ³Ý å³ÛÙ³ÝÝ»ñáõÙ ù³Ý³Ï³å»ë Ýí³½áõÙ ¨
2. úùëǹ³ïÇí íݳëáõÙ
³å³ÑáíÙ³Ý íñ³:
3. Æß»Ùdz/ÑÇåûùëdz
Ýϳïٳٵ, ù³ÝÇ áñ ûÅïí³Í »Ý Ñëϳ ¹»Ý¹ñÇïÇÏ Í³éáí, áñÝ ³é³í»É ѳí³Ý³Ï³Ý ¿ ¹³ñÓÝáõÙ
91
ѻ勉Ýùáí: ê³Ï³ÛÝ áõÕ»óáõÛó áõݻݳÉáí ³Ëï³ýǽÇáÉá·Ç³Ï³Ý
Ý߳ݳÏáõÃÛáõÝÁ áñáß Ñ»ÕÇݳÏÝ»ñ (De Schutter E., 1995, Llinas R. et al., 1997,
4. ¶ÉÇ³É ·áñÍáÝ:
ѳٳñ:
¹»ñ ϳñáÕ ¿ Ë³Õ³É Ý³¨ ³Ûëå»ë Ïáãí³Í Ï»ÝïñáÝ³Ï³Ý Ý»Ûñá·»Ý Ý»Ûñáåñáï»ÏóÇ³Ý (Reis
92
ųÙÏ»ïáõÙ Ýßí³Í óáõó³ÝÇßÝ»ñÇ ÙÇç¨ ³Û¹ ½áõ·³Ñ»é³Ï³ÝáõÃÛáõÝÁ ãÇ ÝϳïíáõÙ:
³×ÇÝ, áõß Å³ÙÏ»ïáõÙ (45ñ¹ ¨ 70ñ¹ ûñ»ñ) ³ÛÝ Ýí³½áõÙ ¿, ѳëÝ»Éáí ÑëÏÇã
93
§ÈdzñÅ»ù ³¹³åï³ódz¦, û± §ÁÝï»É³óáõÙ¦: гñó³¹ñÙ³Ý Ñ³Ù»Ù³ï³µ³ñ ëïáõÛ·
Ó»éù µ»ñ³Í ѳñÙ³ñáõÙÝ, ϳÙ, ³í»ÉÇ ×ß·ñÇïª ÁÝï»É³óáõÙÁ, ³Ûë ¹»åùáõÙ ß³ï ³í»ÉÇ
ë³ÑٳݳÛÇÝ É³ñí³ÍáõÃÛ³Ý:
94
9. º¼ð²Î²òàôÂÚàõÜܺð
ÙÇÝã¹»é áõß Å³ÙÏ»ïáõÙ (70 ûñ) ³é³í»É ïÇå³Ï³Ý ¿ §÷áëÇϳíáñ ï³Ëï³ÏǦ ûëïÇ
óáõó³ÝÇßÝ»ñÇ:
¹ÇëýáõÝÏódzÛÇ Ù³ëÇÝ:
95
½³Ý·í³ÍÇ Ñ³í³ëïÇ Ýí³½áõÙ: àõÕ»ÕÇÏÇ ÑÛáõëí³ÍùÇ ½³Ý·í³ÍÁ ¹ñë¨áñáõÙ ¿ Ýí³½Ù³Ý
ÙÇïáõÙ:
96
10. ¶ð²Î²ÜàôÂÚ²Ü ò²ÜÎ
1. ܳí³ë³ñ¹Û³Ý ¶.²., ºñÇóÛ³Ý Ü.´., ¶ñÇ·áñÛ³Ý ².ê., ê³ý³ñÛ³Ý Î.ê. ÐÇåáϳÙå ¨ áõÕ»ÕÇÏ-
6. Акопян В.П., Симонян М.А., Манукян А.А., Симонян Р.М., Акопян А.А. Дисбаланс
8. Ашрафян К.Б., Баласанян М.Г. Участие системы оксида азота в развитии тревожности,
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