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The monograph gives a description of human organism great adaptation abilities during a long-lasting period in space, in the state of hunger, cold, heat, as well as in different pathological states and so on. The paper elucidates the evolution of stress theory, its classification, morphofunctional changes of organs in the state of stress. The authors discuss biological stresses, physiological (temperature, alimentary, respiratory, cosmic, stress in pregnancy, sport), pathological (chemical, emotional, painful, shock state), pathological processes with stress reaction (infection, fever reaction, anaphylaxis, allergy), pathological disturbances and stress (cardiovascular, endocrine, digestive), stress reactions  conditions of the technico-scicntific progress. The book is meant,for. readers interested, in the problems of stress. It will be of great help for biologists, physiologists, medical workers and students of medical institutes. 5F5=78@>20;8 : @5:><5=4>20;8 : 8740=8N :0=4. 18>;. =0C: . . 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ogt 8 Long (1952), . !. 4E0=0 (1961) 8 4@. AG8B0NB 2K45;5=85. " @57C;LB0B>< 0:B820F88 A8<?0B>-04@5=0;>2>9 A8AB5<K. 0@0AB0=85 F8@:C;8@CNI53> 04@5=0;8=0 2 :@>28 AB8<C;8@C5B 2K45;5=85 ". !>3;0A=> . !. 0E0=0 (1961), 2 ?5@2>9 D075 AB@5AA0 >B<5G05BAO 2K45;5=85 :0B5E>;0<8=>2, 3>@<>=>2 I8B>284=>9 65;57K, 3>@<>=>2 704=59 4>;8 38?>D870 (@8A. 1). > 2B>@>9 D075 ?>2KH05BAO 2K45;5=85 ", :>@B8-:>AB5@>84>2, 8=AC;8=0. 0 ?OB>< MB0?5 8AA;54>20=89 87CG0;8AL A;>6=K5 <5E0=87<K, ?@>8AE>4OI85 2 38?>B0;0<CA5, :>B>@K5 >1JOA=ONB ?@52@0I5=85 =5@2=>3> AB8<C;0 2 =59@>3C<>@0;L-=K9, 3>@<>=0;L=K9. @8 MB>< 206=CN @>;L 83@0NB 04@5=5@38G5A:0O, E>;8=5@38G5A:0O 8 A5@>B>=8=5@38G5-A:0O A8AB5<K 38?>B0;0<CA0 (. !. 0E0=0, 1978; . . 8@C, 1978). K ?@8405< >A>1>5 7=0G5=85 8AA;54>20=8O< @5F5?B>@>2 =59@>=>2 2KH5C:070==KE AB@C:BC@ 38?>B0;0<CA0. 07;8G0NB >A<>@5F5?B>@K, B5@<>@5F5?B>@K, 3;8:>@5F5?B@@K, @5F5?B>@K, @5038@CNI85 =0  1 8 8A. I. $07K AB@5AA0. <<>18;870F8O 60 <8=CB. I  :0B01>- ;8G55:0O; li  0=01>;8G5A:0O. 1  :0B5E>;0<=OK; 2 - B0@5>- 84=K5 3>@<>=K: 3  "; 4  :>@B?:>AB5@A84K 9 3>@<>=K (:>@B8:>AB5@>84K, B8@5>=4=K5 8 ?>;>2K5 ropj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llen et al. (1970). =8 @0745;ONB AB@5AAK =0 425 3@C??K; A8AB5<=K5 8 =5@2=K5.  ?5@2>9 3@C??5 @07;8G0NB 2 ?>43@C??K: 0) AB@5AAK, 2>7=8:0NI85 ?@8 87<5=5=8OE 2=CB@5==59 A@54K (=0?@8<5@, D878>;>38G5A:85 A42838 2 @57C;LB0B5 2;8O=8O D0@<0:>;>38G5A:8E 25I5AB2); 1) AB@5AAK, 2>7=8:0NI85 ?>A;5 2@54=KE 459AB289 2=5H=8E D0:B>@>2. B>@0O 3@C??0 ?>4@0745;O5BAO =0 4 ?>43@C??K: 0) AB@5AAK, 2K720==K5 2@54=K<8 2>7459AB28O<8, ?>@060NI8<8 >@30=87< 8 ?>@>640NI8<8 1>;52K5 O2;5=8O; 1) 2;8O=85< A25B0; 2) 72C:0; 3) M<>F89. . . @H02A:89 A A>02B. (1977 1980) AG8B0NB, GB> A ?>=OB85< 040?B0F88 A;54C5B A2O7K20BL D878>;>3>-<>@D>;>38G5A:85 87<5=5=8O 2 >@30=87<5. -B0 @50:F8O >ACI5AB2;O5BAO 2 >B25B =0 459AB285 AB@5AA>2KE @074@068B5;59 A@54K, :>B>@K5 =07K20NBAO D878>;>38G5A:8<8. >4>1=0O 040?B82=0O @50:F8O A>AB>8B 87 B@5E D07: 1) 0=01>;8G5A:>9, 2) :0B01>;8G5A:>9, >15A?5G820NI59 2>7<>6=>ABL >ACI5AB2;5=8O >G5@54=>9 0:B82=>AB8, 3) 871KB>G=>-0=01>;8G5A:>9. 2B>@ @07;8G05B D878>;>38G5A:89 AB@5AA, :>340 ?>2KH0NBAO =53M=B@>?89=K5 2>7<>6=>AB8 @072820NI53>AO >@30=87<0, 8 ?0B>;>38G5A:89 AB@5AA, :>340, =0?@>B82, >=8 A=860NBAO.  0=B5=0B0;L=>< ?5@8>45 . . @H02A:89 (1980) @07;8G05B D878>;>38G5A:85 38?>:A5<8G5A:=5 AB@5AAK.  @0==89 ?>AB=0B0;L=K9 ?5@8>4 206=K5 AB@5AA>2K5 @5- ? 0:F88 ?@>8AE>4OB ?>4 2;8O=85< B5<?5@0BC@K 2=5H=59 A@54K. 2B>@ C:07K205B =0 B>, GB> ?@8 @50;870F88 0=-| B83@028B0F8>==KE @50:F89 D878>;>38G5A:89 AB@5AA A>?@>2>6405BAO B8<8:>-<8B>B8G5A:8<, 0 ?0B>;>38G5A:89 j B8<8:>-;8B8G5A:8< MDD5:B><.  ?5@8>4 ?>A;5 @50;870F88 C 682>B=KE ?>7K AB>O-j =8O D878>;>38G5A:89 AB@5AA 7028A8B >B >1J5<0 42830B5;L=>9 0:B82=>AB8. 07;8G0NB M2@8>18>=B=KE 682>B=KE A 2KA>:>9 42830B5;L=>9 0:B82=>ABLN 8 AB5=>18>=-B=KE  A >3@0=8G5==>9 42830B5;L=>9 0:B82=>ABLN. (. . @H02A:89, . . 5<5F, 1968; . . >70=>20, . . 070@8=0, . . @H02A:89, 1978). ! =0H59 B>G:8 7@5=8O, ?@82545==K5 :;0AA8D8:0F88 =5 8AE>4OB 87 4>AB0B>G=> G5B:8E :@8B5@852. 07=>>1@07L =K5 AB@5AAK >4=>2@5<5==> ?@82>4OBAO 8 2 B5E 8 2 4@C38E 3@C??0E. 0: 8725AB=>, 3;02=K<8 4;O :;0AA8D8:0F88 4>;6=K 1KBL :@8B5@88, ?>72>;ONI85 @0AA<0B@820BL @07=>>1@07=K5 D>@<K AB@5AA0. >A:>;L:C AB@5AA ?>A;54AB285 >?@545;5==KE 2>7459AB289 =0 >@30=87< 8 =0 &! 682>B=KE 8 G5;>25:0, <>6=> AG8B0BL, GB> AC-| H5AB2CNB 2 >A=>2=>< B@8 3@C??K AB@5AA>2: 0) 18>;>38G5A:85; 1) D878>;>38G5A:85; 2) ?0B>;>38G5A:85. @8 MB>< =5>1E>48<> CG5ABL, GB> AB@5AA>2K5 A>AB>O=8O ?@>-1 O2;ONBAO ?>-@07=><C 2 7028A8<>AB8 >B C@>2=O @0728B8O] 859@>M=4>:@8==>9 A8AB5<K C @07;8G=KE 284>2 8 3@C??! 682>B=KE. !B@5AA>2K5 A>AB>O=8O C 682>B=KE A ?@8<8! B82=>9 =5@2=>9 A8AB5<>9, C :>B>@KE =5 AD>@<8@>20=K' 38?>B0;0<CA 8 38?>D87, >B;8G0NBAO >B B0:>2KE A 38?>-] B0;0<>-38?>D870@=>9 A8AB5<>9. 1. 8>;>38G5A:85 AB@5AAK =01;N40NBAO C 682>B=KE; A ?@8<8B82=>9 =5@2=>9 A8AB5<>9. 07;8G=K5 D>@<K] AB@5AA0 ?@>O2;ONBAO 2 7028A8<>AB8 >B 2;8O=8O @07=>>1-) @07=KE M:7>35==KE D0:B>@>2 (@5A?8@0B>@=>3>, B5<?5! @0BC@=>3>, >A<>B8G5A:>3>). # B0:8E 682>B=KE 8<55BAO 30=3;8>=0@=>-A5B5284=0O =5@2=0O A8AB5<0, 3><5>AB0B8-G5A:85 <5E0=87<K =5AB>9:85. 8?>B0;0<8G5A:0O >1;0ABL 8 38?>D87 >BACBAB2CNB. 8>;>38G5A:85 AB@5AAK 83@0NB 206=CN @>;L 2 040?B0F88 8 M2>;NF88 682>B=KE. ; B0:8E AB@5AA0E ?@5>1;040NB 0=01>;8G5A:85 @50:F88, G0AB> =01;N405BAO ?0@018>7. 2. $878>;>38G5A:85 AB@5AAK >B<5G0NBAO C 682>B=KE A @0728B>9 38?>B0;0<>-38=>D870@=>9 A8AB5<>9 A 2@5<5==K< =0@CH5=85< ?@>F5AA>2 3><5>AB070 8 ?>A;5- 12 4CNI8< 53> 2>AAB0=>2;5=85<. @>O2;5=85 D878>;>38G5A:8E AB@5AA>2 2 7=0G8B5;L=>9 AB5?5=8 >?@545;O5BAO @07;8G=K<8 D>@<0<8 >1<5=0 25I5AB2 8 A>AB>O=85< @07;8G=KE DC=:F89 >@30=87<0. !@548 D0:B>@>2, 8E 2K7K20NI8E, =081>;55 8725AB=K 3>;>4, 38?>B5@<8O, 60640, =0@CH5=8O :@>2>>1@0I5=8O, 4KE0=8O, DC=:F88 =5@2=>9 A8AB5<K 8 B. 4.  D878>;>38G5A:8E AB@5AA0E ?>A;54>20B5;L=> ?@>B5:0NB :0B01>;8G5A:0O 8 0=01>;8G5A:0O @07K  AB@5AA>2K5 <5E0=87<K CG0AB2CNB 2 ?@>F5AA0E A0<>@53C;OF88 8 >15A?5G820NB @07=>>1@07=K5 D>@<K 3><5>AB070, 2 >A=>2=>< M=4>:@8==K9, >1<5==K9, =59@>-2535B0B82=K9. ;02=K< @53C;OB>@>< O2;O5BAO 38?>B0;0<CA, ;8<18G5A:85 AB@C:BC@K 8 :>@0 3>;>2=>3> <>730, 83@0NI85 ?CA:>2CN 8 :>@@838@CNICN @>;O, 3;02=K< MDD5:B>@><  B@8 MB060 F5=B@>2 38?>B0;0<CA0. 5@2K9 MB06  =081>;55 4@52=89, >= @53C;8@C5B 3>-<5>AB07 M=4>:@8==>9 A8AB5<K 8 ?@54AB02;O5B A>1>9 [A2>5>1@07=CN @5B8:C;O@=CN AB@C:BC@C, A2O7K20NICN =59@>A5:@5B>@=K5 F5=B@K. B>@>9 MB06 C?@02;O5B | >1<5=>< 25I5AB2 8 8<55B B5A=CN A2O7L A F5=B@0<8, @53C;8@CNI8<8 38?>D870@=>-M=4>:@8==K5 ?>4A8AB5<K. ("@5B89 MB06  MB> A;>6=0O @5B8:C;O@=0O AB@C:BC@0, :>>@48=8@CNI0O F5=B@K, C?@02;ONI85 M=4>:@8==K<8 8 >1<5==K<8 ?@>F5AA0<8 (. !. 0E0=0, . . 5;L=8:, 1978). 3. 0B>;>38G5A:85 AB@5AAK E0@0:B5@87CNBAO ?@>4>;68B5;L=K< =0@CH5=85< 3><5>AB070. 07;8G0NB AB@5AAK, ?@>B5:0NI85 A =0@CH5=85< ?@>F5AA>2 3><5>AB070, ?5@28G=> 2K7K20NI85 ?0B>;>38N (8=>340 =0@CH5=8O =5>1@0B8<K 8 254CB : A<5@B8), 8 AB@5AAK, 2K720==K5 ?0B>;>38G5A:8< ?@>F5AA><  >B@02;5=85<, 8=D5:F859, 3@02<>9 8 4@. @8 AB@5AA0E =01;N40NBAO A?5F8D8G5A:85 <>@D>;>38G5A:85, >1<5==K5 8 >?@545;5==K5 DC=:F8>=0;L=K5 87<5=5=8O. 4.  $$#&,+ /        !" !! I5 !5;L5 (1953 1960) ?>:070;, GB> =04?>G5G=8:8 >@30=K B8<8:>-;8<D0B8G5A:>9 A8AB5<K, :;5B>G=K9 A>-AB02 :@>28 >1;040NB 2KA>:>9 GC2AB28B5;L=>ABLN : 459 AB28N ;N1>3> AB@5AA>@=>3> D0:B>@0 8 >B25G0NB =0 =535 >?@545;5==K<8 <>@D>DC=:F8>=0;L=K<8 87<5=5=8O<8 "0:, ?>A;5 AB@5AA0 :>@0 =04?>G5G=8:>2 7=0G8BA;L= C25;8G8205BAO 2 @07<5@0E. 01;N405BAO 38?5@;8?>849 70F8O F8B>?;07<K :;5B>:, >A>15==> :;C1>G:>2>9 7>=K ?0@0;;5;L=> A C25;8G5=85< :>;8G5AB20 =59B@0;L=K 68@>2, E>;5AB5@8=0, 53> MD8@>2 8 A2O70==KE A =8< ;8 ?84>2.  40;L=59H5< :>;8G5AB2> ;8?84>2 C<5=LH05BAO  AB0488 B@52>38 2 ?CG:>2>9 7>=5 C25;8G8205BAO >1J5< O45@ 8 8=3818@C5BAO <8B>B8G5A:0O 0:B82=>ABL.  AB; 48= @578AB5=B=>AB8 38?5@B@>D8O O45@ C45@68205BAO =0 AB018;L=>< C@>2=5, <8B>B8G5A:0O 0:B82=>ABL AB8<C ;8@C5BAO.  AB0488 8AB>I5=8O 38?5@B@>D8O O45@ >A;0-1520AB, 0 <8B>B8G5A:0O 0:B82=>ABL >?OBL A=8605B5 (. II. 8@8;;>2 A A>02B., 1973). B<5G05BAO ?>;=0 45;8?>84870F8O :;5B>: :>@K =04?>G5G=8:>2, C:07K20N I0O =0 8E 8AB>I5=85 (. . 0;8=8=0, 1973). C48>35==K9 AB@5AA C :@KA 2K7K205B C25;8G5=85 A>45@60=8O  2 =04?>G5G=8:0E A <0:A8<C<>< G5@57 8 12 G. AE>4=K9 C@>25=L 2>AAB0=02;8205BAO G5@57 48 G (. . 0:>2552, 1977).  >?KB0E . . 0;8=8=>9 (1973) 45;8?>84870F8O :;5B>: :>@K =04?>G5G=8:>2 8<5;0 >G03>2K9 E0@0:B5@ 2B>@ >B<5G0;0 B0:65 >B5: :;5B>:, 2>;>:=5=85 :0?AC; 8 AB@><K. !>AC4K 65;57K 1K;8 @0AH8@5=K A> 7=0G8-B5;L=> =5@02=><5@=K< ?>;=>:@>285<, 2AB@5G0;8AL B@><-1K, =01;N40;8AL @50:F8O =0 =59B@0;L=K5 <C:>?>;8A E0@84K ?@8 8=B5=A82=>9 20:C>;870F88 :;5B>:, ?CABK5 :;5B:8 A ?8:=>B8G=K<8 O4@0<8. !5:@5B>@=0O 0:B82 =>ABL <>73>2>3> 25I5AB20 =04?>G5G=8:>2 A>?@>2>640 14 ABAO 8=B5=A82=>9 20:C>;870F859 :;5B>: 8 C<5=LH5=85< AC<<0@=KE :0BAE>;0<8=>2.  >?KB0E . . "8E>=>2>9 (1973) CA8;820;0AL D;N>@5AF5=F8O :;5B>: <>73>2>3> >B45;0, A>45@60I8E =>@04@5=0;8=. @8 <=>3>:@0B=KE >1>@>=8B5;L=KE @50:F8OE C 682>B=KE <K =01;N40;8 <>@D>;>38G5A:85 8 38AB>AB@C:-BC@=K5 87<5=5=8O I8B>284=>9 65;57K, 2K@060NI85AO 2 C25;8G5=88 55 >B=>A8B5;L=>3> 8 01A>;NB=>3> 25A0, ?>O2;5=88 =51>;LH8E D>;;8:C;>2, @57>@1F88 :>;;>840, =0@0AB0=88 2KA>BK M?8B5;8O (. !. 0E0=0, 1960; $. . $C@4C9, 1980 8 4@.). !>3;0A=> !5;L5 (1934), ?>4 2;8O=85< :>@B8:>AB5-@>84>2 01A>;NB=K9 8 >B=>A8B5;L=K9 25A 28;>G:>2>9 65;57K C<5=LH05BAO. >;8G5AB2> B8<>F8B>2 B0:65 C<5=LH05BAO.  =0AB>OI55 2@5<O CAB0=>2;5=>, GB> ;8<-D>F8B>;87  >48= 87 E0@0:B5@=KE ?>:070B5;59 CA8;5=8O 0:B82=>AB8 :>@K =04?>G5G=8:>2. >4>1=K5 87<5=5=8O >B<5G0;8 8 2 A5;575=:5. > 40==K< . . !028=0 A A>02B. (1973), ?@8 AB@5AA5, 2K720==>< 459AB285< ?>AB>O==KE <03=8B=KE ?>;59, 2 D>;;8:C;0E A5;575=:8 C<5=LH05BAO :>;8G5AB2> ;8<D>F8B>2, :0@8>@A:A8A >BBO38205BAO : F5=B@0< D>;;8:C;>2.  B8<CA5 8 2 A5;575=:5 >B<5G05BAO 38?5@?;078O ?;07<0B8G5A:8E :;5B>:. -. . 8@8;>2, !. . 8 8 . . .@35=A (1973) ?>A;5 <>18;870F88 =01;N40;8 ?@5>1@07>20=85 G0AB8 ?>;8-B;>84=KE :;5B>: 2 42CO45@=K5 8 B>@<>65=85 <8B>B8G5-6>9 0:B82=>AB8. . . '5@=CE A A>02B. (1975) ?>:070;8, GB> 2 3><5>AL0B8G5A:8E @50:F8OE ?@8 AB@5AA0E 87<5=O5BAO A>AC48-AB0O A8AB5<0 >?@545;5==>9 B5@@8B>@88. . - C;038=, .. >20;52, . . @82>@CG5=:>, ! . (5@5?5B>2-A:0O (1980) 2KO28;8 >?@545;5==K5 70:>=><5@=>AB8 @5-B8>=0@=>3> ?5@5@0A?@545;5=8O :@>28 ?@8 459AB288 =0 >@30=87< AB@5AA>@>2 @07;8G=>9 BO65AB8. 2B>@K >?8-AK20NB G5BK@5 B8?0 87<5=5=8O @0A?@545;5=8O :@>28 2 >@30=0E: I B8?  C :@KA A>45@60=85 :@>28 C<5=LH05B-AO 2> 2=CB@5==8E >@30=0E 682>B0 8 <0;>3> B070 (B>=-:0O :8H:0, B>;AB0O :8H:0, A5;575=:0, ?>465;C4>G=0O 65;570), 2 <KH5G=KE 8 :>AB=KE B:0=OE 682>B0 8 <0-;>3> B070, ?5@54=8E 8 704=8E :>=5G=>AB59, 2 :>65 704-=8E :>=5G=>AB59 8 3@C48; II B8?  2> 2=CB@5==8E >@30-=0E; III B8?  2 ?5G5=8; IV B8? =01;N405BAO ?@8 B5@-<8=04L=KE AB048OE H>:0: 2 >@30=0E 3@C48 A:0?;8205B- 15 AO :@>2L, >A>15==> 2 ;53:8E. #:070==K5 87<5=5=8O ?@> 8AE>4OB 2 @57C;LB0B5 CG0AB8O 38?>B0;0<>-38?>D870@=>-04@5=0;>2>9 A8AB5<K. @8 AB@5AA5 E0@0:B5@=K <>@D>;>38G5A:85 87<5=5=8O :@>28  ;8<D>?5=8O 8 M>78=>?5=8O. "0:, A>3;0A=> . . "8E>=>2C (1973), ?@8 38?>:8=5788 =0 20-9 45=L } :@KA A>45@60=85 M>78=>D8;>2 C<5=LH05BAO =0 79,7 ?> A@02=5=8N A :>=B@>;5<. >A;5 0C48>35==>3> AB@AAA3 2 ?5@2K5 60 80 <8= 2 35?0B>F8B0E A=8605BAO :>;8 G5AB2> , 0 G5@57 2 G 8AE>4=K9 C@>25=L 2>AAB0=02] ;8205BAO (. . 0:>2552, 1977).  =59@>=0E 2B>@>9 A;>O 28A>G=>9 :>@K 2 ?5@2K5 12 G 459AB28O AB@5AA0 :> ;8G5AB2>  C<5=LH05BAO.  3;80;L=KE :;5B:0E 87<5=5=89 02B>@ =5 >1=0@C68;. 4=0:> 2 3;80;L=KE :;5B:0E <>765G:0 A>45@60=85  A=878;>AL  <>B>=59@>=0E A?8==>3> <>730 C@>25=L  ?>2KA8;-AO G5@57 2 G ?>A;5 AC4>@>3 (. . 0:>2552, 1974). !. . @0C45 A A>02B. (1973) ?@8 >6>3>-;CG52>O B@02<5 =01;N40;8 87<5=5=8O M:A:@5B>@=>9 DC=:F88 ?>G5:. # A>10: ?>A;5 M:A?5@8<5=B0;L=>3> =52@>70 . $. @8B:>20 (1966) B0:65 >B<5B8;0 2 >4=8E A;CG0OE ?>;8C@8N, 2 4@C38E  >;83C@8N. -:A?5@8<5=B0;L=K9 AB@5AA <>65B 2K720BL <>@D>;>38G5A:85 87<5=5=8O 2 A5@45G=>9 <KHF5, ?>4>1=K5 8=D0@:BC. C48>35==K9 AB@5AA C <KH59 ?@82>48B : <8:@>35<>@@038O< 2 <>73> 2>9 B:0=8 =04?>G5G=8:>2. . !. 0=:>20 (1973) >B<5 G0;0 <>@D>;>38G5A:85 87<5=5=8O N:AB03;><5@C;O@=>? 0??0@0B0 ?>G5: C :@KA, 2K720==K5 4;8B5;L=K< 459AB 285< CA:>@5=8O. @8 MB>< 0:B82878@>20;0AL @588=-0= 38>B58788>20O A8AB5<0, 2 @57C;LB0B5 G53> ?>2KA8;0A 8=B5=A82=>ABL D8;LB@0F88 2 :;C1>G:0E. !. . @0C4 (1973) 2KO28; ?@8 AB@5AA5 =0 8=B0:B=>< 8 =0 87>;8@2 20==>< 65;C4>G:5 ?> 5945=309<C A5:@5B>@=K5 8 429 30B5;L=K5 @0AAB@>9AB20. 02=> 8725AB=>, GB> ?@8 AB@5A-A5 (M:A?5@8<5=B0;L=> 8 :;8=8G5A:8) 2>7=8:0NB O72L 65;C4:0 8 425=04F0B8?5@AB=>9 :8H:8. #<5AB=> =0?><=8BL, GB> AB@5AA>2K5 A>AB>O=8O ?@> B5:0NB ?>4>1=> ?0@018>B8G5A:8< D070<. . . Haconoi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lum, 1951; Proser, Brown, 1967). > <5@5 8AB>I5=8O 70?0A>2 18>35==KE >@30=8G5A:8E A>548=5=89 2A5 1>;LH55 7=0G5=85 ?@8>1@5B0;8 <5E0=87<K A8=B570, 2>7=8:0NI85 ?CB5< <CB0F89. >AB5?5==> 87 ?5@28G=KE 35B5@>B@>D=KE >@30=87<>2 @0728;8AL D>B>-02B>B@>DK 8 >B45;L=K5 E8<8>02B>B@>DK (Frenekcl, 1958). $>B>A8=B57 >1>30B8; 0B<>AD5@C :8A;>@>4><, 8 3;02=CN @>;L AB0;8 83@0BL 0M@>1=K5 ?@>F5AAK. 3  40;L=59H5< MB8 >@30=87<K 8A?KBK20;8 459AB285 '@07;8G=KE AB@5AA>@=KE D0:B>@>2. "0:, ?5@5E>4 >B 0=0M@>1=KE CA;>289 : 0M@>1=K< 2K720; 18>E8<8G5A:85 A42838. >O28;8AL >@30=87<K, A?>A>1=K5 ?>3;>I0BL !3. >7=8:;8 D>B>E8<8G5A:85 =5>@30=8G5A:85 @50:F88. >74=55 2 :0G5AB25 8AB>G=8:0 07>B0 1K;8 8A?>;L7>20=K =8B@0BK 8;8 0<<80: (Saffran, I960). !CI5AB2>20=85 <=>38E ?>:>;5=89 2 A@545 A =87:8< A>45@60=85< :8A;>@>40 ?@825;> : @0728B8N C 682>B=KE @O40 ?@8A?>A>18B5;L=KE <5E0=87<>2, =0?@8<5@ B0:8E, :>B>@K5 >15A?5G820NB 8E ACI5AB2>20=85 ?@8 =87:>< ?0@F80;L=>< 402;5=88 :8A;>@>40. 5A?8@0B>@=K9 AB@5AA =01;N405BAO ?@8 38?>:A88 8 2K7K205BAO @07=8F59 2 ?0@F80;L=>< 402;5=88 307>2 2 0;L25>;0E ;53:8E (Mitchel, Erdman, 1951). @C38< ?@8<5@>< ?@8A?>A>1;5=8O <>65B A;C68BL 38?5@B@>D8O <KHF ;52>3> 8 ?@02>3> 65;C4>G:>2 A5@4-;1?B ,i) @@-3CI-B8B@ :-?"?@?9 >B=>H5=85 25A0 A5@4F0 : 25AC 17 B5;0 2>7@>A;>. -B> =01;N40;>AL 2 @57C;LB0B5 CA8;5=8O 42830B5;L=>9 0:B82=>AB8 682>B=KE (153, >1>@>=8B5;L=K5 @50:F88). !;54AB285< 2;8O=8O M:>;>38G5A:8E AB@5AA>@?K; D0:B>@>2 O2;O5BAO <83@0F8O @K1. 4=8 02B>@K AG8B0NB, GB> @K1K ?5@2>=0G0;L=> 68;8 2 ?@5A=>9 2>45, 4@C 385  2 A>;>=>20B>9, B@5BL8  2 <>@A:>9 (Romer 1935).  ?>;L7C ?5@2>3> ?@54?>;>65=8O 3>2>@OB =87:0O :>=F5=B@0F8O E;>@8AB>3> =0B@8O 2 :@>28 2A5E A>2@5 <5==KE :>AB8ABKE @K1, =87:>5 A>45@60=85 A>;59 2 :@> 28 E@OI52KE @K1, 0 B0:65 =0;8G85 3;><5@C;O@=KE ?> G5:, E>@>H> ?@8A?>A>1;5==KE : 2K45;5=8N 38?>>A<-> B8G5A:>9 <>G8.  <5;>2>< ?5@8>45 :>AB8ABK5 8 E@OI5 2K5 @K1K, >G5284=>, <83@8@>20;8 87 ?@5A=>9 2>4K O <>@5. =>385 A>2@5<5==K5 :>AB8ABK5 @K1K 2B>@8G=>1 70A5;O;8 ?@5A=K5 2>4K; =5:>B>@K5 65 :>AB8ABK5 30=> 84K 2>>1I5 =5 <83@8@>20;8 87 ?@5A=KE 2>4.  ?@>F5A A5 M2>;NF88 A>25@H0;8AL =5>4=>:@0B=K5 <83@0F88, i B5?5@L =5:>B>@K5 <>@A:85 @K1K 2 >?@545;5==>9 AB5?5 =8 7028AOB >B ?@5A=KE 2>4; >B45;L=K5 3@C??K @K1, =0 ?@8<5@ ;>A>A52K5 (0=04@><=K5 @K1K), 2>72@0I0NBAO 2 =8E 4;O @07<=>65=8O. @C385 @K1K, =0?@8<5@ yrpi (:0B04@><=K5 @K1K), >18B0NB 2 ?@5A=KE 2>40E, => @07 <=>60NBAO 2 >:50=5. K1K ?@8A?>A>1;5=K : CA;>28O< A =87:8< A>45@ 60=85< :8A;>@>40. @8 A=865=88 :>=F5=B@0F88 :8A;> @>40 C<5=LH05BAO I5;>G=>9 @575@2 :@>28, 2 B:0=O; ?>2KH05BAO A>45@60=85 <8>3;>18=0 8 @0AH8@O5B5? A5BL :0?8;;O@>2. %5<>@5F5=B>@K :0@>B84=>3> A8=CA3 AB0=>2OBAO 1>;55 GC2AB28B5;L=K<8 : 38?>:A5<88, GB5! ?@82>48B : CA8;5=8N 4KE0=8O (Rahn, 1947; Ottls, 1953) @C3>9 D0:B>@, 2K7K20NI89 18>;>38G5A:89 AB@5AA,-87<5=5=85 B5<?5@0BC@K 2=5H=59 A@54K. >7459AB285 E>;>40 0:B82878@C5B 8=B5@@5=0;>2CN B:0=L. '8A;< 7>7O=>D8;>2 8 ;8<D>F8B>2 C<5=LH05BAO. !>45@6088* 0A:>@18=>2>9 :8A;>BK 2 MB8E B:0=OE 2>7@0AB05B, GB< A?>A>1AB2C5B ?>2KH5=8N E>;>4>CAB>9G82>AB8.  B5G5 =85 ?5@2KE G0A>2 ?@51K20=8O =0 E>;>45 C25;8G8205B5? A>45@60=85 35<>3;>18=0 8 C<5=LH05BAO >1J5< F8@:C ;8@CNI59 :@>28 (Desmarais, 1957). >A;5 8=B5=A8D8:0F88 B5?;>?@>4C:F88 2>7<>6= AB8<C;8@>20=85 >1I53> >1<5=0, 2K720==>5, 2 G0AB=> AB8, 2K45;5=85< =>@04@5=0;8=0.  M:A?5@8<5=B5 8=J is 5F8@>20==K9 =>@04@5=0;8= >1;0405B 1KAB@K< :0;>@8-35==K< 459AB285<. -B> 459AB285 A>E@0=O5BAO 8 ?>A;5 C40;5=8O 2=CB@5==8E >@30=>2. 1I0O <5B01>;8G5A:0O @50:F8O C <>@A:8E 682>B=KE 7028A8B B0:65 >B 0:B82=>AB8 I8B>284=>9 65;57K 8 :>@K =04?>G5G=8:>2.  B5G5=85 ?5@2KE =5A:>;L:8E G0A>2 ?@51K20=8O =0 E>;>45 2 8=B5@@5=0;>2>9 B:0=8 C25;8G8205BAO A>45@60=85 :>@-B8:>AB5@>84>2. @8 2KA>:8E B5<?5@0BC@0E >:@C60NI59 A@54K =01;N405BAO B5A=0O A2O7L <564C B5@<8G5A:8< 8 2>4=K< 10;0=A0<8. 5:>B>@K5 682>B=K5 70?0A0NB 2>4C 8 70I8I0NBAO >B 60@K, 2?040O 2 ;5B=NN A?OG:C. >2545=G5A:85 @50:F88 A?>A>1AB2CNB =0E>645=8N <5AB A ?>4E>4OI59 B5<?5@0BC@>9. ;03>40@O 18>E8<8G5A:8< :><?5=A0F8O<, 8=B5=A82=>ABL <5B01>;87<0 =5 ?>2KH05BAO B0: @57:>, :0: A;54>20;> >6840BL ?@8 ?5@5=>A5 >@30=87<0 87 E>;>4=>9 A@54K 2 B5?;CN. 0I8B0 >B 2KAKE0=8O B5A=> A2O70=0 A 70I8B>9 >B 459AB28O 2KA>:>9 B5<?5@0BC@K, B5< =5 <5=55 =5:>B>@K5 >@30=87<K, 2 G0AB=>AB8 @O4 =0A5:><KE 8 3><>9>B5@<=KE 284>2 682>B=KE, MDD5:B82=> >E;0640NBAO ?CB5< 8A?0@5=8O.  >1I5< 65 C 682>B=KE <5=LH5 ?@8A?>A>1;5=89 4;O 70I8BK >B B5?;0, G5< >B E>;>40 (Burton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ethier, 1953). @C3>5 ?@>O2;5=85 AB@5AA0  87<5=5=85 ?83<5=B0F88 :>68 C <=>38E 157?>72>=>G=KE (=0?@8<5@, <8<8:@8O :0: 70I8B=0O @50:F8O). Woring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oring, 8 C =5:>B>@KE 3@C?? 15A?>72>=>G=KE; 45 AOB8=>385 @0:8, ?@>8AE>645=85 :>B>@KE >B=>A8BAO i B@80AC, >1;040NB 3>@<>=0;L=>9 @53C;OF859, 0 42C60 15@=K5 (3>;>2>=>385 <>;;NA:8), ?>O282H85AO =5 @0 =55 ?;59>F5=0,  8A:;NG8B5;L=> @5D;5:B>@=>9 @53C;OF859. 5D;5:B>@=K5 ?CB8 (15@CI85 A2>5 =0G0;> i A5BG0B:5 3;070), G5@57 :>B>@K5 >ACI5AB2;O5BAO @53C;OF8O 87<5=5=89 >:@0A:8 C 2A5E @0:>>1@07=KE, =0A5:> <KE, 0<D8189 8 <=>38E 284>2 @K1 8 @5?B8;89, 2>2;5 :0NB 2 ?@>F5AA A2>53> DC=:F8>=8@>20=8O 0:B82=CN A5 :@5F8N <5;0=>D>@=KE 3>@<>=>2. 0: F8@:C;8@CNI6 2 :@>28 3>@<>=K, B0: 8 =5?>A@54AB25==0O 8==5@20F4* 83@0NB 206=CN @>;L C 1>;LH8=AB20 4@C38E 284>2; 8A :;NG5=85<, ?>-2848<><C, O2;ONBAO E0<5;5>= 8 4@C38* @5?B8;88. # :>AB8ABKE @K1 @5D;5:B>@=0O @53C;O88! 8<55B ?@5>1;040NI55 7=0G5=85 (Woring, 1963). !B@5AA>2K5 @50:F88, ?@>O2;ONI85AO G5@57 38?5@ ?83<5=B0F8N :>68, >15A?5G820NB @07=>>1@07=>5 ?@8 A?>A>1;5=85 682>B=KE : >:@C60NI59 A@545. %@><0B>D>@=>9 A8AB5<5 ?@8?8AK20NB >ACI5AB2;5 =85 70I8B=>9 8 03@5AA82=>9 >:@0A:8, DC=:F88 B5@<>@5 3C;OF88 8 70I8BK >B 8=B5=A82=>3> >A25I5=8O 8, =0:> 85F, 1@0G=>3> =0@O40 C A0<F>2 2 ?5@8>4 @07<=>65=! (Yellewag, 1911). # =0A5:><KE =0945=K 3>@<>=0;L=K5 25I5AB20 040 D0- 8 15B0-M:487>8, 83@0NI85 206=CN @>;L 2 ?@>=5A A0E <5B0<>@D>70. 07=>>1@07=K5 D0:B>@K 2=5H=5 A@54K (;CG8AB0O M=5@38O, B5<?5@0BC@0) 2;8ONB =0 <5 B0<>@D>7 8 ;8=L:C =0A5:><KE. -:487>=K A8=B578@CNB AO 87 E>;5AB5@8=0, 8E AG8B0NB ?@54H5AB25==8:0<8 :>@ B8:>AB5@>84>2. K;> CAB0=>2;5=>, GB> <>;5:C;O@=K! 25A M:487>=0 @025= 464, M<?8@8G5A:0O D>@<C;0 C2CH44Q (Car)son, 1964}. 20 5I5AB2>, 1;87:>5 : M:487>=C =0A5:><KE, 1K;> 2K45;5=> 8 87 8E >@30=87<0. >:070=>, GB> M:487>= C =0A5:><KE 2>7459AB2C5B =0 E@><>A><K 8 0:B828@C5B >?@545;5==K5 35=K. @O<>5 2;8O=85 =5:>B>@KE 3>@<>=>2 =0 35=K >1=0@C65=> C 15A?>72>=>G=KE. 5 8A:;NG5=>, GB> 2 18>;>38G5A:8E AB@5AA0E CG0AB2CNB 8 M:487>=K.  =5:>B>@KE 18>;>38G5A:8E AB@5AA0E 1>;LH>5 7=0G5=85 8<5NB DC=:F88 =5@2=>9 8 <KH5G=>9 A8AB5<. ! MB>9 B>G:8 7@5=8O ?@54AB02;ONB 8=B5@5A @50:F88 M;5:B@8G5A:>3> A:0B0. "0:, ?5@540G0 2>71C645=8O 2 =5@2=>-<KH5G=KE A8=0?A0E >ACI5AB2;O5BAO E8<8G5A:8<8 <5480B>@0<8: 0F5B8;E>;8=><, =>@04@5=0;8=><, 5->:A8-B@8?B0<8=><. 5<1@0=0 =5@2=>-<KH5G=>3> A8=0?A0 >B;8G05BAO ?> A2>8< M;5:B@8G5A:8< A2>9AB20< >B ?@>2>4OI59 <5<1@0=K <KH5G=>9 :;5B:8. -;5:B@8G5A:85 >@30=K 8<5NB @07;8G=>5 ?@>8AE>645=85. 5:>B>@K5 87 =8E ?@54AB02;ONB A>1>9 284>87<5=5==K5 <5<1@0=K =5@2=>-<KH5G=KE, 0 4@C385  <KH5G=KE 2>;>:>=. KA>:>5 M;5:B@8G5A:>5 =0?@O65=85, @0728205<>5 =5:>B>@K<8 M;5:B@8G5A:8<8 >@30=0<8, >1JOA=O5BAO AC<<0F859 @07=>AB59 ?>B5=F80;>2, 2>7=8:H8E 2 >B45;L=KE ?>A;54>20B5;L=KE :;5B:0E 2 @57C;LB0B5 0A8<<5B@8G5A:>9 45?>;O@870F88 (Dijkgraaf, 1963).  AB@5AA>2KE @50:F8OE C 15A?>72>=>G=KE 1>;LHCN @>;L 83@0NB <5480B>@K. "0:, 0F5B8;E>;8= >1=0@C65= C ?@>AB59H8E, =0?@8<5@, C @5A=8G=KE 8=DC7>@89 8 B@8=0=>A><, => >BACBAB2C5B C =5?>4286=KE ?;07<>4852 (A?>@>28:>2). K;> ?>:070=>, GB> B:0=8 MB8E >@30=87<>2 A>45@60B A?5F8D8G5A:CN 0F5B8;E>;8=MAB5@07C, @0AI5?;ONICN 0F5B8;E>;8= A> A:>@>ABLN 0,08 <:3/G "0 1 <3 ACE>3> 25I5AB20 (Seamen, 1951). "@8?0=>A><K A8=B578@CNB 4> 71,5 <:3 0F5B8;E>;8=0 =0 I 3 ACE>3> 25I5AB20 70 75 <8= ?@8 B5<?5@0BC@5 37,64 (Bulbring, 1954). 2865=85 ?5G5=>G=>9 42CCAB:8 (Faciola hepatica) ?>402;O5BAO 0F5B8;E>;8=>< :0: 2 =>@<5, B0: 8 ?@8 C40;5=88 30=3;852. -B8 ?;>A:85 G5@28 A>45@60B >B 0,19  1,7 <:3 0F5B8;E>;8=0 =0 I 3 AK@>3> 25I5AB20, 0 B0:65 0F5B8;E>;8=MAB5@07C 8 E>;8=0F5B8;B@0=AD5@07C. 5<5@B8=K A>45@60B <=>3> E>;8=MAB5@07K (Chance,  A5=A>@=KE =5@2=KE :;5B:0E 0==5;84 =0945= 0F5-2OE>;8= (Umreth, 1952). 5@2K, 8=85@28@CNI85 =5:>- 21 B>@K5 <KHFK 0==5;84, A8?C=:C;84 8 A25BOI85AO >@30 =K 0==5;84, E>;8=5@38G=K.  @5A=8G=KE :;5B:0E <>; ;NA:>2 >?@545;5=K 0F5B8;E>;8= 8 E>;8=MAB5@07; (Bulbring, 1954).  30=3;8OE <>730 3>;>2>=>38E <>;;NA:>2 A>45@68B AO <=>3> 0F5B8;E>;8=0; 2 <>73C >AL<8=>30 53> :>=F5= B@0F8O A>AB02;O5B 77 <:3/r; <>73 :0;L<0@0 384@>;87C5 200 400 <3 0F5B8;E>;8=0 2 I G/100 <3 15;:0. & 2;8O=85< E>;8=0F5B8;B@0=AD5@07K 2 3>;>2=>< 30=3;8 0F5B8;E>;8= A8=B578@C5BAO A >G5=L 2KA>:>9 A:>@>ABLN 40 80 <:<>;L/G =0 I <3 15;:0 (Berman, 1953). F5B8;E>;8=>2K5 =59@>=K =0A5:><KE 8AA;54>20; 2 A2O78 A B5<, GB> <=>385 8=A5:B8F84K O2;ONBAO 0=B* E>;88MAB5@070<8.  3>;>25 <54>=>A=>9 ?G5;K =0945= <=>3> 0F5B8;E>;8=0; :@><5 B>3>, 5ABL 8 4@C3>9 E>;8=> 2K9 MD8@, 0 2>7<>6=>, 8 420 (Augustinson, 1954; Met calf, 1956).  @07;8G=KE AB@5AA>2KE @50:F8OE 1>;LH>5 7=0G5=8 83@0NB :0B5E>;0<8=K, 04@5=0;8= 8 =>@04@5=0;8=. Tai 2 =043;>B>G=>< 30=3;88 4>6452>3> G5@2O >1=0@C65=! E@><0DD88=K5 :;5B:8. -:AB@0:BK 87 =8E 40NB F25B =CN @50:F8N =0 04@5=0;8=. 5@2=0O A8AB5<0 G5@2O 55 45@68B 1,4 <:3 04@5=0;8=0 8 0,32 <:3 =>@04@5=0;8= ?0 I 3 AK@>3> 25I5AB20.  A;N==KE 65;570E >AL<8=>3! >1=0@C65=> 1  10 <:3 =>@04@5=0;8=0 =0 1 3 AK@>3 25I5AB20 8 A>2A5< =5 =0945=> 04@5=0;8=0.  =5@2=> A8AB5<5 101>G:8 A>45@68BAO 0,19 <:3 =>@04@5=0;8=0 <5=55 0,01 <:3 04@5=0;8=0 =0 1 3, 0 2 =5@2=>9 A8AB5<C ;8G8=:8 6C:0  1,3 2,0 <:3 04@5=0;8=0 =0 1 3 =ep2-=>9 B:0=8 (Ostlang, 1954).  @50:F8OE AB@5AA0 C 15A?>72>=>G=KE >?@545;5==CN @>;L 83@05B 8 <5480B>@ 5->:A8B@8?B0<8=. -B> 25I5AB2> 845=B8G=> A5@>B>=8=C (A>AC4>AC6820NI55 25I5AB2 A>45@60I55AO 2 AK2>@>B:5) 8 M=B5@0<8=C, 2K45;5==>< 87 M=B5@>E@><0DD8==KE :;5B>: A;N==KE 65;57 3>;>2! =>38E <>;;NA:>2 (Page, 1954). 5-:A8B@8?B0<8= >1@0-7CABAO ?>4 459AB285< B@8?B>D0=45:0@1>:A8;07K 2 5->:A8B@8?B>D0=0. = A8=B578@C5BAO 2 1>;LH8E :>;! G5AB20E 2 A;878AB>9 65;C4>G=>-:8H5G=>3> B@0:B0 8 ?>465;C4>G=>9 65;575 C 0:C;>2KE @K1 (Eccles et a 1954) 8 ?@54AB02;O5B A>1>9 25AL<0 MDD5:B82=K9 AB: <C;OB>@ F5=B@0;L=>9 =5@2=>9 A8AB5<K (Von Eule 1958). 5-:A8B@8?B0<8= =0945= 2 B:0=OE 0AF8489, ? 22 @54=56015@=KE <>;;NA:>2, 2 :>6=KE 65;570E O4>28BKE 0<D8189, 2 IC?0;LF0E :8H5G=>?>;>AB=KE (Erspa-mer, 1954). = A;C68B <>I=K< AB8<C;OB>@>< 4;O A5-B5284=>9 D>@<0F88 :8H5G=>?>;>AB=KE (Rois, 1957), C :>B>@KE >1=0@C65= 8 B8@0<8=. # =5:>B>@KE ?;0AB8=G0B>6015@=KE <>;;NA:>2 5->:A8B@8?B0<8= <>65B 459AB2>20BL :0: ?5@540BG8:, CA:>@ONI89 45OB5;L=>ABL A5@4F0 (Weleh, 1958). @8 2>7459AB288 48MB8;0<840 ;875@38=>2>9 :8A;>BK =0 ?0C:>2 =0@CH05BAO >1KG=K9 A8<<5B@8G=K9 @8AC=>: ?0CB8=K (Witt, 1951). # :>H5: MB> 25I5AB2> 2K7K205B A>AB>O=85 O@>AB8, C 3C??8 8 C;8B>:  ?>AB>O==CN ;>:><>B>@=CN 0:B82=>ABL. !5@>B>=8= ?@82>48B 0:B82=KE @K1 2 A>AB>O=85 ?>:>O, 0 ?>A;54CNI55 2>7459AB285 48MB8;0<84>< ;875@38=>2>9 :8A;>BK >A;01;O5B 53> 2;8O=85 (Abramson, 1954; Tront, 1957).  18>;>38G5A:8E AB@5AA0E C 15A?>72>=>G=KE CG0AB2CNB 8 4@C385 25I5AB20, =0720==K5 2>71C640NI8<8 D0:B>@0<8 (Abbie, 1940). %8<8G5A:8 >=8 =5 845=B8D8F8@>20=K (Florey, 1960). "0:, D0:B>@ , 2K45;5==K9 87 A5=A>@=KE =5@2>2 8 87 A<5H0==KE =5@2>2 <=>38E 15A?>72>=>G=KE, ?>1C6405B ?G5;C :@C68BLAO 8 2>71C6405B @5F5?B>@K @0ABO65=8O C @0:0. !E>4=K9 M:AB@0:B 2K7K205B A>:@0I5=85 @07;8G=KE <KHF C 0==5;84 (Umrath, 1952). 7 B:0=59 F5=B@0;L=>9 =5@2=>9 A8AB5<K ?>72>=>G=KE 2K45;5= D0:B>@, :>B>@K9 ?>402;O5B @07;8G=K5 DC=:F88 =5@2=>9 A8AB5<K # @0:0. ">@<>7=K5 =5@2K, 84CI85 : A5@4FC 8 : @5F5?B>@0< @0ABO65=8O, C @0:0 A>45@60B D0:B>@, :>B>@K9 AE>45= A 30<<0-0<8=><0A;O=>9 :8A;>B>9, => =0<=>3> 0:B82=55 55 (Florey, 1960). 0<<0-0<8=><0A;O=0O :8A;>B0 >?@545;O5BAO 2 =5@2=>9 B:0=8 <=>38E 682>B=KE; 2 A5@>< 25I5AB25 55 1>;LH5, G5< 2 15;><, 0 2 :>@5 3>;>2=>3> <>730 =0<=>3> 1>;LH5, G5< 2 ?5@8D5@8G5A:8E =5@20E. =>3> 55 2 30=3;8OE ><0@0. 0<<0-0<8=><0A-;O=0O :8A;>B0 C3=5B05B @5F5?B>@K @0ABO65=8O <KHF 0:0, => =5 2KO2;O5BAO 2 ?5@DC70B5 2> 2@5<O =5@2=>3> B>@<>65=8O (Edwards, 1959). 0<<0-0<8=><0A;O=0O :8A;>B0 2K7K205B B>@<>65=85 2 <KHF0E A5@4F0 C @0:0, 8 MB> B>@<>65=85 1;>:8@C5BAO ?8:@>B>:A8=>< :0: 8 B>@<>65=85, 2K720==>5 @074@065=85< B>@<>7=>3> =5@20 .(Florey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| 3@81>284=KE B5;0E G;5=8AB>=>38E. # =0A5:><KE AB5@5>B8?=>ABL ?>2545=8O A:;04K205B A O 87 4>2>;L=> A;>6=KE 2@>645==KE @5D;5:A>2. !@548 15A?>72>=>G=KE 040?B0F8>==K< ?>2545=85< 2K45;O5BAO >AL<8=>3. ';5=8AB>=>385 C=8:0;L=K 2 B>< >B=>H5=88, GB> @07=>>1@0785 8E ?>2545=8O >1CA;>2;5= 45OB5;L=>ABLN =51>;LH>3> G8A;0 =59@>=>2. !;>6=>ABL ?>2545=8O =5 ?@>?>@F8>=0;L=0 G8A;C =59@>=>2, A>AB>8B 87 D8:A8@>20==KE B8?>2 @50:F89 8 ?>A;54>20B5;L=KE 0:B>2- =8 <>3CB @53C;8@>20BLAO AB@>3> ;>:0;87>20= =K<8 =5@2=K<8 F5=B@0<8, =0?@8<5@ AB@5:>B0=85 ] A25@G:0 2K7K205BAO @074@065=85< ;>:0;L=KE B>G5: i =043;>B>G=>< 30=3;88 (Prosser, 1957). @><5 AB@5AA>2KE @50:F89, :>B>@K5 =01;N40NBAO C 15A?>72>=>G=KE 8 @K1 ?>4 2;8O=85< 5AB5AB25==KE D0:B>@>2, B0:85 65 87<5=5=8O <>6=> 2K7=0BL M:A;5@8<5= B0;L=>. "0:, Kromlik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| 2;8O=85< B5<?5@0BC@=KE :>;510=89, 3>;>40=8O, 606! 4K, 38?>:A88, D878G5A:8E =03@C7>:, M<>F8>=0;L=KE ?^ @56820=89 8 B. 4. 0B5< 3><5>AB07 2>AAB0=02;8205B5 . X. 0@:028, . . 20:8=0, . . #:>;>2; (1968 1980) CAB0=>28;8, GB> 2 7028A8<>AB8 >B A8; 2>7459AB289 2 >@30=87<5 <>3CB @072820BLAO @07;8G8 D878>;>38G5A:85 @50:F88: 1) ?@8 A;01KE 2>7459AB2 OE  @50:F8O B@5=8@>2:8, 2) A@54=59 A8;K  @50:F 0:B820F88, 3) A8;L=KE  @50:F88 AB@5AA0. 2B>@K 87CG8] ;8 2;8O=85 =0 >@30=87< 04@5=0;8=0, <03=8B=>3> ?>;. <C<8Q 8 =01;N40;8 0:B820F8N M;5<5=B>2 @5B8:C;O =>9 D>@<0F88, <56CB>G=>3> <>730, 38?5@E@><0B>7 F8 B>?;07<K 8 =59@>3;80;L=KE :;5B>:.  -- ?@5>1;04; ;8 1>;55 2KA>:>G0AB>B=K5 :>;510=8O, >4=0:> 2>71C4! <>ABL 2 AB0488 AB>9:>9 0:B820F88 =5 87<5=O;0! 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MB8< F5=B@0< =0?@02;O5BAO 8 8=D>@<0F8O >B >A<>@5F5?B>@>2 :@C?=KE A>AC4>2, 2=CB@5==8E >@30=>2, => 3;02=K< D0:B>@>< 2 <5E0=87<5 38?>384@0B0F8>==>3> AB@5AA0 O2;O5BAO 459AB285 >A<>B8G5A:>3> 402;5=8O :@>28 =0 >A<>-@5F5?B>@K 38?>B0;0<CA0 (Verney, 1949). -B8 2>7459AB28O, >A>15==> =0 C@>2=5 AC?@0>?B8G5A:8E F5=B@>2, ?@82>4OB : 2K45;5=8N 207>?@5AA8=0, 2 @57C;LB0B5 G53> ?@>O2;O5BAO 53> 0=B848C@5B8G5A:>5 459AB285, @501A>@1F8O ?5@28G=>9 <>G8 =0@0AB05B, 48C@57 C<5=LH05BAO; ?>O2;O5BAO >;83C@8O.  A;CG05 38?>384@5<88 45A8=E@>8=70-=O @8B<0 8 2>71C645=85 AC?@0>?B8G5A:8E F5=B@>2 @0A?@>AB@0=O5BAO =0 <8=40;8=K 8 38??>:0<?  =01;N40- 37  8A. 6. !E5<0 AB@5AA>@=>3> 459AB28O 6064K (?>;848?A8O) : 1  38?>384@0B0F8O :@>28; 2  >A<>@5F5?B>@K A>AC4>2 8 >@30=>2; 3  38?>B0;0<CA: !  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C :@KA A>45@60=85< 8E 157 ?8BLO =0 ACE>< :>@<5 2 v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}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i :;5B:0E, ?>MB><C ?@>O2;ONI89AO A8=4@>< A;@0254;82< =07K20NB 38=>:A8G5A:8< 38?5@:0?=8G5A:8<. . . !5@>20 A A>02B. (1973) ?>4=8<0;8 :@KA 2 25= B8;8@C5<>9 10@>:0<5@5 =0 2KA>BC 10 000 <.  ?5@2>5 >?KB5 2K68;> 6%, ?>A;5 2B>@>3> ?>4J5<0  51'%. A 384@0B0F8O A=860;0 ?@>F5=B A<5@B=>AB8. -B8 8AA;54> 20=8O ?>4B25@640NB @01>BK . . >@87>?B>20 (1967)' Barton (1968). 2B>@K >B<5G0;8 C25;8G5=85 :>=F5=B@0 F88 35<>3;>18=0, M@8B@>F8B>2 8 A=865=85 8E >A<>B8G5 A:>9 @578AB5=B=>AB8. 8?>:AOG5A:=5 ?@>1K ==4CF8@> 20;8 AB@5AA>2K5 @50:F88 A 87<5=5=8O<8 DC=:F88 =04 ?>G5G=8:>2. Bertach et al. (1973) 2K720;8 C :@KA 38 ?>:A8G5A:8A-38?5@:0?F=G5A:85 @0AAB@>9AB20 ?@8 A>45@ 0888 8E 2 70<:=CBKE :0<5@0E A ?>AB5?5==> 87<5=ON; I8<AO 307>2K< A>AB02><. 8AB>E8<8G5A:8 =01;N40;! 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A. - 0E0@>2 (1973) ?>:070;, GB> @01>B0 >?5@0B>@>2, =5 B@51CNI0O D878G5A:8E CA8;89, A>?@>2>6405BAO C25;8G5=85< ;53>G=>9 25=B8;OF88 4> 13 ; 2 <8=CBC, @>AB>< ?>B@51;5=8O :8A;>@>40 4> 150%, A=865=85< ?0@F80;L=>3> 402;5=8O C3;5:8A;>3> 3070 2 0;L25>;O@=>< 2>74CE5 ?@8 >4=>2@5<5==>< A=865=88 ?>B@51;5=8O :8A;>@>40. !>3;0A=> . . >;:>2C (1973), ?>2KH5=85 @578AB5=B=>AB8 A0<>: 8 :0AB@8@>20==KE A0<F>2 : >AB@>9 38?>:A88 >1JOA=O5BAO 2;8O=85< MAB@>35=>2, A=860NI8E DC=:F8N I8B>284=>9 65;57K 8 0:B82878@CNI8E @5-3?:C;>M=4>B5;80;L=CN A8AB5<C.  ?@>F5AA0E 4578=B>:A8:0F88 >4=>2@5<5==> =01;N40;8 ?>2KH5=85 DC=:F88 :>@K =04?>G5G=8:>2 8 A5:@5B>@=>9 45OB5;L=>AB8 ?0=:@50A0. B. !. 8DH8F (1949), . . 2>@K:8= (1951), - . @5AB:8= (1952), . . !8@>B8=8= (1954) >B<5@;8 =0@CH5=8O ?>2545=G5A:8E 8 CA;>2=>-@5D;5:B>@=KE @50:F89 ?@8 =0@0AB0NI59 :8A;>@>4=>9 =54>AB0B>G=>-:B8. 0= 8@ 8 A>02B. (1967) =01;N40;8, GB> ?@8 38- >:A8G5A:>< AB@5AA5 C ;N459 2>7=8:0NB >B@8F0B5;L=K5 <>F88. . . 0;N6=K9 8 . . >@>O= (1973) >1=0@C68;8, ?> ?@8 38?>:A8G5A:>< AB@5AA5 @50:F88 A0<>@074@065- O 73=5B0;8AL, >1;53G0;0AL @50:F8O 871530=8O. ! ?>- )LN M;5:B@>D=78>;>38G5A:>3> <5B>40 02B>@K >1=0- 41 @C68;8 2>71C645=85 2 004=5<5480;L=KE >B45;0E 38?A B0;0<CA0, 04@5=5@38G5A:8E D>@<0F8OE 8 O4@^ ?>4:>@:8. 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(1977 1978), =5A<>B@O A>1AB25==K9 AB@5AA>@=K9 E0@0:B5@, ?>2KH05B pel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inculescu (1963), Simagi, Bernathini (1966) 8 4@. 2 =0G0;L=>< ?5@8>45 459AB28O <03=8B=>3> ?>;O >1=0@C68;8 ?>2KH5=85 0:B82=>AB8 :>@K =04?>G5G=8:>2, >4=0:> ?@8 4;8B5;L=KE 2>7459AB28OE (4> 35 ACB>:) =0ABC?0;0 0B@>D8O ?CG:>2>9 7>=K. . . !028= A A>02B. (1973) 8AA;54>20;8 459AB285 ?>AB>O==KE <03=8B=KE ?>;59 =0 15;KE :@KA0E 8 ?>4B25@48;8 40==K5 F8B8@>20==KE 2KH5 02B>@>2. 4=>2@5<5==> 2 ;8<D>84=KE >@30=0E, 2 B8<CA5 8 A5;575=:5 >=8 =01;N40;8 C<5=LH5=85 :>;8G5AB20 D>;;8:C;>2 8 ;8<D>F8B>2. A>15==> 1>;LH85 87<5=5=8O 2>7=8:0;8 ?>A;5 15-ACB>G=>3> 2>7459AB28O <03=8B=>3> ?>;O.  >?KB0E . !. 0E0=0 A A>02B. (1965) ?>4 2;8O=85< <03=8B=>3> ?>;O C A>10: >B<5G0;8 87<5=5=8O 2 18>M;5:B@8G5A:>9 0:B82=>AB8 O45@ ?5@54=59 8 704=59 G0AB59 38?>B0;0<CA0.  =5:>B>@KE A;CG0OE O2=> ?>2KH0;0AL 0:B82=>ABL 04@5=5@38G5A:8E D>@<0F89 8 I8B>284=>9 65;57K. 725AB=>, GB> 0=B83@028B0F8>==K5 @5D;5:AK 2 :>A<>A5 8AG570NB, =0@CH05BAO DC=:F8O 25AB81C;O@=>3> 0??0@0B0, ?>MB><C ?@8 =525A><>AB8 =0ABC?05B 25AB81C;O@=K9 AB@5AA. -:A?5@8<5=B0;L=> ?>4>1=K9 AB@5AA <>6=> 2K720BL C 682>B=KE @07@CH5=85< ;018@8=B>2, 22545=85< 2 ?>;>ABL ;018@8=B>2 20B:8, A<>G5==>9 10'%-=K< @0AB2>@>< =>2>:08=0. 3. !. >=F>20, . . @83>@5=:> (1973) =01;N40;8 G5@57 30 <8= E0>B8G=>ABL 2 <>=>A8=0?B8G5A:8E >B25B0E, @0AAB@>9AB2> M:AB5=7>@=KE 8 D;5:A>@=KE @5D;5:A>2, @5F8?@>:=>9 8==5@20F88. 0: 284=>, =5:>B>@K5 D878G5A:85 2>7459AB28O, B0:85 :0: <03=8B=K5 2>;=K, 2;8ONB =0 38?>30-;0<8G5A:85 F5=B@K. 6.7. !'! !" !! >A<8G5A:89 AB@5AA A>AB>8B 87 @07=KE :><?>=5=B>2.  =0G0;5, =01;N405BAO ?5@53@C7:0, 0 2 40;L=59H5< 2 7028A8<>AB8 >B 4;8B5;L=>AB8 ?>;5B0 8 B@5=8@>2:8 :>A<>=02B0  2;8O=85 A;54CNI8E D0:B>@>2: M<>F89, =525A><>AB8, 38?>48=0<88, 25AB81C;O@=KE =0@CH5=89. 081>;55 F5;5=0?@02;5==K5 8AA;54>20=8O ?> :>A<8G5A:>9 D878>;>388 >ACI5AB2;5=K . . 0@8=K< (1968), . . 075=:> (1974) 8 4@. =8 ?>:070;8 2>7<>6=>ABL ?@>3=>78@>20=8O 2 :>A<8G5A:>9 18>;>388.  43 40;L=59H5< . . 052A:89 (1978 1980) CB>G=8; ?@8* F8?K 8AA;54>20=8O 040?B0F88 >@30=87<0 : CA;>28O* 4;8B5;L=>3> :>A<8G5A:>3> ?>;5B0. < CAB0=>2;5= ?@>3 =>AB8G5A:89 0A?5:B ?>=OB89 74>@>2L5, 1>;57=L 040?B0F8O. >:070=>, GB> AB5?5=L =0?@O65=8O perjj ;OB>@=KE <5E0=87<>2  >48= 87 DC=40<5=B0;L=KE 48:0B>@>2 B5:CI53> A>AB>O=8O >@30=87<0. 2BA AG8B05B, GB> A>AB>O=85 :@>2>>1@0I5=8O O2;O5BAO ?>:073 B5;5< 040?B0F8>==>9 45OB5;L=>AB8 >@30=87<0. !>3;0A =>. . . 052A:><C, ?@>3=>78@>20=85 >@B>AB0B8G5A:>| CAB>9G82>AB8 >@30=87<0  F5==K9 M;5<5=B :>A<8G5A:A <548F8=K. >6=> ?@>3=>78@>20BL ?5@5=>A8<>ABL AB81C;O@=KE 2>7459AB289, AB5?5=L C<AB25==>3> =0?@* 65=8O 8 CB><;5=8O 2 :>A<8G5A:8E ?>;5B0E. . . 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"0:85 65 A2545=8O ?@82>4OB Yngle, Nezemi (1949), Hort (1951), . . >-3020=>20 8 . . >B02:8= (1975), . . 8BNG:>20 (1977). #25;8G5=85 :>;8G5AB20 <8B>E>=4@80;L=>3> 15;:0 2 =04?>G5G=8:0E >1=0@C68;8 Merry (1975), . . 8@C (1977). ! 4@C3>9 AB>@>=K, ?>4 2;8O=85< ' ?@>8AE>48B 18>35=57 <8B>E>=4@89 8 AB5@>84>35857 (Nus-sdorfer, 1974). >@B8:>AB5@>84K 2;8ONB =0 A8=B57 15;:>2, 8=B5=A82=>ABL 2>7459AB28O "$ =0 <8>78=, 2K7K20NB 38?5@B@>D8N <8>:0@40, 87<5=ONB DC=:F8N 8>==KE =0A>A>2. . . 8@C A A>02B. (1977) @0745;ONB <=5=85 K.rauss, 0??5 (1970), A>3;0A=> :>B>@><C I8B>284=0O 5;570 83@05B 1>;55 206=CN @>;L 2 38?5@B@>D88 <KHF <8>:0@40, G5< :>@0 =04?>G5G=8:>2. @><5 B>3>, 2 MB8E ?@>F5AA0E CG0AB2C5B 8 A><0B>B@>?8= (Goldbery, 1962). @8 A?>@B82=>< AB@5AA5, :@><5 " 8 :>@B8:>AB5- >84>2, 1>;LH>5 7=0G5=85 8<5NB 8 :0B5E>;0<8=K. "0:, "> <=5=8N . . 0AA8;O, -. (. 0B;8=>9 (1980) 8 @., :0B5E>;0<8=K CG0AB2CNB 2 ?@>F5AA0E 3;8:>35=>- 4* 51 ;870 8 3;8:>;870, . $. 5@56=>2 A A>02B. 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bi 4>2 682>B=KE =01;N40NBAO 83@0B5;L=KA @5D;5:A! :>B>@K5 A>?@>2>640NBAO ?>;>68B5;L=K<8 M<>F8O 5 8A:;NG5=>,' GB> ?@8 A?>@B82=KE C?@06=5=8OE 2;5:0NBAO 7>=K C4>2>;LAB28O 2 38?>B0;0<>-;8<18 A:8E AB@C:BC@0E (. . 3=0B>2, 1950). ?@545;5=* @>;L ?@8 MB>< 83@0NB E>;8=5@38G5A:85 8 A5@>B>HHA 38G5A:85 AB@C:BC@K 38?>B0;0<CA0, => 1>;LH>5 7=0G5=] 25@>OB=>, 8<5NB 8 8>@04@5=0;8=>2K5 =59@>=K, 0 B08 AB@C:BC@K, 2K45;ONI85 =>@04@5=0;8= (-. 5;L3>@ 1969). 6.11. !" !!  % "  !" 1I5?@8=OB>, GB> 87<5=5=85 DC=:F89 >@30=87^ ?@8 <KH5G=>9 =03@C7:5 ?@>B5:05B ?>4>1=> AB@AAA>2> @50:F88. 0:>?;5=K <=>3>G8A;5==K5 =01;N45=8O] 2;8O=88 @07;8G=KE 284>2 <KH5G=>9 =03@C7:8 =0 B@ =8@>2:C DC=:F88 >@30=87<0, >A>15==> A:5;5B=KE 8 A5 45G=>9 <KHF, >@30=>2 4KE0=8O 8 4@. G5=L F5=B 40==K5 >1 87<5=5=8OE >1<5==KE ?@>F5AA>2 2 @57C;LB B5 B@5=8@>2:8 ?>;CG5=K M:A?5@8<5=B0;L=>. 7 =0H8 ?@54K4CI8E @01>B A;54C5B, GB> 87CG5=85 AB@5AA>2L @50:F89 ?@8 <KH5G=>9 =03@C7:5 <>65B @0A:@KBL =5| B>@K5 <5E0=87<K B@5=8@>20==>AB8 (. !. 0E08| . . 0@0=>2, 1979). 5B A><=5=8O, GB> =5@2=0O A8AB5<0 83@05B 206=| HCN @>;L 2 ?@>F5AA0E B@5=8@>20==>AB8. 725AB=>, 157CA;>2=K5 8 CA;>2=K5 42830B5;L=K5, B>=8G5A:85, <! 65G:>2K5, ?@54AB0@B>2K5, AB0@B>2K5 @5D;5:AK @07=>A @07=K. -<>F8>=0;L=K5 8 2535B0B82=K5 @50:F88 20B=> @072820NBAO 8 >B =8E 7028A8B AB5?5=L B@5=8@ 20==>AB8. <5AB5 A B5< 87<5=5=8O, ?@>8AE>4OI85 >@30=87<5, =54>AB0B>G=> 87CG5=K. =0;87 M;5:B@>D873 ;>38G5A:8E ?0@0<5B@>2 ?@8 >1>@>=8B5;L=>< AB@5AA5 :070;, GB> 45A8=E@>=870F8O @8B<0 2>7=8:05B 2 :> <>730 8 @0A?@>AB@0=O5BAO =0 ;8<18G5A:CN A8AB5< ?5@53>@>4:C 8 38?>B0;0<8G5A:85 F5=B@K. 8AE>4 I85 =5@2=K5 8<?C;LAK >E20BK20NB @5B8:C;O@=CN D>$ 54 ,0F=N 8 >AB0;L=K5 >B45;K F5=B@0;L=>9 =5@2=>9 A8AB5<K 0:B82878@CNB ?5@8D5@8G5A:85 >@30=K, <KH5G=CN A8AB5<C, 4KE0=85, :@>2>>1@0I5=85.  40;L=59H5< >1-?OB=0O 0DD5@5=B0F8O, 8=D>@<0F8O >1 8=B5=A82=>AB8 ?@>F5AA>2 =0 ?5@8D5@88 ?>2KH0NB 8;8 C<5=LH0NB 0:B82=>ABL F5=B@0;L=KE @53C;OB>@>2. 1I5?@8=OB>, GB> 9" =8AE>4OI85 8 2>AE>4OI85 <5E0=87<K A>G5B0NBAO A =59@>3>@<>=0;L=K<9 8 3>@<>=0;L=K<8 D0:B>@0<8. "0:, 1>;LH>5 7=0G5=85 ?@8405BAO C@>2=N 0:B82=>AB8 =04->G5G==:>2, 2K45;ONI8E :0B5E>;0<8=K 8 :>@B8:>AB5-@>84K, :>B>@K5 2;8ONB =0 =5@2=CN A8AB5<C 8 =0 ?5@8D5@8G5A:85 >@30=K. <5AB5 A B5<, ?> 40==K< >@-?53 (1967), Berdanier (1972), . . CAA8= (1977), 04@5-=0;M:B><8@>20==K5 682>B=K5 A>E@0=ONB 2 >?@545;5==>9 <5@5 A?>A>1=>ABL ?@8A?>A01;820BLAO : ?>2B>@ONI8<AO D878G5A:8< =03@C7:0<, => 4>AB83=CB0O 040?B0F8O =5 >ACI5AB2;O5BAO ?>;=>F5==>. Tipton (1972) ?>:070;, GB> C MB8E 682>B=KE ?@8 B@5=8@>2:5 A>45@60=85 3;8:>35=0 2 <KHF0E =5 C25;8G8205BAO. . . 8@C (1976) ?>A;5 22545=8O 45:A0<5B07>=0 70 420 G0A0 4> B@5=8@>2>G=KE =03@C7>: =01;N40; =5?>;=>F5==>5 @0725@BK20=85 <5E0=87<0 >1I59 040?B0F88 8 >BACBAB285 8=4C:F88 15;:>2>3> A8=B570. 4=0:> <=>385 @01>BK ?>:070;8, GB> 45:A0<5B07>= 2;8O5B =5 B>;L:> =0 38?>B0;0<CA, => 8 =0 :>@C =04?>G5G=8:>2 8 38?>D87. >4 2;8O=85< 45:A0<5B07>=0 2 ?5@54=59 4>;5 38?>D870 A=860NBAO A8=B57 ", AB5@>84>35=57, A8=B57 B8@5>B@>?-=>3> 3>@<>=0, 2 38?>B0;0<CA5 =5 >1@07C5BAO :>@B8:>-;815@8=. !>3;0A=> !5;L5 (1950), C 38=>D87M:B><8@>20==KE 682>B=KE ?@8 2K?>;=5=88 ?>2B>@ONI8EAO D878G5A:8E =03@C7>: =0ABC?05B 1KAB@>5 8AB>I5=85. Tipton et al. (1968, 1971, 1972) ?>:070;8, GB> 38?>D87M:B><8O ?@54>B2@0I0;0 38?5@B@>D8N A5@4F0 2 ?@>F5AA5 B@5=8@>2:8, ?@>G=>ABL 65 ACE>68;89 =5 C25;8G820;0AL.  B> 65 2@5<O 22545=85 :>@B8:>B@>?8=0 8;8 A><0B>B@>?8=0 >:070;>AL =5MDD5:B82=K<. Goldberg (1967) ?@8H5; : 2K2>4C, GB> A><0B>B@>?8= =5>1E>48< 4;O 2>7@0AB=>9 38?5@B@>D88 <KHF, => =5 4;O 38?5@B@>D88 <KHF, >1CA;>2;5==>9 B@5=8@>2:>9. Golmik, Lanuzze (1975) >1=0@C68;8 ?>2KH5=85 :>;8G5AB20 AC:F8=0B45384@>35=07K 8 <8B>E>84@80;L=>3> 15;:0 2 A:5;5B=KE <KHF0E 38?>- 55 I D87M:B><8@>20==KE 8 0;;>:A0=48015B8G5A:8E :@L8 @57C;LB0B5 B@5=8@>2:8. ;O 2KOA=5=8O <5E0=87<>2 B@5=8@>2:8 =5>1E>4 BI0B5;L=K9 0=0;87 <5E0=87<>2 AB@5AA0. 7 =0H8E 4 =KE A;54C5B, GB> =01;N40NBAO 425 D07K AB@5AA0. & C65 >B<5G0;>AL 2KH5, 2 ?5@2>9, :0B01>;8G5A:>9 D 2K45;ONBAO :0B5E>;0<8=K 8 B8@5>84=K5 3>@<>=K,| 2B>@>9, 0=01>;8G5A:>9  ", :>@B8:>AB5@>84K, ; AC;8=, A><0B>B@>?8=. 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ACI5AB2CNB @07=K5 <=5=8O 8 A0< 2>?@>A =54>AB0B>G 87CG5=. "0:, 2 M2>;NF8>==>< 0A?5:B5 =5 CAB0=>2;5=A :>340 8 =0 :0:>< MB0?5 @0728B8O F5=B@0;L=>9 =5@2I A8AB5<K D>@<8@C5BAO M<>F8>=0;L=0O @50:F8O. 1I ?@8=OB>, GB> M<>F88 O2;ONBAO 18>;>38G5A:8< MDD5:B5 ?>;>68B5;L=>3> 8;8 >B@8F0B5;L=>3> 2>7459AB28O i >@30=87<. 4=0:>, E>BO B0:8< 2>7459AB28O< ?>425@6 =K 2A5 682K5 >@30=87<K, > ?>4>1=>9 @50:F88 <K 3>2 @8< ;8HL B>340, :>340 >1J5:B82=> 2 :0:>9-B> D>@\ <>6=> 55 >1=0@C68BL. 8I52K5 8 ?>;>2K5 @5D;5:A 4065 C 682>B=KE A ?@8<8B82=>9 =5@2=>9 A8AB5<>9, A B5284=>9 8;8 30=3;8>7=>-A5B5284=>9, A>?@>2>640NBA ?@87=0:0<8 C4>2;5B2>@5=8O 8;8 (1>;55 =03;O4=>) = C4>2;5B2>@5=8O. 0I8B=K5 @5D;5:AK C =0A5:><KE, <4 ;NA:>2 8 B. 4. 2K@060NB 8=>340 B@52>3C, =5C4>2;5B2] @5=85. "0:, 2A5< 8725AB=0 1C@=0O >1>@>=8B5;L=0O @50: F8O C <CE. # ?>72>=>G=KE  @K1, 75<=>2>4=KE  2KO2;ONBA ?>2545=G5A:85 @50:F88, 2K@060NI85 ?>;>68B5;L= ?8I52K5 8 ?>;>2K5 @5D;5:AK (4>1K20=85 ?8I8, 1@< =K9 =0@O4 8 B. 4.). # B0:8E 682>B=KE B0;0<8G5A:85 AB@80@=K5 D>@<0F88 =0E>4OBAO 2 70G0B>G=>< A>AB> =88, 2 38?>B0;0<CA5 >1@07>20= ?5@2K9 MB06 F5=B@> @53C;8@CNI8E DC=:F88 M=4>:@8==KE 65;57, 8 @072; 205BAO 2B>@>9 8 B@5B89 MB068 F5=B@>2, @53C;8@CNB] >1<5= 25I5AB2. # 75<=>2>4=KE ?>O2;ONBAO 3>;>A>2! @5D;5:AK  >48= 87 A?>A>1>2 2K@065=8O ?>;>2KE ? 72 0>:8B5;L=KE @C48<5=B0@=KE M<>F89. "0:85 ?@8<8B82=K5 $ + M<>F89 1>;55 =03;O4=> >1=0@C6820NBAO C 75<=>2>4=KE, @5?B8;89 8 ?B8F. !0<FK ?B8F 8740NB @07=K5 3>;>A>2K5 72C:8, ?>NB, 27J5@>H820NBAO, C =8E ?>O2;O5BAO ?>;>2>5 ?>2545=85, ?@>O2;ONI55AO 2 >?@545-45=?>9 M<>F8>=0;L=>9 @50:F88.  MB8E @50:F8OE CG0AB2CNB F5=B@K B0;0<CA0, :>B>@K5 2>A?@8=8<0NB >1I55 1;03>?>;CG85 >@30=87<0.  38?>B0;0<CA5 D>@<8@CNBAO  5=B@><5480;L=K5 8 25=B@>;0B5@0;L=K5 F5=B@K, >B :>B>@KE 7028AOB GC2AB20 =0AKI5=8O 8 3>;>40; 2 ?>4:>@:>2KE C7;0E  F5=B@K, A2O70==K5 A 2K@065=85< M<>F89, >A>15==> A >1>@>=8B5;L=K<8 @5D;5:A0<8. # @07;8G=KE 284>2 682>B=KE 2K@010BK20NBAO @07=>>1@07=K5 >@85=B8@>2>G=K5 @50:F88, A83=0;878@CNI85 >1 >?0A=>AB8 8 2K7K20NI85 C 03@5AA>@0 AB@0E. >;>68B5;L=K5 M<>F88 2K@060NBAO >A>1K<8 72C:0<8, ?>7>-B>=8G5A:8<8 8 4@C38<8 ?>2545=G5A:8<8 @50:F8O<8. @0A82> 8 =03;O4=> ?@>O2;ONBAO ?>;>68B5;L=K5 ?>2545=G5A:85 M<>F88 ?@8 ?>;>2KE 8 1@0G=KE 83@0E, ?@54H5AB2CNI8E >?;>4>B2>@5=8N C @07=KE 284>2 ?B8F. >;>68B5;L=K5 M<>F88 8<5NB 284>2>9 8 @>48B5;LA:89 .E0@0:B5@. >4>1=K5 O2;5=8O B0:65 >B<5G0NBAO 2 83@0-B5;L=KE @5D;5:A0E 2 @0==5< >=B>35=575 C ?B8F 8 <;5:>?8B0NI8E (!. . 07C<>2, 1978). # 682>B=KE 4> >1@07>20=8O ?0;5>- 8 =5>:>@B5:A0 <>6=> @07;8G0BL M<>F8>=0;L=>5 C4>2;5B2>@5=85, A2O70==>5 A DC=:F8O<8 38?>B0;0<CA0 (F5=B@K 3>;>40, 6064K, B5?;>?@>4C:F88). BACBAB285 ?8I8, 2>4K, E>;>4 ?@5645 2A53> 2;8ONB =0 38?>B0;0<CA  3;02=K5 87<5=5=8O ?@>8AE>4OB 2 04@5=5@38G5A:8E F5=B@0E, >B@8F0B5;L=K5 M<>F88 2>7=8:0NB 2 @57C;LB0B5 2K45;5=8O 04@5=0;8=0. @85< ?8I8, 2>4K, B5?;> 459AB2CNB G5@57 38?>B0;0<8G5A:85 F5=B@K, 2K7K20O GC2AB2> C4>2;5B2>@5=8O. >MB><C 2 MB>9 >1;0AB8 D>@<8@CNBAO ?5@2K5 7>=K C4>2;5B2>@5=8O, >?8A0==K5 Olds, Miller (1950) " @. E =5>1E>48<> >B;8G0BL >B F5=B@>2 2K@065=8O M<>F89, 70;>65==KE 2 ?>4:>@:>2KE C7;0E 8 2 <8=40;8=5. !;54C5B >B<5B8BL, GB> ?@8 ?5@5E>45 682>B=KE =0 A#HC >3@><=>5 18>;>38G5A:>5 7=0G5=85 ?@8>1@5;8 >1>-OB5;L=K5 @5D;5:AK (?8I52K5, >1>@>=8B5;L=K5, ?>;>A5). 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M:A?5@8<5=B0;L=K< 40==K<, CG0AB2CNB 2 <5E0=87< M<>F89.  ?5@53>@>4:5 ACI5AB2C5B @O4 F5=B@>2, >15> G820NI8E ?5@5:;NG5=85 8 2E>4 2 @07;8G=K5 7>=L8 ?>B0;0<CA0, A2O70==K5 A F5=B@0<8 C4>2>;LAB28O 1 =5C4>2>;LAB28O. '5@57 A2>4 ?5@540NBAO 8<?C;LA^ <0<8;;O@=K< 04@5=5@38G5A:8< F5=B@0< 8 ?@5645 I 3> : AB@C:BC@0<, 8<5NI8< >B=>H5=85 : >1>@>=8B? =K< @5D;5:A0<. !2>4 B0:65 >ACI5AB2;O5B A2O7L A\ 74 aWn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D8;>35=575 8 >=B>35=575 1>;52>3> >ICI5=8O =0:>?;5=K 4>AB0B>G=K5 A2545=8O. >;52>9 MDD5:B O2;O5BAO A83=0;>< >?0A=>AB8 8 >15A?5G8205B ?@8=OB85 682>B=K< ?@525=B82=KE <5@ 70I8BK. I5 =01;N45=8O Ri-chet (1902), . . @15;8 (1935) 8 4@. C:07K20;8 =0 18>;>38G5A:CN @>;L 1>;8 2 70I8B=KE @5D;5:A0E. - . AB20F0BC@>2 (1939) AG8B05B, GB> 1>O7=L 1>;8 8<55B 18>35=5B8G5A:85 :>@=8  C 682>B=KE 2>7=8:0-'B M<>F88 AB@0E0 8 2K@010BK205BAO AB@5<;5=85 C40;8BL ?@8G8=K 1>;8. 5==>= 5I5 2 1922 3. @07;8G0; D8- 85 78>;>38G5A:8A 8 A><0B8G5A:85 87<5=5=8O ?@8 1>;O >1J5:B82=> 87CG0; D878>;>38N 1>;8 8 M<>F89. . . @15;8 (1935) 8 4@. 2KO28;8, GB> 4065 KpJ :>2@5<5==>5 1>;52>5 @074@065=85 2K7K205B <=>3>-@07=K5 8 A;>6=K5 87<5=5=8O 2 >@30=87<5 (0=8<0;K^ 2535B0B82=K5, 3C<>@0;L=K5 8 3>@<>=0;L=K5). O4 02B>@>2 CAB0=>28;8, GB> 1>;52>5 @074@065 @0A?@>AB@0=O5BAO =0 @07;8G=K5 >B45;K F5=B@0;L9 =5@2=>9 A8AB5<K 8 2 ?@>F5AA 2 >A=>2=>< 2>2;5:05< A8<?0B>04@5=0;>20O A8AB5<0 (Eulcr, 1958; . . 3 A8;L, 1965, 1968, 1975; . . 5=LH8:>2, 1975; 8 4& <5NBAO <=>3>G8A;5==K5 40==K5 >1 87<5=5=8OE :>| G5AB20 :0B5E>;0<8=>2 2 18>;>38G5A:8E 684:>ABOE B5@<0;L=>9, 25=>7=>9 :@>28, A?8==><>73>2>9 684:>5? ?@8 1>;52KE @074@065=8OE. > 40==K< Kety et al. (1955), Franco (196 . . >38=>20 (1969), Rosenkranz (1969) 8 4@., 2;8O=85< @07;8G=KE M:AB@5<0;L=KE D0:B>@>2 2>71C; 45=85 F5=B@>2 3>;>2=>3> <>730 A>?@>2>6405BAO A=8> =85< A>45@60=8O :0B5E>;0<8=>2 2 53> B:0=OE. >;/ AB25==>5 87<5=5=85 :0B5E>;0<8=>2 2 >B45;0E 3>;>< 3> <>730 <>65B 1KBL @57C;LB0B>< ?>2KH5==>9 npoi F05<>AB8 35<0B>M=F5D0;8G5A:>3> 10@L5@0 (Axelrc 1962; . . 0AA8;L, 1963; 8 4@.). ". . "C@?052 (196 ?@8 1>;8 >1=0@C68; 2 F5=B@0;L=>9 =5@2=>9 A8AB5 ?@>4C:BK >:8A;5=8O :0B5E>;0<8=>2. :8A;5=85 :0| E>;0<8=>2 ?@>B5:05B ?> E8=>84=><C ?CB8, !>3;0A=> "@C,  #"+ !5/ C" '0!"' !": 4} AB28B5;L=>ABL 04@5=>@5F5?B>@>2 : 04@5=0;8=C. >;LH>5 7=0G5=85 2 1>;52KE @5D;5:A0E 8<55B @59 :C;O@=0O D>@<0F8O (. - =>E8=, 1973; 8 4@.). Tg =>F8F5?B82=K5 2;8O=8O 2K7K20NB 2>71C645=85 pel :C;O@=>9 D>@<0F88, ?> 2>AE>4OI8< ?CBO< 2>2;5:0NB!] 2KAH85 >B45;K F5=B@0;L=>9 =5@2=>9 A8AB5<K, ?> AE>4OI8<  0:B82878@CNBAO DC=:F8O <>73>2>9 4ij =04?>G5G=8:>2 8 2K45;5=85 :0B5E>;0<8=>2; ?@8 of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g . (0;O?8=0 (1972) A2O7K205B A=865=85 A>45@60=8O =>@04@5=0;==0 2 3>;>2=>< <>735 A ?@5>1;040=85< @0AE>40 3>@<>=0 =04 A8=B57><.  1>;52KE ?@>F5AA0E >A>1>5 2=8<0=85 C45;O5BAO F5=B@0< 38?>B0;0<CA0. "0:, ?> . . 0AA8;N (1975), 8<?C;LAK ?5@540NBAO 2 38?>B0;0<CA, 87 =5@2=KE :;5B>: :>B>@>3> 2KA2>1>6405BAO =>@04@5=0;8=, 0 B0:65 0:B82878@CNBAO ;8<18:>-@5B8:C;O@=K5 =>@04@5=5@38-G5A:85 AB@C:BC@K 8 A8<?0B>04@5=0;>20O A8AB5<0. -B> 2545B : 2K1@>AC 04@5=0;8=0 87 <54C;;O@=>9 7>=K, >1>30I5=8N :@>28 04@5=0;8=><, :>B>@K9 G5@57 35<0B>M=-F5D0;8G5A:89 10@L5@ ?@>=8:05B 2 F5=B@K ;8<18:>-@5-B8:C;O@=>9 D>@<0F88 8 38?>B0;0<CA A ?>A;54CNI59 0:B820F859 04@5=5@38G5A:8E M;5<5=B>2 MB8E >B45;>2. Spector et al. (1965) AG8B0NB, GB> A=865=85 A>45@60=8O =>@04@5=0;==0 2 F5=B@0;L=>9 =5@2=>9 A8AB5<5 A>?@>2>6405BAO B>@<>65=85< 42830B5;L=>9 0:B82=>AB8, A 4@C3>9 AB>@>=K, =>@04@5=0;8= 8 4>D0<8= O2;ONBAO >B25BAB25==K<8 70 ?>445@60=85 =5>1E>48<>3> C@>2=O ?A8E><>B>@=>9 0:B82=>AB8. -. (. 0B;8=0, . !. CE>20 (1976) 87CG0;8 A>45@60=85 :0B5E>;0<8=>2 2 B:0=OE C :@KA G5@57 24 G ?>A;5 22545=8O AB>;1=OG=>3> B>:A8=0 2 704=85 @>30 A?8==>3> <>730 (A?8=0;L=K9 1>;52>9 A8=4@>< ?> . @K60=>2-A:><C A A>02B., 1972). 2B>@K 2KO28;8, GB> 2 38?>B0;0<CA5 A=8605BAO :>;8G5AB2> 04@5=0;8=0, $ 8 =>@-<5B0=5D@8=0.  =04?>G5G=8:0E C<5=LH05BAO :>;8G5AB2> 04@5=0;8=0, =>@04@5=0;==0 8 $, 2 A5@45G=>9 <KHF5  A>45@60=85 =>@04@5=0;==0. -B8 40==K5 @0AA<0B@820NBAO :0: >A;01;5=85 3>@<>=0;L=>3> 8 <5480-B>@=>3> 725=L52 A8<?0B>04@5=0;>2>9 A8AB5<K. 2B>@K B0:65 >B<5G0;8 ?>A;5 M;5:B@8G5A:>3> @074@065=8O A540;8I=>3> =5@20 C<5=LH5=85 :>=F5=B@0F88 04@5=0;8=0 2 38?>B0;0<CA5, A>G5B0NI55AO A 2K1@>A>< 04@5=0;8=0 87 <>73>2>3> A;>O =04?>G5G=8:>2 2 :@>2L ?@8 >BACBAB288 87<5=5=89 A>45@60=8O 04@5=0;8=0 2 A0<>9 65;575. . . 0B@>A>2 (1980) 2K7K20; 1>;52>9 A8=4@>< =0;>65=85< =0 A540;8I=K9 =5@2 ?;0AB8=>: 87 ?;5:A8- 67 3;0A0 (<5B>4 . . @02F>20, 1956) 8 8AA;54>20; ;8G5AB2> :0B5E>;0<8=>2 2 @07;8G=KE >B45;0E 3>;>9! 3> <>730 8 =04?>G5G=8:>2 2 B5G5=85 60 4=59.  ?5 7 4=59 >?KB0 :>;8G5AB2> 8>@04@A=0;8=0 2 38?>B0;024 A5 =5A:>;L:> C25;8G820;>AL, G5@57 14 8 30 4=59 C<5=LH0;>AL. !>45@60=85 04@5=0;8=0 B0:65 ?040| 8 MB> 65 2@5<O 2 =04?>G5G=8:0E :>;8G5AB2> :0: 05B =0;8=0, B0: 8 =>@04@5=0;8=0 C25;8G820;>AL, >4=0 G5@57 60 4=59 :>;8G5AB2> 04@5=0;8=0 C<5=LH0;>AL.!1 . !. 0;048 (1978) ?@8 A402;5=88 <O3:8E mi G5@57 10 <8= =01;N40;0 C<5=LH5=85 :>;8G5AB20 0| =0;8=0 2 =04?>G5G=8:0E =0 31%, G5@57 >48= G0A 41, G5@57 5 G  =0 60%; :>;8G5AB2> =>@04@5=0;8=0 @57 10 <8= C<5=LH8;>AL =0 16%, G5@57 G0A  =0 G5@57 5G  =0 79%.  =0H8E >?KB0E <K 2K7K20;8 1>;52>9 >1>@>=8B5 =K9 @5D;5:A C :@>;8:>2. ! 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CE>20 (1976) =01;3Q 40;8, GB> G5@57 24 G ?>A;5 =0G0;0 >?KB0 C :@KA-] 1>;52K< A8=4@><>< ?> A@02=5=8N A 8=B0:B=K<8 7=0G8-^ 92 I 3 B5;L=> A=865=> A>45@60=85 =5 B>;L:> 04@5=0;8=0, =>  $ 8 =>@<5B0=5D@8=0. . . 0B@>A>2 (1980) ?@8 8@>=8G5A:>< 1>;52>< @074@065=88 >1=0@C68; C :@KA #<5=LH5=85 :>;8G5AB20 ?@>4C:B>2 >:8A;5=8O :0B5E>;0-ji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ahlstrom, 1964; Greer, 1970; Stoner, 1971).  >1571>;820=88 CG0AB2CNB M=4>@D8=K 8 M=:5D0;8=K.  7028A8<>AB8 >B 8=B5=A82=>AB8 8 4;8B5;L=>AB8 1>;52>3> AB@5AA0 =0ABC?05B 2B>@0O D070  0=01>;8G5A:0O. <5AB5 A B5<, 5A;8 :><?5=A0B>@=0O @50:F8O =5 >:065B 4>;6=>3> MDD5:B0, =0ABC?05B H>:>2>5 A>AB>O=85, 2K720==>5 8AB>I5=85< DC=:F88 :>@K =04?>G5G=8:>2, 8=>340 A> A<5@B5;L=K< 8AE>4><. 7.4. ( !!"/  !" !! !@548 ?0B>;>38G5A:8E AB@5AA>2 >A>1>5 <5AB> 70=8<05B B@02<0B8G5A:89 H>: (. . 2>7452, . . C;038=, 1970). (>:>2K5 A>AB>O=8O ?@>B5:0NB ?>4>1=> AB@5AA>2K< @50:F8O< A ?@54><8=8@CNI8<8 F8@:C;OB>@-=K<8 @0AAB@>9AB20<8. 07;8G=K5 02B>@K >?8AK20NB =5A:>;L:> 284>2 H>:0: B@02<0B8G5A:89, 35<>@@038G5A:89, 04@5=0;8=>2K9, M=4>B>:A8=>2K9 8 B. 4. (Herhey, I960; Weil, 1971).  ?0B>35=575 MB8E ?@>F5AA>2, :@><5 =0@CH5=8O :@>2>>1@0I5=8O, >B<5G05BAO 87<5=5=85 >1<5=0 25I5AB2.  >F5=:5 53> 48=0<8:8 83@0NB @>;L ?@8A?>A>18B5;L=K5 @50:F88 (. . 5B@>2,  -. C;038=, 1973).  ?0B>35=575 H>:0 2KO2;5=K <5E0=87<K, ?@>B5:0NI85 2 F5=B@0;L=>9 =5@2=>9 A8AB5<5 A ?>4:;NG5=85< " "8?>D870@=>-=04?>G5G=8:>2KE :><?>=5=B>2.  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AB5@>84>2. @8 AB@5AA>2KE @50:F8OE (D8:A0F8O :@K2 :0B5B5@870F8O 154@5==>9 0@B5@88) C@>25=L 3;N:>:2 B8:>84>2 @57:> 2>7@0AB05B 8 C45@68205BAO 2 B5G59 2A53> ?5@8>40 H>:0 2?;>BL 4> 3815;8 682>B=! (. !. 07C@:528G, . . C;8:>20, 1976). 50DD5@! B0F8O 38?>B0;0<CA0, ?> %0;0A7C 8 C?? (1965), =5 <5=O5B DC=:F88 :>@K =04?>G5G=8:>2. 1<5==K5 =0! H5=8O ?@8 H>:5 7028AOB B0:65 >B 3>@<>=0;L=KE D| B>@>2, 2KO2;5==KE 8 ?@8 4@C38E AB@5AA0E. >;K 7=0G5=85 8<55B 2K45;5=85 :0B5E>;0<8=>2 8 3>@<>? I8B>284=>9 65;57K. #AB0=>2;5=>, GB> C :>H5: 7=0| B5;L=> C25;8G8205BAO A>45@60=85 04@5=0;8=0 8 >A>H => =>@04@5=0;8=0 2 0@B5@80;L=>9 :@>28. ;>:040 -4 ;8=>@50:B82=KE AB@C:BC@ (?>A;5 22545=8O 0B@>?8=0|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llen et al, 1968; Stoner e al., 1969; 8 4@.). -B>B ?@>F5AA 4;8BAO >B 20 4> 60 < ?>A;5 =0G0;0 AB@5AA0. @8 ?5@5@57:5 A?8==>3> <>73 2K45;5=8O :>@B8:>AB5@>84>2 =5 =01;N405BAO. 2B>@ AG8B0NB, GB> =5@2=K5 8<?C;LAK 4>E>4OB 4> 38?>B0;0< A0 G5@57 2>AE>4OI85 ?CG:8 A?8==>3> <>730. 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Hillarp (1952) 8 4@. @07;8G0NB 2 <>73>2>9 G0AB8 =04?>G5G=8:>2 04@5=0;8=>2K5 8 =>@04@5=0;8=>2K5 :;5B:8. 5B>4>< $0;L:0 %8;;0@?0 G5@57 G0A ?>A;5 @50:F88 2KO2;ONBAO ?5@28G=K5 0<8=K (4>D0<8=, =>-@04@5=0;8=). !?CABO 3 G ?>A;5 >1@07>20=8O D;C>@>-D>@>2 87 ?5@28G=KE :0B5E>;0<8=>2 (4>D0<8=, =>@04@5-=0;8=) >?@545;O5BAO 2B>@8G=K9 0<8=  04@5=0;8=. 7 =0H8E 40==KE A;54C5B, GB> 04@5=0;8=>2K5 =59@>=K ?@5>1;040NB 2 O4@0E <0<8;;O@=>3> :><?;5:A0, 25?B@>;0B5@0;L=>9 G0AB8 8 2 O4@0E <8=40;8=K, <5=LH5 8E 2 O4@0E ?5@54=59 G0AB8 38?>B0;0<CA0 8 38??>:0<?5. >@04@5=0;8=>2K5 =59@>=K >B<5G0NBAO 2 >A=>2=>< 2 O4@0E ?5@54=59 G0AB8 38?>B0;0<CA0 8 2 38??>:0<?5, <5=LH5  2 <0<8;;O@=KE F5=B@0E 8 <8=40;8=5; E>;< =5@38G5A:8A =59@>=K  2 O4@0E ?5@54=59 8 25?B@><5-=0;L=>9 G0AB59 38?>B0;0<CA0 8 2 38??>:0<?5, <5=LH5 2 O4@0E 704=59 G0AB8 38?>B0;0<CA0 8 2 <8=40;8=5. !5@>B>=8=5@38G5A:85 =59@>=K ?@5>1;040NB 2 O4@0E A@54=59 G0AB8 38?>B0;0<CA0 8 2 38??>:0<?5, 2 <5=LH5< :>;8G5AB25 >=8 >1=0@C65=K 2 >AB0;L=KE D>@<0F8OE <8=40;8=K 8 38?>B0;0<CA0. 8AB>E8<8G5A:8< <5B>4>< . .. C40=F52 (1976) >1=0@C68; =>@04@5=0;8=>2K5 =59@>=K 2 ?5@825=B@8:C-9O@=KE O4@0E AB5=>: B@5BL53> 65;C4>G:0. 5A:>;L:> :;5B>: >= >B<5G0; 2 >1;0AB8 704=53> 38?>B0;0<CA0 8 2 AC?@0<0<8;;O@=>9 >1;0AB8; =51>;LHCN 3@C??C =59@>=>2 A> A;01>9 ;N<8=5AF5=F859  2=CB@8 0@:C0B?>3> O4- 103 pa, >A>15==> 2 ?5@54=5< >B45;5 8 2 25=B@0;L=>9 G0, ?5@54=53> ?5@825=B@8:C;O@=>3> O4@0. :A>=K MB>9 3>1 ?K >:0=G820NBAO 2 >1;0AB8 ?5@28G=>3> :0?8;;O@! A?;5B5=8O ?>@B0;L=>9 A8AB5<K. 5;:85 ?;>B=> @0^' ;>65==K5 =59@>=K 2KO2;ONBAO 2 4>@A><5480;L=>< j' @5 38?>B0;0<CA0. G5=L <=>3> =>@04@5=5@38G5A:8E <8=0;59 >?@545;O5BAO 2 4>@A><5480;L=><, AC?@0>2 G5A:><, ?5@825=B@8:C;O@=><, ;0B5@0;L=>< O4@0E rjj B0;0<CA0. >;LH8=AB2> O45@ 2 <0<8;;O@=KE B5;L 8<5NB >G5=L =87:CN ?;>B=>ABL MB8E B5@<8=0;59. I B@><5480;L=>< 8 ?5@54=5< 38?>B0;0<CA5 B0:65 @C65=0 =87:0O ?;>B=>ABL =>@04@5=5@38G5A:8E B5@<8| ;59, 0 2 AC?@0E807<0B8G5A:>< O4@5 >=8 >BACBAB2C9 >D0<8=5@38G5A:85 AB@C:BC@K 87CG5=K Brownste et al. (1974), . .. C40=F52K< (1976) 8 4@. 8AB>E <8G5A:8< <5B>4>< 2 <5480;L=>< O4@5 <0<8;;O@=KE -O ;5F >1=0@C68;8 1>;55 =87:CN :>=F5=B@0F8N 4>D0<8| G5< 2 ;0B5@0;L=><. >D0<8=5@38G5A:85 ?59@5 =0945=K 2 >1;0AB8 A@548==>3> 2>72KH5=8O, 2 ?0@) =>< A;>5 :>B>@>3> =01;N40NBAO ?;>B=> C?0:>20=* 4>D0<8=5@38G5A:85 B5@<8=0;8, :>B>@K5 ?>4E>4OB : ?8;;O@0< ?>@B0;L=>9 A8AB5<K (. 10. C40=F52, 19^  <5480;L=>< O4@5 <0<8;;O@=KE B5;5F >B<5G5=0 :0O :>=F5=B@0F8O 4>D0<8=0. >D0<8=5@38G5A:85 =5 @>8K B0:65 >1=0@C65=K 2 >1;0AB8 A@548==>3> 2>72L B5=8O. Rose et al. (1974) 8 4@. 38AB>E8<8G5A:8 8AA;54>20;} 0F5B8;E>;8=MAB5@07C 2 38?>B0;0<CA5 4;O 2KO2;5= E>;8=5@38G5A:8E =59@>=>2 8 >A>1>5 7=0G5=85 ?@840; <0<8;;O@=><C :><?;5:AC. 5480;L=0O 3@C??0 A0< 1>;LH0O 8 A>AB02;O5B 65,3%, ;0B5@0;L=K5 O4@0 11,6%, 4>@A0;L=K5  23,2%. =B5=A82=0O @50:F8O 0F5B8;E>;8=MAB5@07C =01;N405BAO 2 B5;0E =59@>=>2 45=4@8B0E ;0B5@0;L=>3> O4@0. 5480;L=>5 8 ;0B5@0;L =>5 O4@0 B0:65 >:@0H820NBAO 8=B5=A82=> =0 0F5B8; E>;8=MAB5@07C. KA>:0O 0:B82=>ABL 0F5B8;E>;8=MAB5@0 7K :0: 2 =59@>=0E, B0: 8 2 >:@C60NI59 B:0=8 <548 0;L=>3> <0<8;;O@=>3> O4@0 ?>72>;O5B AG8B0BL, G4 MB8 =59@>=K O2;ONBAO E>;8=>F5?B82=K<8 (. .. ) 40=F52, 1976; . . ">=:>3;0A, 1980). Aprison, Hungten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prugs et al. (1966) =01;N40;8 A>2?045=85 2 @0A?>;>65=88 2 A;>OE 38??>:0<?0 -B@0=AD5@07K 8 0F5B8;E>;8=MAB5@0-7K.  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K45;5=85 <5480B>@>2 2 A8=0?A0E >15A?5G8205B ?@>2545=85 8<?C;LA>2 A =59@>=0 =0 =59@>= B>9 5 A8AB5<K; ?@>F5AA 2KA2>1>645=8O <5480B>@0 87 B5@<8=0;59 <0;> 87CG5=. 5@>OB=>, 2 38?>B0;0<=G5A:>9 1;0AB8 MB>B ?@>F5AA >15A?5G8205B A2O7L >4=>9 A8AB5<K =59@>=>2 A 4@C3>9 8;8 2;8O=85 >?@545;5==>9 105 A8AB5<K =0 A?5F8D8G5A:85 F5=B@K, >B :>B>@KE 7028A?B ?@>F5AAK =59@>A5:@5F88 8 A8=B570 ;815@8=>2 8;8 AB0-B8=>2. > 40==K< Iversen (1974) 8 4@., ACI5AB2C5B 2;8O=85 >1@0B=>9 A2O78 2 <5E0=87<5 2KA2>1>645== <5480B>@>2 2 A8=0?A0E 8 B5@<8=0;OE. -DDA:B>@=0 :;5B:0 <>65B 2K45;OBL >?@545;5==K5 25I5AB20, @5 ;8@CNI85 :>;8G5AB2> 2KA2>1>645==>3> <5480B>@0 8; >1@0B=>3> 53> ?>3;>I5=8O 2 A8=0?A0E 8;8 2 B5@<8= ;OE (uptake).  MB>< ?@>F5AA5 83@0NB @>;L 8>=K ) +, !0++  >=8 2;8ONB =0 <5<1@0==K9 ?>B5=F8 =59@>=>2, 0 :>;8G5AB2> 04@5=0;8=0 2 38?>B0;0<CA5 I 28A8B >B A>45@60=8O (^<5B8;8@>20==KE :0BAE>;0<8 =>2 8 =5:>B>@KE AB5@>84>2 (:>@B8:>AB5@>=, MAB@048A B5AB>AB5@>=). 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(1946), . !. 0E0=0 (1947) 8 4@. ?>:070;8, GB> ?=B@025=B@8:C;O@=>5 22545=85 A>;59 :0;LF8O 2K7K205B B>@<>65=85 2 F5=B@0E 38?>B0;0<CA0, ?@82>4OI55 : A>==><C A>AB>O=8N (25@>OB=>, 1;>:8@>20=85 :0;LF89@5-45=B>@>2 2K:;NG05B 04@5?5@38G5A:=5 AB@C:BC@K), 8, =0>1>@>B, 22545=85 A>;59 :0;8O 2K7K205B 2>71C645=85. . . 0@E>=, . !. 0E0=0 (1929) =01;N40;8 C<5=LH5=85 A>45@60=8O :0;LF8O 2 B:0=OE <>730 ?>A;5 C40;5=8O >:>;>I8B>284=KE 65;57. 2545=85 ?0@0B-3>@<>-=0 405B ?@>B82>?>;>6=K9 MDD5:B. 725AB=0 @>;L :0;8O 8 =0B@8O, 2;8ONI8E =0 <5<1@0=K 5@38G5A:8E =59@>=>2, :>B>@0O =01;N405BAO 8 2 38?>B0;0<CA5. Hag-gendal (1970) 8 4@. 2KO28;8, GB> >A>1> 206=CN @>;L 83@0NB 8>=K !0++. =8 A?>A>1AB2CNB 2KE>4C =>@04@A-0;8=0 87 ?@5A8=0?B=G5A:>9 <5<1@0=K. @8B>: 8>=>2 !8++ 2 =5@2=CN B5@<880;L >AB0=02;8205B 2K45;5=85 <5480B>@0. B :>=F5=B@0F88 Mg++, Fe++, Fe+++ 7028A8B ?>3;>I5=85 <5480B>@>2 3@0=C;0<8 2 ?@5A8=0?B8- 107 G5A:>9 <5<1@0=5. =B5@5A=K5 40==K5 8<5NBAO > @>;F Mg++, Ca++, AI+++, :>B>@K5 C25;8G820;8 0:B82=>^ 45:0@1>:A8;07K (Christenson ct al., 1971) :B82=>5! :0B5E>;--<5B8;B@0=AD5@07K 7028A8B >B ?@8ACBAB2? Mg++ 2 A@545, E>BO ?++, !0++, !1_ <>3CB 53> 708 =8BL.  18>E8<8G5A:8E ?@>F5AA0E 0:B82=> CG0AB2CN^ D>AD>@, D>AD0B=K5 A>548=5=8O, @53C;8@CNI85 I5;>G=>9 @575@2 9 2E>4OI85 2 A>AB02 $, $, "$, D>A-D>;8?84>2. 0;> 87CG5=0 @>;L <8:@>M;5<5=B>2 (J, Zn, !8 4@C38E) 2 >1<5==KE ?@>F5AA0E 38?>B0;0<CA0. . . >3>4052 (1970 1980) ?>:070;, GB> 2 ochoij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a+, B>340 :0: 2 =5?'" @>=0E A>45@60=85 + A=8605BAO, a Na+ ?>2KH05BAO >;8G5AB2> +- 8 Na-"$-07K =5 87<5=O5BAO 8;8 cm 605B5O. 8?>:A8O 8 4@C385 ?@>F5AAK 2K7K20NB C ?>B><AB20 :@KA ?>2KH5==CN AC4>@>6=CN 0:B82=>ABL, =0@CH5=* M=5@35B8G5A:>3> >1<5=0 2 =59@>=0E, A2O70==K5 A> A=865=85< 2>A?@>872>4AB20 "$. @>D8;0:B8G5A:8 M<1@8>=0< <>6=> 225AB8 3;N:>7C, "$, AC:F8=0B :0;8O 3;CB0<0B 8 <5B8>=8= 4;O >15A?5G5=8O ?;0AB8G5A:>9 >1<5=0. #3;52>4=K9 >1<5= 2 38?>B0;0<CA5 ?@54AB02;O5B A0 1>9 >A>1K9 8=B5@5A, ?>A:>;L:C 2 25=B@><5480;L=2 O4@0E 8<5NBAO 3;8:>@5F5?B>@K, 2>A?@8=8<0NI85 :>;5 10=8O 3;8:5<88, 2K7K20NI85 GC2AB2> 3>;>40 8;8 =0AKI5=8O. >>1I5 2A5 =5@2=K5 :;5B:8 GC2AB28B5;L=K ?>=865=8N 3;N:>7K 2 :@>28; A 4@C3>9 AB>@>=K, 2 =: 108 <0;> 3;8:>35=0. = =0E>48BAO 2 A2O70==>< 8 A2>1>4=>< A>AB>O=88.  =0H8E >?KB0E ?>4 2;8O=85< 8=AC;8=0 :>;8G5AB2> A2>1>4=>3> 8 A2O70==>3> 3;8:>35=0 2 ?5@54=59 G0AB8 38?>B0;0<CA0 ?>2KH0;>AL 7=0G8B5;L=>, 2 704=59 G0AB8  <5=55 2K@065=85 >A;5 22545=8O 0<8=078=0 :>;8G5AB2> 3;8:>35=0 A=860;>AL 8 2 ?5@54=59 8 2 704=59 G0ABOE 38?>B0;0<CA0. =AC;8= 2>AAB0=02;820; 70?0AK 3;8:>35=0, 0<8=078=, =0>1>@>B, C<5=LH0;. @8 AB@5AA5, 2K720==>< <=>3>:@0B=K<8 >1>@>=8B5;L=K<8 @50:F8O<8, :>;8G5AB2> 3;8:>35=0 :0: 2 ?5@54=59, B0: 8 2 704=59 G0ABOE 38?>B0;0<CA0 C<5=LH05BAO. 0A?04 3;8:>35=0 ?@>8AE>48B ?>4 2;8O=85< F8:;8G5A:>3> 045=078=<>=>D>AD0B0 ($). > <=5=8N Sutherland et al. (1970), $ O2;O5BAO @5F5?B>@>< 4;O 18>;>38G5A:8E 0<8=>2 ?@8 8E D878>;>38G5A:>< 459AB288 =0 MDD5:B>@=K5 :;5B:8, -04@5=A@38G5A:85 1;>:0-B>@K <>3CB ?@5?OBAB2>20BL MB><C 2>7459AB28N. $ ?@8=8<05B CG0AB85 :0: 2 <5480B>@=KE, B0: 8 2 3>@<>=0;L=KE DC=:F8OE 18>35==KE 0<8=>2. $ 2 38?>B0;0<CA5 B0:65 0:B82878@C5B "$, :>B>@K9 AB8<C;8@C5B 2KE>4 0<8=>2 87 2578:C; ?@5A8=0?B8G5A:8E AB@C:BC@. ! 4@C3>9 AB>@>=K, D5=>B8078=K, ?C@8=K, 8>=K :0;LF8O 1;>:8@CNB A8=B57 $. 8?84=K9 >1<5= 2 F5=B@0;L=>9 =5@2=>9 A8AB5<5 8 >A>15==> 2 38?>B0;0<CA5 87CG5= =54>AB0B>G=>. "0:, ?> 40==K< Yatsu, Moss (1972), D@0:F8O <8B>E>=4@89 3>;>2=>3> <>730 :@KA A?>A>1=0 :0B0;878@>20BL C4;8=5=85 ;8=>;52>9 8 ;8=>;5=>2>9 :8A;>B 4> !22 !33 A>>B25BAB25==> 2 ?@8ACBAB288 $-, -, "$ 8 14!-0F5B8; >.  >?KB0E A @0AB2>@8<K<8 D5@<5=B0<8 @07;8G=KE >B45;>2 <>730 ?>:070=>, GB> 2 68@=K5 :8A;>BK 2:;NG0NBAO 425 !-548==FK ?@8 CG0AB88 <0;>-=8;->. Akino, Shimajo (1970) 2KO28;8, GB> 2 <>735 :C@8=>3> M<1@8>=0 ?@>8AE>48B A8=B57 D>AD0B848;-8=>78B84>2, 2 :>B>@>< CG0AB2CNB ;8?>=C:;5>B84F8B848= 8 48D>A-D0B-483;8F5@84.  <>735 FK?;OB =0945= D5@<5=B, :0B0;878@CNI89 0F8;8@>20=85 AD8=3>78=0 B8>MD8@0<8 > 4> >1@07>20=8O F5@0<840, B> 5ABL 0F8;5D8=3>78=0. . . @>E>@>20 A A>02B. (1972) 87>;8@>20;8 87 <>730 :@KA 8 D@0:F89 30=3;8>784>2. Dicessare, Dain (1971), 87CG0O 0:B82=>ABL B@0=AD5@07K, :0B0;878@CNI59 %-0F5B8;-30;0:B070<8= =0 30=3;8>784, ?>:070;8, GB> 109 1>;LH0O G0ABL 55 ;>:0;87>20;0AL 2 <8:@>A><0E (441 8 <8B>E>=4@8OE (6'%) 3>;>2=>3> <>730 :@KA, ?@8G5! 2 >=B>35=575 0:B82=>ABL 55 A=860;0AL. > 40==K< -. A5 et al. (1971), AC1:;5B>G=K5 G0AB8FK <>730 A?>A>1=K : A8=B57C 30=3;8>784>2 A 8A?>;L7>20=85< M=4>35 0:F5?B>@>2, ?@8G5< :064K9 87 =8E (<>=>-, 48- 8 B@8-A80;>30=3;8>784K) A8=B578@CNBAO =57028A8<> >B 4@C. 38E. 7CG5=85 @0A?@545;5=8O 68@=KE :8A;>B 8 >A=>20 =89 2 4;8==>9 F5?8 30=3;8>784>2 3>;>2=>3> <>730 ?A 72>=>G=KE ?>72>;8;> Avrova 8 Zallelinsky (1971 A45;0BL 2K2>4 > ?>O2;5=88 C <;5:>?8B0NI8E (2 >B;* G85 >B =87H8E ?>72>=>G=KE) A0<>AB>OB5;L=>3> A8=B57 30=3;8>784>2. >7<>6=K5 ?CB8 A8=B570 30=3;8>784>2 | 4@C38E AD883>3;8:>;8?84>2 87CG5=K Eichberg et al (1969).  ;8B5@0BC@5 8<55BAO @O4 @01>B >1 >1<5=5 D>AD>; ?84>2 2 B:0=OE =5@2=>9 A8AB5<K 2 ?@>F5AA5 <85;24 70F88. @8 701>;520=8OE =5@2=>9 A8AB5<K 87CG5=K D>AD>;8=84K <85;8=0 (. . @5?5, 1967; . . '52 25@8:>2, 1967; . . @>E>@>20 A A>02B., 1972). 0: ?>:070;8 Smith 8 Mayer (1911  1912), E>;5AB/ @8= O2;O5BAO 206=59H8< :><?>=5=B>< =5@2=>9 B:0=8, 345 >= A>AB02;O5B >:>;> 20% ;8?84>2. 081>;LH55 : ;8G5AB2> E>;5AB5@8=0 =0E>48BAO 2 A2>1>4=>< A>AB>O=88, MB> E0@0:B5@=> 4;O 27@>A;KE 682>B=KE. 0 @0==8E AB048OE M<1@8>35=570 2 <>735 G5;>25:0 8 682>B-?KE 8<55BAO MAB5@8D8F8@>20==K9 E>;5AB5@8=. Eichberg el al. (1969) =01;N40;8 2 @0==5< 2>7@0AB5 C 45B59 2 ?5@8>4 <85;8=870F88 C25;8G5=85 A>45@60=8O MD8@>2 E>;5AB5@8=0 4> 20%. =0G8B5;L=>5 :>;8G5AB2> MD8@>2 E>;5AB5@8=0 ?5@5E>48B 2 A?8==><>73>2CN 684:>ABL, AI AB02;OO 50'% >B >1I53> :>;8G5AB20 E>;5AB5@8=0 2 =59 (0,26 <3 =0 100 <;).  G8A;5 >A=>2=KE 68@=KE :8A;>B 2 MD8@0E E>;5AB5@8=0 <>730 =0E>48;8 >;58=>2CN, ?0;L<8B8=>2CN 8 ?0;L<8B8;>;58=>2CN, 0 ?>B>< ?>A;5 7025@H5=8O <85;8=870F88  0@0E84>8>2CN. Smith,Mayer (1972) ?>:070;8 =0;8G85 MAB5@=D8F8-@CNI8E 8 384@>;87CNI8E D5@<5=B>2 2 3>;>2=>< <>735 27@>A;KE 682>B=KE. !>3;0A=> 02B>@0<, E>;5AB5@>;MA8 @070 8 384@>;07K =0E>4OBAO 2 <8B>E>=4@8OE 8 <8:@>A><0E. E 0:B82=>ABL ?>2KH05BAO A 2>7@0AB><, ?0@0;;5;L=> ?@>F5AAC <85;8=870F88. -D8@K E>;5AB5@8=0 =0* :0?;820NBAO 2 =5@2=>9 A8AB5<5 4> 25% >B >1I53>  :>;8G5AB20 E>;5AB5@8=0 2 @57C;LB0B5 4535=5@0F88. Chc-valier, Lutten (1971) AG8B0NB, GB> A8=B57 E>;5AB5@8=0 ?@>8AE>48B 2> 2@5<O >1@07>20=8O 8 >1=>2;5=8O =5@2=KE :;5B>:, ?@8G5< 4;O =53> E0@0:B5@5= 70<54;5==K9 B5<?. CB8 8 A:>@>AB8 A8=B570 E>;5AB5@8=0 7028AOB =5 B>;L:> >B 2840 8 DC=:F8>=0;L=>9 0:B82=>AB8 B:0=8, => ; >B 8=4CF8@CNI53> 2;8O=8O ?0B>;>38G5A:>3> ?@>F5AA0. !>3;0A=> . !. 0E0=0 8 . - 0BC;>2>9 (1962), C :@>;8:>2 A 2>7@0AB>< (>B 8 4> 200 4=59) C25;8G8205BAO A>45@60=85 D>AD>;8?84>2 2 ?5@54=59 8 704=59 G0ABOE 38?>B0;0<CA0. # 27@>A;KE :@>;8:>2 22545=85 04@5=0;8=0 C25;8G8205B :>;8G5AB2> D>AD>;8?84>2 2 38?>B0;0<CA5; 0<8=078=0  =5A:>;L:> 53> C<5=LH05B. !>45@60=85 E>;5AB5@8=0 2 38?>B0;0<8G5A:>9 >1;0AB8 1>;LH5, G5< 2 :>@5 <>730; 2 ?5@54=59 G0AB8 38?>B0;0<CA0 1>;LH5, G5< 2 704=59. >A;5 22545=8O 0<8=078=0 A>45@60=85 E>;5AB5@8=0 2 38?>B0;0<CA5 C<5=LH05BAO, ?@8G5< 1>;55 7=0G8B5;L=> 2 53> 704=59 G0AB8. /2;5=8O AB@5AA0, 2K7K205<K5 <=>3>:@0B=K<8 >1>@>=8B5;L=K<8 @5D;5:A0<8, C<5=LH0NB :>;8G5AB2> E>;5AB5@8=0 :0: 2 ?5@54=59, B0: 8 2 704=59 G0ABOE 38?>B0;0<CA0. 54>AB0B>G=> 8AA;54>20= 2 38?>B0;0<CA5 >1<5= =C:;58=>2KE :8A;>B 8 15;:>2. . !. 0E0=0 8 . . 0-BC;>20 (1962) >1=0@C68;8, GB> A 2>7@0AB>< (>B 8 4> 200 4=59) C :@>;8:>2 :>;8G5AB2>  8  2 38?>B0;0<CA5 C<5=LH05BAO. >;55 2K@065==>5 C<5=LH5=85 =01;N405BAO C 6-<5AOG=KE :@>;8:>2 8 2 ?5@54=59 8 2 704=59 G0ABOE 38?>B0;0<CA0. > Jalima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rpynn. SH-3@C??0 2 38?>B0;0<CA5 8<55B >A>1>5 7=0G5=85, ?>A:>;L:C >=0 2E>48B 2 A>AB02 D5@<5=B=KE A8AB5<, :@><5 B>3> A>45@60B SH-3@C?8K >:B>?5?B84K 8 =5:>B>@K5 ;815@8=K 8 AB0B8=K. ! 4@C3>9 AB>@>=K, SH-3@C??K 2E>4OB 2 A>AB02 B0:8E D5@<5=B>2, :0: <>=>0<8=>>:A84070 8 045=>78;<5B8>?8=. Fri-edenwald et al. (1942) AG8B0NB, GB> <>=>0<8=>>:A84070 A>45@68B SH-3@C??K, >15A?5G820NI85 B@0=A?>@B M;5:B@>=>2. =3818@>20=85 MB>3> D5@<5=B0 ?@54>B2@0I05BAO 22545=85< 3;CB0B8>=0 8 F8AB58=0, :>B>@K5 2>AAB0=02;820NB SH-3@C??K (Lagnada et al., 1956). Axel-rod (1957), Blashko (1968), Weinshilboim (1972) 8 4@. C:07K20NB =0 @>;L SH-045=>78;<5B8>=8=0 2 :0G5AB25 4>=0B>@0 <5B8;KN9 3@C??K ", :>B>@K9 >ACI5AB2;O5B -<5B8;8@>20=85 0<8=>2 8 8E 8=0:B820F8N 2 A8=0?B8G5A:>9 I5;8 8;8 2 MDD5:B>@=>9 :;5B:5.  A2O78 A B5<, GB> @O4 B8>;>2KE @5035=B>2 (P-CL-7-<5@:C@82 15=7>0B, 8>40F5B0<84 8 4@.) O2;ONBAO 8=3818B>@0<8 112 -5:0@1>:A8;07K, 2>7<>6=> CG0AB85 SH-3@C?? 2 0:B82=>AB8 :>M=78<?8@84>:A8; D>AD0B-45:0@1>:A8;07K (Christensen, 1970).  8AA;54>20=88 DC=:F88 38?>B0;0<CA0 <K ?@84020;8 >A>1>5 7=0G5=85 >1<5=C SH-rpynn. 4;O MB>3> 8A?>;L7>20;8 ?>;O@>3@0D8G5A:89 8 38AB>E8<8G5A:89 <5B>4K (?> Barnet). 7 =0H8E 40==KE A;54C5B, GB> :>;8G5AB2> SH-rpynn <5=LH5 2 704=59 G0AB8  ' 1>;LH5 2 ?5@54=59 G0AB8 38?>B0;0<CA0. -B> <>6=> >1JOA=8BL B5<, GB> 2 ?5@54=8E O4@0E F8AB58= 8A?>;L7C5BAO 4;O A8=B570 >:B>?5?B84>2. 0 A>45@60=85 SH-rpynn 2;8ONB @07;8G=K5 3>@<>=K, A2O70==K5 A 8E >1@0B=K< 459AB285< =0 38?>B0;0<CA. "0:, . !. 0E0-=0 8 . . 0?>?>@B (1967) ?>:070;8, GB> A 2>7@0AB>< :>;8G5AB2> SH-3@C?? 2 38?>B0;0<CA5 C25;8G8205BAO. 2545=85 B8@5>84=KE 3>@<>=>2 2K7K205B C25;8G5=85 :>;8G5AB20 SH-3@C?? C <>;>4KE :@>;8:>2, 0 C 27@>A;KE (3 4-<5AOG=KE)  C<5=LH5=85. 8AB>E8<8G5A:8< <5B>4>< Barnet 2KO2;5=>, GB> 2 F8B>?;07<5 =59@>=>2 AC?@0>?B8G5A:8E 8 ?0@025=B@8:C;O@=KE O45@ <5=LH5 SH-3@C??, 2 =59@>A5:@5B5, 2 <56:;5B>G=KE ?@>AB@0=AB20E  1>;LH5, GB> <>6=> >1JOA=8BL CA8;5==>9 0:B82=>ABLN =5@2=KE :;5B>: C <>;>4KE :@>;8:>2. 0>1>@>B, C 27@>A;KE 682>B=KE SH-3@C?? 1>;LH5 2 F8B>?;07<5 =59@>=>2, <5=LH5 2 <56:;5B>G=>< ?@>AB@0=AB25, GB> C:07K205B =0 A=865=85 8=B5=A82=>AB8 =59@>A5:@5-:H8 (. "0;8:8=0, 1970). 2545=85 D>;;8:C;8=0 8 ?@>35AB5@>=0 42CE;5B=8< :@>;8:0< C<5=LH05B :>;8G5AB2> SH-3@C?? 2 =5@2=KE :;5B:0E C:070==KE O45@ ?> B>9 65 ?@8G8=5. # 27@>A;KE :@>;8:>2 ?>4 2;8O=85< ?>;>2KE 3>@<>=>2 1>;LH5 =0:0?;8205BAO SH-3@C?? 2 F8B>?;07<5. 5 8A:;NG5=>, GB> B0:85 87<5=5=8O 8<5NB >B=>H5=85 : A8=B57C >:B>?5?B84>2 8 B8@5>;815@8=0. . !. 0E0=0, . !. C48<0 (1973) ?>:070;8, GB> C :@KA 18;0B5@0;L=0O 04@5=0;M:B><8O A=8605B :>=F5=B@0F8N SH-3@C?? 2 ?5@54=59 G0AB8 8 =5A:>;L:> ?>2KH05B 2 704=59 G0AB8 38?>B0;0<CA0. >A;5 22545=8O 384@>:>@B87>=0 2>AAB0=02;8205BAO ?@56=89 C@>25=L SH-B@C??. 0: A>;52>5, B0: 8 1572>4=>5 ?8B0=85 8 >A>15==> 22545=85 38?5@B>=8G5A:>3> @0AB2>@0 E;>@8AB>3> =0B@8O 2K7K20NB A=865=85 :>;8G5AB20 SH-3@C?? 2 ?5@54=5< 8 704=5< >B45;0E 38?>B0;0<CA0. 725AB=>, GB> 3?>384@0B0F8O :@>28 CA8;8205B 2K45;5=8O 207>?@5A- 8 0:07 ! 457.  A8=0, 0=B848C@5B8G5A:>5 459AB285 :>B>@>3> C<5=LI05 2K45;5=85 2>4K 87 >@30=87<0. >MB><C CA8;5=85 A8=. 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Martini (1954) AG8B0;, GB> 0F5-B8;E>;8= AB8<C;8@C5B A8=B57 207>?@5AA8=0, :>B>@K9 459AB2C5B =0 2K45;5=85 ". >A;5 22545=8O 04@5=0;8=0 :@KA0< 2 38?>B0;0<8G5A:>9 >1;0AB8 >B<5G05BAO C25;8G5=85 :>;8G5AB20 SH-3@C??. 5 8A:;NG5=>, GB> B0:85 87<5=5=8O A2O70=K A 0:B820F859 <>=>0<8=>>:A8-407K 8 SH-045=>78;<5B8>=8=0. 2545=85 =>@04@5=0;8=0 :@KA0< C<5=LH05B A>45@60=85 SH-3@C?? 2 38?>B0;0<CA5. "8@5>84=K5 3>@<>=K 8<5NB B0:>9 65 MDD5:B. >4 2;8O=85< 0<8=078=0 :>;8G5AB2> SH-3@C?? 2 38?>B0;0<CA5 B0:65 C<5=LH05BAO, GB> <>6=> >1JOA=8BL A=865=85< 2K45;5=8O 04@5=0;8=0. 2545=85 A5@>B>=8=0 :@KA0< ?@82>48B : =0@0AB0=8N :>;8G5AB20 SH-3@C??. =B5@5A=> >B<5B8BL, GB> 04@5=0;8= 8 A5@>B>88= 2K7K20NB ?@>B82>?>;>6=K5 MDD5:BK ?> A@02=5=8N A =>@04@5=0;8=>< 8 04@5=0;8=><. 5B>4K >?@545;5=8O SH-3@C?? 2 38?>B0;0<CA5 ?> ACI5AB2C @02=>F5==K <5B>40< >?@545;5=8O :>;8G5AB20 F8AB58=0. 5>1E>48<> ?>4G5@:=CBL, GB> F8AB58=  F8AB8= 2E>4OB 2 A>AB02 >:B>?5?B84>2 8 3>@<>=>2 38?>- 114 .40<CA0. "0:8< >1@07><, ?> :>;8G5AB2C SH-rpynn |0} AC48BL >1 0:B82=>AB8 =5:>B>@KE D5@<5=B>2, A>45@60I8E MB8 @048:0;K. ! 4@C3>9 AB>@>=K, ?> :>;8G5AB2C SH-rpynn <>6=> AC48BL >1 C@>2=5 >1@07>20=8O S A.3@C??, >1 8=B5=A82=>AB8 A8=B570 >:B>?5?B84>2 8 =5-:0B>@KE 3>@<>=>2 38?>B0;0<CA0 (. $. '5@=>:0=, 1980). Klegg and Klegg (1960) ?@82>4OB C1548B5;L=K5 >?KBK, ?>:07K20NI85, GB> <5G5=K9 F8AB58=, 22545==K9 O 38?>B0;0<CA, G5@57 =5:>B>@K9 8=B5@20; 2@5<5=8 2KO2;O5BAO 2 704=59 4>;5 38?>D870 2 A>AB025 >:B>?5?B84>2. Safran, Sayers (1960) CB25@640NB, GB> >:B>?5?B84K 2;8ONB =0 A8=B57 :>@B8:>;815@8=0, E8<8G5A:0O AB@C:BC@0 :>B>@>3> 1;87:0 : 207>?@5AA8=C. 5@>OB=>, 8=B5=A82=>ABL A8=B570 >:B>?5?B84>2 2 38?>B0;0<CA5 8<55B 8 4@C3>5 7=0G5=85. "0:, 4> >B:@KB8O :>@B8:>;815@8=0 8 4@C38E 3>@<>=>2, ?> 40==K< Sawyer (1960) 8 4@.", AG8B0;>AL, GB> 207>?@5AA8= O2;O5BAO AB8<C;OB>@>< A8=B570 .". ABL >A=>20=8O 4C<0BL, GB> >:B>?5?B84K A;C60B <0B5@80;><, 87 :>B>@>3> A8=B578@CNBAO =5:>B>@K5 ;8-15@8=K 8 AB0B8=K 38?>B0;0<CA0. 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elye, 1949). ! 4@C3>9 AB>@>=K, 0448A>=>-20 1>;57=L 8;8 ?>4>1=K5 O2;5=8O =5 ?>@>640NB A:;>==>AB8 : CA8;5=8N 2>A?0;8B5;L=>9 @50:F88 (Rose, 1953; Hylmeyer, 1953). >A?0;8B5;L=K9 035=B 8 >G03 2>A?0;5=8O 2;8ONB =0 :>@C =04?>G5G=8:>2 G5@57 CA8;5==>5 2K45;5=85 ", :>B>@K9 2 A2>N >G5@54L @53C;8@C5B C@>25=L 3;N:>:>@-B8:>84>2 2 :@>28. >A;5 38?>D87M:B><88 @578AB5=B=>ABL 682>B=>3> : 459AB28N 2>A?0;8B5;L=>3> 035=B0 125 70<5B=> A=8605BAO (Tonuffi, 1949 1954). 2B>@ =014N. 40;, GB> AB@5AA>@ 2 A>AB>O=88 AB8<C;8@>20BL DC=:F8N :>@K =04?>G5G=8:>2 8 2K7K20BL 8=2>;NF8N B8<8:>-;8<D0B8G5A:>3> 0??0@0B0. 2545=85 " AB8<C;8@C9 :>@C =04?>G5G=8:>2 8 CA8;8205B ;8<D>F8B>;87 C 38?>-D87M:B><8@>20==KE 682>B=KE.  7028A8<>AB8 >B AB5?5=8 >E;0645=8O ?>A;54>20B5;L=> 87<5=O5BAO ?@>4C:F8O :>@B8:>848KE 3>@<>=>2 2>A?0;8B5;L=0O @50:F8O <>48D8F8@C5BAO (Pepper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obson et al., 1950 1952; Kramer, 1953). >4 2;8O=85< <8=5@0;>:>@B8:>84>2 CA8;8205BAO M:AAC40F8O (Rindau, 1953). #:070==K5 3>@<>=K ?>2KH0NB ?@>=8F05<>ABL :0?8;;O@>2. "0:, 22545=85 457>:A8:>@-B8:>AB5@>=0 ?@8 2>A?0;5=88 @57:> CA8;8205B >B5: 8. G0AB>BC =03=>5=8O (. . 5<CA, 1958). >4 2;8O=85< 457>:A8:>@B8:>AB5@>=0 ?>2KH05BAO 8=B5=A82=>ABL 2>A?0;8B5;L=>9 @50:F88 2 F5;>< (. . >@87>=B>2, 19571). 2545=85 0;L4>AB5@>=0 2K7K205B C :@KA A 01AF5AA0<8 2K@065==CN ?@>;8D5@0F8N 8 @0728B85 :0?8;;O@=>9 A5B8 (Rjomani, 1955). >;LH8=AB2> M:A?5@8<5=B0;L=KE =01;N45=89 ?>:070;8, GB> @07;8G=K5 :>@B8:>AB5@>84K 2 7028A8<>AB8 >B 8E A2>9AB2 <>3CB >:07K20BL ?>402;ONI55 459AB285 =0 ?@>F5AA 2>A?0;5=8O. KO2;5=>, GB> C65 ?@8 <5AB=>< ?@8<5=5=88 457>:A8:>@B8:>AB5@>= ?>402;O5B 2>A?0;5=85, 2K720==>5 8=>@>4=K< B5;>< 8;8 :@>B>=>2K< <0A;><. <5AB5 A B5< ?@8 2=CB@825==>< 22545=88 >= CA8;8205B 2>A?0;8B5;L=CN @50:F8N. # 04@5=0;M:B><8@>- I .4==KE 682>B=KE 457>:A8:>@B8:>AB5@>= CA8;8205B >B5: Meyer et al., 1952). > :;8=8G5A:8< 40==K<, 457>:A8:>@B8:>AB5@>= 2K-" ji,isacT >1>AB@5=85 ACAB02=>3> @52<0B87<0, => =5 459AB-C5B =0 E@>=8G5A:CN D>@<C 53>. ;L4>AB5@>= B>;L:> 2 ' 1>;LH8E 4>70E CA8;8205B 2>A?0;8B5;L=CN @50:F8N. O4 40==KE C:07K205B =0 >B@8F0B5;L=>5 459AB285 A><0B>--@>?=>3> 3>@<>=0 (Taubenhaus, 1954). @>B82>2>A?0;8B5;L=K5 2>7459AB28O >:07K20NB 3;N-:>:>@B8:>84K ?@8 22545=88 BC15@:C;8=0 8 4@C38E 0;- 1 ;5@35=>2 (Bradley, 1950).  >?KB0E A 2>A?0;5=85<, 7K720==K< ?@8 ?><>I8 2>74CH=>3> <5H:0, MB8 3>@<>- I 8K 8=3818@CNB ?@>=8F05<>ABL A>AC4>2, M:AAC40F8N 8 I @0728B85 >B5:>2, C3=5B0NB @>AB D81@>F8B>2, >1@07>20=85 >A=>2=>9 AC1AB0=F88 8 2>;>:>=, CA8;820NB ;878A ;8<D>- 8 ?;07<>F8B>2, 87<5=OO B0:8< >1@07>< @0728- (| B85 >A=>2=KE D07 2>A?0;5=8O (. . !B@>30=>20, 1936; I . . 54=8:, 1953; . . !<8@=>2, 1955; Cooper et al., I 1957; E. P. 03@0<O=, 1959).  A2O78 A MB8< 3;N:>:>@-B=:>84K 1K;8 8A?>;L7>20=K ?@8 ;5G5=88 2>A?0;5=8O ACAB02>2, @>3>2>9 >1>;>G:8 3;070, ;8<D0B8G5A:8E C7- j ;>2. >@B87>= B0:65 ?>402;O5B 2>A?0;8B5;L=CN @50:F8N I C 682>B=KE (Taubenhaus, 1953). !2O7L 3>@<>=>2 :>@K =04?>G5G=8:>2 A 2>A?0;8B5;L=K< ?@>F5AA>< 4>:07K20- I 5BAO 8E CA8;5==K< 2K45;5=85< C :@>;8:>2 ?>4 2;8O=8- 1 5< 0A5?B8G5A:>3> 2>A?0;5=8O (. . .4052, .. . !C@8:>2, 1957). >@B87>= 7045@68205B @0728B85 A0<KE @0==8E ?@87=0:>2 2>A?0;5=8O: 87<5=5=85 ?@>=8F05<>AB8 A>AC4>2, 70AB>9 :@>28, M<83@0F8O ;59:>F8B>2, ?>O2;5=85 2>A?0;8B5;L=>3> >B5:0. @8 2K7K20=88 01A- | 8AAA>2 C 682>B=KE, ?>;CG0NI8E ?>4:>6=> A:8?840@ 2 I A<5A8 A :>@B87>=><, <>6=> =01;N40BL @0728B85 G@57<5@=> B>=:>3> A;>O 3@0=C;OF8>==>9 B:0=8, A=865=85 >1@07>20=8O :0?8;;O@>2, C3=5B5=85 @>AB0 D81@>F8B>2, C<5=LH5=85 8E @07<5@>2 (Sahaeffer, 1951). 84@>:>@B87>= >1;0405B 2 80 @07 1>;55 A8;L=K< ?@>B82>2>A?0;8B5;L=K< 459AB285<, G5< :>@B87>=. @8 2:;NG5=88 D;C>-@0 2 <>;5:C;C :>@B8:>AB5@>84>2, =0?@8<5@ 45:A0<5B0-7>=0, 8E 0=B82>A?0;8B5;L=>5 459AB285 CA8;8205BAO.  <5E0=87<5 2>A?0;8B5;L=>9 @50:F88 1>;LH>5 7=0G5=85 8<55B =0@CH5=85 @53C;OF88 A>AC48AB>9 ?@>=8F05<>AB8. 725AB=>, GB> " 8 :>@B87>= C<5=LH0NB ?@>- 127 =8F05<>ABL A>AC4>2 8 <5<1@0= :;5B>:. > 40==K< 24. :>B>@KE 02B>@>2, ?@>=8F05<>ABL <5<1@0= 7028A8B 8 >B 380;C@>=8407K, :>B>@0O ?>402;O5BAO " 8 :>@B87>=>< (Seifter, 1950). !>3;0A=> Menkins (1950), :>@B8:>ABA@>=4K C3=5B0NB 459AB285 ;59:>B>:A8=0 8 4@C38E 0:B82=KE ?>;8?5?-B84>2 8 15;:>2. " >1;0405B A2>9AB2>< =5?>A@54AB25==> 459AB2>20BL =0 M:AAC48= 8 C 04@5=0;M:B><8@>20=-=KE 682>B=KE ?>402;O5B 53> 459AB285. <5NBAO 40==K5 >1 8=3818@CNI5< 2;8O=88 " 8 :>@B87>=0 =0 459AB285 M:AAC48=0 8 D81@8=>;878=0 (Ungar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|B> 2>7@0AB0=85 ?>B@51;5=8O :8A;>@>40 ?>4 2;8O=85<  4@5=0;8=0 A2O70=> A 53> 459AB285< =0 ?5G5=L. >A;5 I' /2545=8O 682>B=K< 04@5=0;8=0 >B<5G05BAO 42CD07?K9 ?@8@>AB ?>B@51;5=8O :8A;>@>40.  B5G5=85 ?5@2>9 D07K |1 F> C25;8G820;>AL =0 20%, 2B>@>9  =0 .'%. @8 8=- B@0F8AB5@=0;L=>< 22545=88 1>;LH8E 4>7 04@5=0;8=0 87<5=5=8O B5<?5@0BC@K B5;0 =0?><8=0;8 ;8E>@04:C ! /Skoric, 1962), 0 4@C385 =01;N40;8 38?>B5@<8N I (- $5;L415@3, 1971). ">;5@0=B=K5 : ?8@>35=C :@>;8:8 40NB =>@<0;L=CN I B5<?5@0BC@=CN @50:F8N =0 04@5=0;8=. 4@5=0;8=>20O :5\:?5@0BC@=0O @50:F8O =5 C3=5B05BAO :>@B87>=>< 8 4", B>340 :0: 2>7=8:=>25=85 ;8E>@04:8 :0: A;54AB-I 2=5 10:B5@80;L=KE ?8@>35=>2 7=0G8B5;L=> 70<54;O5BAO MB8<8 ?@5?0@0B0<8. #25;8G5=85 2 :@>28 04@5=0;8=0 I =01;N405BAO C ;8E>@04>G=KE 1>;L=KE (?=52<>=8O, BC15@:C;57), 0 B0:65 C :@>;8:>2 ?@8 8=D5:F88 (Atkins, 1960). > 40==K< Feldman et al. (1941), ;8E>@04:0, 2K40==0O 22545=85< B8D>?0@0B8D>7=>9 20:F8=K, ?@82>48B : 0:B820F88 :0: A8<?0B8:>-04@5=0;>2>9, B0: 8 703>-8=AC;8=>2>9 A8AB5<K, >4=0:> A> 7=0G8B5;L=K< ?@5>1;040=85< 2 >1KG=KE CA;>28OE B>=CA0 A8<?0B>-04@5-?0;>2>9 A8AB5<K. # 682>B=KE 65, ;8H5==KE =04?>G5G=8:>2 A >4=>2@5<5==>9 203>B><859 (2 F5;OE 45=5@20F88 8=AC;O@=>-3> 0??0@0B0), :0:8E-;81> ACI5AB25==KE A4283>2 B5<?5@0BC@K B5;0 ?>A;5 22545=8O 20:F8= =5 =01;N40;8. @8 45=5@20F88 =04?>G5G=8:>2 (?5@5@57:8 G@52=KE =5@-I 7>2) =8:0:8E 87<5=5=89 A5:@5F88 04@5=0;8=0 2 >B25B I =0 22545=85 ?8@>35=0 =5 >B<5G0;8, 2KA>B0 65 8 ?@>4>;68B5;L=>ABL B5<?5@0BC@=>9 @50:F88 ?@8 MB>< =5 87<5=8;8AL.  >?KB0E =0 A>10:0E A :0=N;8@>20==>9 =04?>-G5G=8:>2>9 25=>9 ?>:070=>, GB> ?@8 22545=88 <0;KE .10.'! ;8?>?>;8A0E0@840 ;8E>@04:0 A>?@>2>6405BAO CA8;5==>9 A5:@5F859 :0B5E>;0<==>2. @8 1>;LH8E 4>78@>2:0E A5:@5F8O 04@5=0;8=0 CA8;820;0AL. 5@5@57:0 % A?8==>3> <>730 =0 C@>2=5 !7 70 420 4=O 4> 22545=8O ?8@>35=0 ?>;=>ABLN CAB@0=O;0 @50:F8N <>73>2>3> 25I5AB20 =04?>G5G=8:>2 =0 459AB285 ?8@>35=0, => =5 2;8O;0 =0 @0728B85 B5<?5@0BC@=>9 @50:F88. >A;5 22545=8O 2 :@>2O=>5 @CA;> A>10: ;8?>?>;8A0E0@840 :>;8-:H;0 >B<5G5=> C25;8G5=85 :>=F5=B@0F88 04@5=0;8=0 '%> 0:07 ! 457. 129 J 130 2 :@>28 =04?>G5G=8:>2>9 25=K 2 35 @07, => A5:@5FF =>@04@5=0;8=0 =5 87<5=O;0AL. 5=5@20F8O =04?>G5GF8' :>2 CAB@0=O;0 C:070==K5 87<5=5=8O (Mikiel et al., l96h -=4>B>:A8=K, ;8?>?>;8A0E0@84K 2K7K20NB ;59:>?5=8N 8 ;8<D>?5=8N, 8<5NI8E >?@545;5==>5 AE>4AB2> A 040?B0F8>==K< A8=4@><><. 0;L=59H85 8AA;54>20-=8O CAB0=>28;8 7=0G8B5;L=CN 0:B820F8N A8AB5<K 38. ?>B0;0<CA 38?>D87 :>@0 =04?>G5G=8:>2 ?@8 459AB288 ?8@>35=>2 (Atkins, I960; 8 4@.). 4=0:> ;8E>@04>G=0O @50:F8O 2 >B25B =0 22545=85 ?8@>35=0 @0728205BAO 8 C 38?>D87M:B><8@>20==KE :>H5: (Whale, 1950). 5>1E>48<> B0:65 CG8BK20BL, GB> ?@5?0@0BK " 8 :>@B87>=0 C 682>B=KE 8 ;N459 =5 ?>2KH0NB, 0 ?>=860NB B5<?5@0BC@C B5;0. -B> 459AB285 " ?@>O2;O5BAO C :@>;8:>2 C65 ?@8 4>78@>2:0E 0,1 54./:3. @8 MB>< B5<?5@0BC@0 B5;0 A=8605BAO =0 1 2 2 B5G5=85 B@5E G0A>2. 57:> >A;01;O5BAO 8 ;8E>@04>G=0O @50:F8O, 2K7K205<0O 10:B5@80;L=K<8 ?=@>-35=0<8 (Douglas et al., 1952). =B8?8@AB8G5A:85 8 38?>B5@<878@CNI85 MDD5:BK " 8 :>@B87>=0 C 682>B=KE 8 ;8E>@04OI8E 1>;L=KE 1K;8 ?>:070=K <=>38<8 02B>@0<8.  CA;>28OE 8=D5:F8>==>9 ;8E>@04:8 MB> 459AB285 A2O7K20NB A B>@<>65=85< 2>A?0;5=8O 8 2K@01>B:>9 M=4>35==>3> ?8@>35=0 (Recant et al., 1950). <5NBAO 40==K5 > =0;8G88 ?@O<>3> C3=5B0NI53> 459AB28O :>@B87>=0 =0 F5=B@K B5@<>@53C;OF88 (. . KF:89, 1961).  ?>A;54=55 2@5<O A8=B5B8G5A:8 ?>;CG5=K <5B01>;8BK (MB8>E>;0=, 384@>:A8?@53=0= 8 4@C385), ?@8 22545=88 :>B>@KE C G5;>25:0 ?@>O2;O5BAO ;8E>@04:0 (Palmer, 1961). 2545=85 M=4>B>:A8=0 :>;810F8;0 2K7K205B 2K45;5=85 " G5@57 2 3 G 8 MB> 459AB285 >ACI5AB2;O5BAO G5@57 1070;L=>-<5480;L=CN G0ABL 38?>B0;0<CA0, ?>A:>;L:C 2K45;5=85 " =01;N405BAO 8 ?>A;5 450D-D5@5=B0F88 38?>B0;0<CA0. -=4>B>:A8=, 25@>OB=>, 2;8O5B 8 =0 38?>D87 (Makara et al., 1971). -DD5:B M=4>B>:A8=0 =0 2K45;5=85 " ?>A;5 38?>D87M:B><88 =5 =01;N405BAO. @8 MB>< M=4>B>:A8= 2K45;5=8O :>@B8:>AB5-@>84>2 =5 AB8<C;8@C5B (Wexler, 1957; Melby, I960). 5-0DD5@5=B088O 38?>B0;0<CA0, =0>1>@>B, =5 ?@5?OBAB2C5B 2K45;5=8N ". ! 4@C3>9 AB>@>=K, 2;8O=85 M=4>B>:A8=0 =0 " =5 A>2?0405B A ?8@>35==K< MDD5:B><. I >@065=85 <5480;L=>3> 38?>B0;0<CA0, 1;>:040 MB>9 D>@<0F88 E;>@?@><078=><, <>@D8=><, ?5=B010@18B0-1(,< ?@5?OBAB2CNB C25;8G5=8N :>@B8:>AB5@>=0 2 ?;07<5. 5:A0<5B07>= (20 <:3) 1;>:8@C5B 459AB285 25 <:3 -^4>B>:A8=0 =0 2K45;5=85 ". =:C10F8O ?5@54=59 qacTH 38?>D870 A M=4>B>:A8=>< ?>402;O5B A8=B57 &". "0:8< >1@07><, ?@8 ;8E>@04:5 =01;N405BAO 2K45;5=85 :0B5E>;0<8=>2 8 :>@B8:>AB5@>84>2, ?@>B5:0NI55 ?>A;54>20B5;L=> 2 2845 D07. 10.3. #""  !" !! ">B D0:B, GB> A?5F8D8G5A:85 8<<C==K5 @50:F88 A>?@>2>640NBAO =5A?5F8D8G5A:8<8 3C<>@0;L=>-3>@<>-=0;L=K< ?@>F5AA0<8, 4>:070= 402=>. "0:, @07;8G=K5 D>@<K 8<<C=8B5B0, ?@8>1@5B5==K5 ?>A;5 1>;57=8 8;8 ?>A;5 20:F8=0F88, B5A=> A2O70=K A> A2>9AB20<8 0=B=-I 35=0. 0AA82=K9 8<<C=8B5B 7028A8B >B >A>15==>AB59 22545==KE 0=B8B5;, >4=0:> ?@8 MB>< <>3CB 2>7=8:0BL " N1>G=K5 @50:F88, GB> 7028A8B >B GC65@>4=KE 15;:>2. 'B> :0A05BAO D0:B>@>2 8<<C=8B5B0, =0 ?5@2>< ?;0=5 @0AA<0B@820NB A2>9AB20 <8:@>1>2, 8E 15;:>2>-=C:;5-8=>2K5 :><?>=5=BK, 70B5<  E8<8G5A:85 >A>15==>AB8 B>:A8=>2. 1@0I05BAO 2=8<0=85 =0 ?CB8 ?@>=8:=>25=8O 2 >@30=87<, <0:@>D038G5A:CN A8AB5<C, =0 :;5B:8 <575=E8<0;L=>3> ?@>8AE>645=8O, @5B8:C;>M=4>B5;80;L-=CN A8AB5<C ?5G5=8, A5;575=:8, :>AB=>3> <>730, A>548=8B5;L=>9 B:0=8, ;8<D0B8G5A:8E 65;57 8 =0 ?>4286=K5 M;5<5=BK, ;59:>F8BK :@>28 8 ;8<DC. 725AB=>, G3> @5F5?B>@K :>68, A;878ABKE >1>;>G5:, 0=38>@5F5?B>@K CG0AB2CNB 2 @5D;5:B>@=KE 8<<C=>-18>;>38G5A:8E ?@>F5AA0E. !G8B0NB, GB> 8<?C;LAK 4>AB830NB F5=B@0;L=>9 =5@2=>9 A8AB5<K, 2 G0AB=>AB8 38?>B0;0<CA0, 0 70B5< 84CB : 8A?>;=8B5;L=K< >@30=0<, : <0:@>D038G5A:>9 A8AB5<5. 5AB0 ?5@5:;NG5=8O 0DD5@5=B=KE 8<?C;LA>2 ?>A;5 22545=8O 0=B835=0 @0A?>;>65=K =0 @07;8G=KE MB060E =5@2=>9 A8AB5<K, => 1>;LH55 7=0G5=85 8<55B 5480;L=0O, G0ABL 38?>B0;0<CA0 (Atkins et al., 1966; .. M. 5?8= A A>02B., 1969). 2B>@K ?@54?>;030;8, GB> 0DD5@5=B=K9 ?CBL 2K@01>B:8 0=B8B5; >ACI5AB2;O5BAO G5@57 ?0@0A8<?0B8G5A:CN 8 A8<?0B8G5A:CN =5@2=CN A8AB5<C. K;> ?>:070=>, GB> 65;57K 2=CB@5==59 A5:@5- -* 13! F88 (I8B>284=0O 65;570, =04?>G5G=8:8, 38?>D87 8 4@ 385) ?@8=8<0NB CG0AB85 2 @53C;OF88 8<<C=>-18>;>3-F. G5A:8E ?@>F5AA>2. 07;8G=K5 D0:B>@K  >E;0645=^ ?5@53@520=85, CB><;5=85, 028B0<8=>7, =0@CH5=85 ?8B0=8O, >B@02;5=85, =0@:>7 8 4@. 87<5=ONB <5E0=87< 8<<C=8B5B0. ;02=K<8 D0:B>@0<8, >B :>B>@KE 7028AOB 8<<C==K5 ?@>F5AAK, O2;ONBAO 3C<>@0;L=K5 :;5B>G=K5 25I5AB20 8 ;8<D>F8BK. >;LH>5 2=8<0=85 C45;O5BAO ;8<D>F8B0< B8<CA0 ("-B8<>F8BK) 8 ;8<D>F8B0< :>AB=>3> <>730 (-;8.G. D>F8BK). > 40==K< <=>38E 02B>@>2, 2 2K@01>B:5 0=B8B5; CG0AB2CNB :;5B:8 ;8<D>84=>9 A8AB5<K B8<CA0 ("-;8<D>F8BK) 8 :;5B:8 :@>25B2>@=KE >@30=>2 (-;8<-D>F8B+'). >;L B8<CA0 2 8<<C=8B5B5 1K;0 4>:070=0 2 >?KB0E Sherman (1963). "0:, C =>2>@>645==KE <L? ?>A;5 C40;5=8O B8<CA0 @072820;0AL @>:>20O 1>;5 Wasting disease, A=860;0AL 0:B82=>ABL 8<<C== ?@>F5AA>2, ?>O2;O;0AL 30<<0-3;>1C;8=5<8O, <KH8 3810;8 >B 8=B5@:C@@5=B=KE 701>;520=89. Nepssal et al. (1968) 8 4@. ?>:070;8, GB> 8<<C=>@ 0:B82=>ABL B8<>M:B><8@>20==KE <KH59 ;53:> 2>A< =02;8205BAO ?@8 22545=88 ?>4>?KB=K< 682>B=K< <>F8B>2 8;8 ;8<D>F8B>2 3@C4=>3> ?@>B>:0, => =5 :>A =>3> <>730. <<C==K9 >B25B =0 ?@>1C A M@8B@>F8B0* 10@0=0 C ?>4>1=KE B=<>M:B><8@>20==KE <KH59 ?> cf 59 25;8G8=5 @02=O5BAO >B25BC =>@<0;L=KE. 4=0:>, 40==K< Conture (1970), 'Wagner (1972) >1@07>20=85 0=B8B5; 7028A8B >B :>>?5@0F88 <564C |  -:;5B:0<8. 07;8G0NB 420 B8?0 "-:;5B>:: "-:;5B:* ?><>I=8:8 (>=8 :>>?5@8@CNBAO A -;8<D>F8B0<8 ?@>4C:F88 3C<>@0;L=KE 0=B8B5;) 8 "-;8<D>F8BK :F ;5@K (:;5B:8 MDD5:B>@>2 g @50:F8OE 8<<C=8B5B0). <0;>5 7=0G5=85 8<5NB =57@5;K5 :;5B:8 B8<CA0, >G5=L GC2AB28B5;L=K : :>@B8:>AB5@>840< ( + + ), 2 2@5<O :0: ", ( + ). "2 (+)  <5=55, 0 :;5B:8  2>5 I5 =5 GC2AB28B5;L=K. <5AB5 A B5< A;54C5B CG5ABL, ) ?@>F5AA 8<<C=8B5B0 =0G8=05BAO A <><5=B0, :>340 <0:@>D03 D03>F8B8@C5B 0=B835= (:;5B:0 ). @54?>;0305BAO, GB> >B <0:@>D03>2 : :;5B:0< " 8 , ?@>4CF8@CNI8E 0=B8B5;0, ?5@5=>AOBAO 3>B>2K5 <0B@8FK 4;O A8=B570 A?5F8D8G5A:>3> -3;>1C;8=0. 5>1E>48<> >B<5B8BL 8 B>, GB> ?>4 2;8O=85< 0=B835=>2 =0@0AB05B 2K45;5=85 ;59:>?>MB8=>2, 2 @57C;LB0B5 2>7=8:05B ;59:>F8B>7, 0:- 132 ^2878@C5BAO <0:@>D038G5A:0O A8AB5<0.  ?>445@60=88 ]0<A>AB070 ;59:>F8B>70 ?@840NB 7=0G5=85 AB2>;>2K< I5B:0< :>AB=>3> <>730 (35<>?>MB=G5A:85 AB2>;>2K5 :;5B:8 F8@:C;8@CNB 2 ?5@8D5@8G5A:>9 :@>28) (Lewis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arat de Sousa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ohen (1972) 8 4@. 0 B><, GB> 2 @57C;LB0B5 :>>?5@8@>20==>3> <56:;5B>G=>3> 2708<>459AB28O "-;8<D>F8BK (:;5B:8-?><>I=8:8) 2>2;5:0NB 2 8<<C==K9 >B25B -:;5B:8 :>AB=>3> <>730.  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"0:, ?>A;5 42CAB>@>==53> ?>2@5645=8O A@54=53> >B45;0 38?>B0;0<CA0 2 >1;0AB8 25=B@>- 8 4>@A><5480;L=KE O45@ =01;N405BAO @57:>5 C3=5B5=85 >1@07>20=8O 0=B8B5; C <>@A:8E A28=>: 2 >B25B =0 22545=85 ;>H048=>9 AK2>@>B:8 (Fillip, 1966, 1973). >;=>5 C3=5B5=85 8<<C=8B5B0 >B<5G0;>AL 8 ?>A;5 ?>2@5645=8O 704=8E 38?>B0;0<8G5A:8E O45@ (Fillip, 1966, 1973; . . >@=52 A A>02B., 1967, 1970). -;5:B@>AB8<C;OF8O @07;8G=KE O45@ 38?>B0;0<CA0 40;0 @07=>>1@07=K5 @57C;LB0BK, AB8<C;OF8O <0<8;;O@=KE B5;  C25;8G5=85 033;NB8=8=>2 2 :@>28 (. . 5B@>2A:89, 1961). . . >@=52, - %08B (1962) =01;N40;8, GB> ?>A;5 @074@065=8O 704=8E 38?>B0;0<8G5A:8E O45@ 8<<C==K9 >B25B CA8;8205BAO. . . !5@>20 A A>02B. (1973) =5 >B<5G0;8 87<5=5=8O 8<<C=>35=570 ?>A;5 M;5:B@>;8B8G5A:>3> ?>2@5645=8O 704=8E 38?>B0;0<8G5A:8E O45@. >6=> ?>;030BL, GB> 2 ?@>F5AA5 AB0=>2;5=8O 8<<C=8B5B0 2>7=8:0NB >?@545;5==K5 2708<>>B=>H5=8O <564C A?5F8D8G5A:>9 8 =5A?5F8D8G5A:>9 @50:F8O<8. =8 1>;55 O@:> 2KO2;ONBAO 2 8<<C==KE @50:F8OE ?@8 8=D5:F88. 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'5@=O:>2, J954), ?>@>3>2^ 4>70 04@5=0;8=0 =5 ?>2KH05B :@>2O=>3> 402;5=8O (. . 8A5;52, 1954).  B> 65 2@5<O =01;N40NBAO ?>B>=8O, 70<54;5==0O A:>@>ABL :@>2>B>:0, C<5=LH5=;)! 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upulescu, Sahleanu (1961) 8 4@. =07K20NB 8E M:7>0;;5@3859. "0:, ?>A;5 22545=8O 8=AC;8=0 >B<5G0NBAO <5AB=K5 @50:F88, :>B>@K5 8=>340 ?@>B5:0NB A 35=5@0;87>20==K<8 ?@87=0:0<8 (:@0?82=8F0, >B5: 28=:5, 1@>=E80;L=0O 0AB<0, 0=0D8;0:B8G5A:89 H>:). =AC;8=>-@578AB>=B=>ABL B0:65 2E>48B 2 MBC 3@C??C O2;5=89 (Tuft, Wiliams, I960). 01;N40NBAO ?>4>1=K5 @50:F88 8 ?@8 459AB288 3>@<>=>2 ?5@54=59 4>;8 38?>D870, 3>@<>=>2 @>AB0, 3>- 137 =04>B@>?8=0, ". "0:, Charpin, Lafiropulos (196IB = 4@. G0I5 2A53> =01;N40;8 ?>;>68B5;L=K5 8=B@045 <8G5A:8A @50:F88: :@0?82=8FC, >B5: 28=:5, 0=0D8;0:B8G5A:85 @0AAB@>9AB20. 565 B0:85 @50:F88 2AB@5G0NBAO ?>A;5 22545=8O >:B>?5?B84>2, MAB@>35=>2, ?@>35AB5@>=0, 0=4@>35=>2, AB5@>84=KE 3>@<>=>2. 5:>B>@K5 02B>@K =01;N40;8 0;;5@38G5A:85 @50:F88 2 @57C;LB0B5 2K45;5=8O A>1AB25==KE 3>@<>=>2 8 =07K20;8 8E M=4>0;;5@3859. "0:85 O2;5=8O >B<5G5=K 2 ?5@8>4 ?@54<5=AB@C0;L=>3> F8:;0.  20 53% A;CG052 2 MB>< ?5@8>45 C 65=I8= =01;N40NBAO ?>;>68B5;L=K5 :>6=K5 @50:F88 =0 ?>;>2K5 3>@<>=K (@50:F8O @0-C=8F, NAB=5), 8=>340 2>7=8:0NB ?@8ABC?K 1@>=E80;L=>9 0AB<K. Turiat et al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A;CG05 0;;5@388 8 >A>15==> ?@8 0=0D8;0:B8G5A:>< H>:5 <>6=> 3>2>@8BL, ?> 2K@065=8N !5;L5, > =5045:20B=>9 D>@<5 AB@5AA0 :0: 1>;57=8 040?B0F88. @54AB02;ONB 8=B5@5A @01>BK Till, McCullogh (196I, 1970), Burch, Gold, Cline (1974) 8 4@., A>3;0A=> :>B>,. @K< 8=D5:F88, B>:A8=K, B@02<K, >1;CG5=85 ?@82>4OB : >1@07>20=8N :>;>=8AB8<C;8@CNI53> D0:B>@0 (Colony stimulating factor, CSF), 2K7K20NI53> @07<=>65=85 AB2>;>2>9 :;5B:8 :>AB=>3> <>730, GB> 2 A2>N >G5@54L AB8<C;8@C5B M@8B@>?>M7 8 ;8<D>?>M7. $0:B>@ CSF ?@54AB02;O5B A>1>9 25I5AB2>, 1;87:>5 : M@8B@>?>MB8?C 8 ;59:>?>MB8=C, A2O70==>5 A 3;8:>?@>B5840<8, A>45@60I8<8 A80;>2CN :8A;>BC. > 40==K< Burch (1974), 87>;8@>20= ;8?>?@>B584=K9 D0:B>@, 1;>:8@CNI89 459AB285 CSF. >6=> ?>;030BL, GB> ?@8 0;;5@388 <>3CB >1@07>2K20BLAO 1;>:8@CNI85 D0:B>@K 8<<C=8B5B0. 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MB8E CA;>28OE A8=B57 8=AC;8=0 2 15B0-:;53-:0E >AB@>2:>2>3> 0??0@0B0 C<5=LH05BAO, 8 :>;8G5AB2> ?>ABC?0NI53> 2 :@>2L 8=AC;8=0 A=8605BAO, ?>A:>;L:C 87<5=ONBAO ?CA:>2K5 203CA=K5 F5=B@0;L=K5 <5E0=87<K, >ACI5AB2;ONI85 AB8<C;OF8N A8=B570 8 2K45;5=8O 8=AC;8=0 87 -:;5B>: >AB@>2:>2>3> 0??0@0B0 2 :@>2L.  @0728B88 A0E0@=>3> 48015B0 8<5NB 7=0G5=85 >15 D07K AB@5AA>@=>3> 2;8O=8O =0 >@30=87<.  ?5@2>9 D075 AB@5AA0 (:0B01>;8G5A:>9) =0 >@30=87< =51;03>?@8OB=> 2;8ONB :0B5E>;0<8=K (04@5=0;8=, =>@04@5=0;8=), :>B>@K5 C25;8G820NB C@>25=L A0E0@0 2 :@>28, 0 B0:65 8<5NB 0=B88=AC;O@=>5 459AB285. 725AB=>, GB> 0:B82870F8O DC=:F88 I8B>284=>9 65;57K B0:65 A?>A>1AB2C5B @0728B8N A0E0@=>3> 48015B0, :@><5 B>3>, >?@545;5==CN @>;L 83@0NB 8 4@C385 3>@<>=K  3;N:03>=, 207>-?@5AA8= 8 4@. > 2B>@>9 D075 (0=01>;8G5A:>9) A0E0@=K9 48015B CAC3C1;O5BAO ?@8 2K45;5=88 :>@B8:>AB5@>84>2, :>B>@K5 5I5 1>;LH5 =0@CH0NB C3;52>4=K9 >1<5=.  MB>9 D075 CG0AB2CNB 8 4@C385 3>@<>=K (A><0B>B@>?=K9 8 ?@>AB0-3;0=48=K), >A>15==> 8725AB=> :>=B@08=AC;O@=>5 2;8O=85 A><0B>;815@8=0. 11.4. !" !!+ ) ", !!"+ >;>4  B8?8G=K9 ?8I520@8B5;L=K9 AB@5AA. Huls-man (1978) AG8B05B, GB> ?@85< 1>;LH>3> :>;8G5AB20 ?8I8, >A>15==> 68@0, B0:65 AB8<C;8@C5B AB5@>84>35-=57. #A8;5=85 8;8 >A;01;5=85 A5:@5B>@=>9 8;8 <>B>@- 149 =>9 DC=:F88 ?8I520@8B5;L=>9 A8AB5<K >A>15==> ?>4 2;8O=85< D0@<0:>;>38G5A:8E 25I5AB2 (22545=85 0B@>?8=0, 38AB0<8=0, 8=AC;8=0, 30AB@8=0, A5:@5B8=0, >:A8-B>F8=0 8 4@.) <>6=> >F5=8BL :0: AB@5AA>2K5 @50:F88. 0 A5:@5B>@=K5 8 <>B>@=K5 DC=:F88 65;C4>G=>-:8H5G=>3> B@0:B0 >:07K20NB 2;8O=85 8=4C:F8>==K5 0=01>;8G5A:85 3>@<>=K (. . (;K38=, 1979). !B@5AAK 2K7K20NB >@30=8G5A:85 87<5=5=8O 2 ?8I520@8B5;L=>< B@0:B5 C ;N459, 8 MB> 1K;> ?>4B25@645=> M:A?5@8<5=B0;L=>. 0?@8<5@, C A>10: =52@>35==K9 AB@5AA (M:A?5@8<5=B0;L=K9 =52@>7) E0@0:B5@87C5BAO @07=>>1@07=K<8 =0@CH5=8O<8 DC=:F89 @07;8G=KE >B45;>2 ?8I520@8B5;L=>3> B@0:B0, =5@54:> DC=:F8>=0;L=K5 ?@87=0:8 A>?@>2>640NBAO AB@C:BC@=K<8 87<5=5=8O<8 2 65;C4:5, :8H5G=8:5, ?>465;C4>G=>9 65;575 (. ". C@F8=, 1952; 8 4@.). #AB0=>2;5=>, GB> ?>A;5 8<<>18;870F88, >A>15==> C :@KA, @0728205BAO B8?8G=0O O720 65;C4:0 (. . 0@:5;L, . . >;?0:>2, 1973; . . 0@8=, 1973; 8 4@.). <<>18;870F8O C<5=LH05B 0:B82=>ABL 045=>78=B@8D>AD0B07K ("$-07K) 2 3>;>2=>< <>73C, 2 @57C;LB0B5 G53> =0@CH05BAO @53C;OB>@=>5 2;8O=85 F5=B@0;L=>9 =5@2=>9 A8AB5<K =0 DC=:F88 65;C4:0. B>@8G=> 2 65;C4:5 =0ABC?05B 70AB>9 :@>28, >BB>@305BAO A;878AB0O >1>;>G:0, =0@0AB05B :>;8G5AB2> :>@B8-:>AB5@>=0 (. . 0@8=, 1973). #AB0=>2;5=>, GB> 22545=85 :>@B8:>AB5@>84>2 682>B=K< 2K7K205B O72C 65;C4:0 8 425=04F0B8?5@AB=>9 :8H:8. 5<0;>5 7=0G5=85 8<55B >BACBAB285 ?@85<0 ?8I8 2 MB>9 D075. # G5;>25:0 ?>2B>@=K5 8 4;8B5;L=K5 AB@5AAK, ?@82>4OI85 : ?>2KH5=8N 2K45;5=8O :>@B8:>AB5@>84>2, 2K7K20NB 38?5@A5:@5F8N 65;C4>G=>3> A>:0, 38?5@E;>@384@8N 8 2 40;L=59H5<  O725==CN 1>;57=L. @><5 B>3>, C B5E ;8F, :>B>@K5 C65 AB@040NB 38?5@0F84=K< 30AB@8B><, ?>O2;O5BAO O725==0O 1>;57=L  AB@5AA>2K5 @50:F88 <>3CB CEC4H8BL B5G5=85 O725==>9 1>;57=8 8 A?>A>1AB2>20BL 2>7=8:=>25=8N ?@54@0:>2KE 87<5=5=89. #EC4H05BAO 8 B5G5=85 4@C38E 1>;57=59, B0:8E :0: A?0AB8G5A:85 :>;8BK, 48A:8=5788 65;G=>3> ?C7K@O; C 1>;L=KE, AB@040NI8E ?0=:@50B8B><, @0728205BAO A0E0@=K9 48015B. !. . @0C45 A A>02B. (1973) 2KO28;8, GB> ?@8 >6>3>-;CG52>9 B@02<5 C A>10:, :@><5 =0@CH5=8O DC=:F88 8=B0:B=>3>, 45=5@28@>20==>3>, 87>;8@>20==>3> 65;C4:0, ?>2KH05BAO 0:B82=>ABL :>@:>2>3> 8 <>73>2>3> 150 A,N52 =04?>G5G=8:>2, 2K45;ONBAO "  8 A><0B>B@>?-FK9 3>@<>=. @C385 284K AB@5AA0 >B@060NBAO =0 ?@>F5AA5 2A0AK20=8O 2 B>=:>< :8H5G=8:5 ( . . $09B5;L-15@3, 1970; ". . 0<0;89, 1973), =0 DC=:F8OE ?5G5=8 8 ?>465;C4>G=>9 65;57K (. $. >;?0:>20, 1955). >A;5 AB@5AA>2 ?@54AB02;ONB 8=B5@5A F8B>A85:B@>-D>B><5B@8G5A:85 8AA;54>20=8O =C:;58=>2KE :8A;>B ?5G5=8 (. . C@=8= A A>02B., 1967, 1973): >1=0@C65=0 ?5@20O D070 (1 7 4=59)  A=865=85 8 2B>@0O D070 (7 60 4=59)  C25;8G5=85 :>=F5=B@0F88 =C:;58=>2KE :8A;>B 2 F8B>?;07<5 8 2 O4@5 35?0B>F8B>2. ;87:85 : AB@5AAC 87<5=5=8O 2K7K20NB ?8I52K5 8 0;;5@38G5A:85 @50:F88. =B5@5A=> >B<5B8BL, GB> ?8I520O 0;;5@38O A>?@>2>6405BAO >?@545;5==K<8 87<5=5=8O<8 DC=:F88 :8H5G=8:0. 0?@8<5@, C 45B59 ?@8 :8H5G=>9 0;;5@388 87<5=O5BAO :>;8G5AB2> ?;07<0B8G5A:8E :;5B>: 2 B>=:>< :8H5G=8:5 8 ?@>F5AA 2A0AK20=8O (Linarez, Lopez et al., 1977). ! ?><>ILN 18>?A88 ?>:070=>, GB> C 1>;L=KE :>;8G5AB2> ?;07<0B8G5A:8E :;5B>: ?>2KH5=>. @><5 B>3>, C =8E >1@07C5BAO 8 2K45;O5BAO 1>;LH5 8<<C=>3;>18=0. ;;5@38G5A:85 @50:F88 C 45B59 2K7K20NBAO ?>A;5 ?>ABC?;5=8O ?8I8, A>45@60I59 15B0-;0:B>3;>1C;8= :>@>2L53> <>;>:0, 0;L1C<8=K 8 3;>1C;8=K O8G=>3> 15;:0 8 65;B:0, 15;:8 8 4@C385 25I5AB20 =5:>B>@KE O3>4 8 D@C:B>2 (:;C1=8:0, <0;8=0, ?5@A8:8). @8 0;;5@38G5A:>9 @50:F88 87<5=O5BAO ;59:>F8B0@=K9 8=45:A 840;O 030=0, A=8605BAO G8A;> ;59:>F8B>2 2 :@>28. 01;N405BAO B@><1>?5=8G5A:89 MDD5:B. ;;5@38G5A:85 @50:F88 65;C4>G=>-:8H5G=>3> B@0:B0 A>?@>2>640NBAO AB@5AA>2K<8 @50:F8O<8, =0@CH0NI8<8 38?>B0;0<>-?H>D870@=>-04@5=0;>2K5 2708<>>B=>H5=8O. > 40==K< Strandbcrg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B5>@5B8G5A:>9 B>G:8 7@5=8O MB>B 2>?@>A <>65B 1KBL @5H5= A CG5B>< 8=48284C0;L=KE >A>15==>AB59 :064>9 ;8G=>AB8. !8;L=K9 B8? 2KAH59 =5@2=>9 45OB5;L=>AB8 AG8B05BAO =081>;55 A?>A>1=K< : 040?B0F88 <2 AB@5AA>2KE CA;>28OE. 4=8< 87 >A=>2=KE D0:B>@>2, ?>2KH0NI8E @578AB5=B=>ABL 8 040?B0F8>==K9 ?>B5=F80; >@30=87<0 2 A8BC0F8OE AB@5AA0, O2;O5BAO A?>@B82=0O B@5=8@>20==>ABL.  >1I8E G5@B0E <>6=> 3>2>@8BL > ;5G5=88 AB@5AA>2KE ?>A;54AB289, ?>A:>;L:C >=8 ?@>B5:0NB :0: >?@545;5==K9 ?0B>;>38G5A:89 ?@>F5AA. >;LH>5 7=0G5=85 8<5NB =>2K5 40==K5 > @5F5?B>@0E 3>@<>=>2 8 =59@>=>2 38?>B0;0<CA0, > @>;8 =58@>?5?B=4>2, M=4>@D8=>2 8 ;59:8=>2 2 @53C;OF88 DC=:F88 M=4>:@8==>9 8 =59@>25-35B0B82=>9 A8AB5<.  70:;NG5=85 A;54C5B >B<5B8BL, GB> 2 @57C;LB0B5 <=>3>G8A;5==KE A>2@5<5==KE 8AA;54>20=89 1K;8 >B:@KBK =>2K5 >1;0AB8 ?@>O2;5=8O AB@5AA0, =5:>B>@K5 AB>@>=K 53> <5E0=87<0, =0<5G5=K >A=>2=K5 ?CB8 ?@5>4>;5=8O 2@54=KE ?>A;54AB289 AB@5AA0, 2KO2;5=K G5BK@5 53> D07K: 1) :0B01>;8G5A:0O, 2) ?5@5E>4=0O (1;>:040 04@5=5@38G5A:8E @5F5?B>@>2), 3) 0=01>;8G5A:0O, '1) MDD5:B>@=0O (2>AAB0=>2;5=85 3><5>AB070, 040?B0F8O, B@5=8@>20==>ABL 8;8 ?0B>;>38O). F* 163 14. #/ " "#  14CA0<0B>20 . .   :=.: !B@5AA 8 040?B0F8O. 8H8=52, )I. ==F0, 1978. 10H8;>20 . .   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 1978. 10H8;>20 . ., 5:A;5@ /. . 8 4@.   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 1978. 30460=O= . ., =4@>=8: /(. ., ><5=40=B>2 . ., 07A>-;>2 . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. 30D>=>20 . ., >=><0@520 ". $., ;59=8: . !.   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 1978. 4> . ., !<8@=>2 . .  N;;. M:A?5@. 18>;. 8 <54., 1913, ! 16, A. 1 2. 68?0 /. .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (BHHF0, 19730. 68?0 /. ., 0;L:>2 . ., 3>@>20 . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 19736. ;5H8= . ., 0<8=0 . ., #A . .   :=.: >2>5 > 3>@<>=0E < <5E0=87<5 8E 459AB28O. 852, 0C:>20 4C<:0, 1977. =>E8= . . @8=F8?80;L=K5 >A=>2K >1I59 B5@0?88 DC=:F8>=0;L=KE A8AB5<. ., 548F8=0, 1971. @H02A:89 . .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 1980. E<5B>2 . 3., 1A0<0B>2 .   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 1978. 052A:89 . .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 19800. 052A:89 . .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 19806. 0@0=>2 . ., 0E0=0 . !. 59@>3>@<>=0;L=K5 <5E0=87<K B@5=8@>20==>AB8. 8H8=52, (B88=F0, 1979. 5=5B0B> ., ?@8H8C - 8 4@.  @>1;5<K M=4>:@8=>;>388 8 3>@<>=>B5@0?88, 1959, B. 5, ! 2, A. 43 51. 5@B0H . ., @C:8=0 . ., !5@35520 . !.   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. >3><>;5F . .  548:>-18>;>38G5A:89 6C@=0;, 1926, B. 1, A. 15-34. >=40@5=:> . ., !?>48= . -, 5I5B=N: . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0! 1973. 164 Spayde !. ., 0@3C=8=0 . ., ;870@L52 . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0. 1973. &C40=F52 . .. >=>0<8=5@38GAA:85 A8AB5<K <>730. ., 548F8=0, 1976. ,K:>2 . . ?KB 87CG5=8O @53C;OF88 D878>;>38G5A:8E DC=:F88. .-., 74.  !!! , 1949. K:>2 . ., C@F8= . ". >@B8:>-28AF5@0;L=0O ?0B>;>38O. ., 548F8=0, 1960. 0A8;L52 . II.   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B8?=F0, 1973. 0A8;L:>2 . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B8?=F0, 1973. 0AN:>20 . ., !>A:8= . !.   :=.: >2>5 > 3>@<>=0E 8 <5E0=87<5 8E 459AB28O. 852, 0C:>20 4C<:0, 1977. 54O52 $. ., /:>2F>20 . $. 8 4@.   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B8?=F0, 1978. 8@C . . 53C;OF8O D5@<5=B=KE A8AB5< 3>@<>=0<8 I8B>284=>9 65;57K 8 =04?>G5G=8:>2. "0;;8=, 1974, A. 41 43. 8@C . .  "5>@8O 8 ?@0:B8:0 D878G5A:>9 :C;LBC@K, 19770, !9, A. 28 31. 8@C . .  -=4>:@8==K5 <5E0=87<K @53C;OF88 ?@8A?>A>1;5=8O >@30=87<0 : <KH5G=>9 45OB5;L=>AB8.  "0@BC, 19776, B. 7, A. 11 20. 8@C . .  #A?5E8 D878>;>38G5A:8E ?0C:, 1980, 4, A. 27. >;:>2 . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B?HH0, 1973. 0<0;89 ". ., @0GC: . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B8?=F0, 1973. 0@:028 . X.. 20:8=0 . ., #:>;>20 . .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B8?=F0, 1980. 5@H5=>28G 3. ., %5@C28<>20 . .   :=.: $C=:F8>=0;L=0O M2>;NF8O =5@2=>9 A8AB5<K. . ., 74.  !!! , 1965. >@485=:> . .   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B=8=-F0, 1978- >@87>=B>2 . .   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B8?=F0, 1978. >@87>=B>2 . .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B8?=F0, 1980. >B;>15@ . ., 0A8;L520 .. .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B8?=F0, 1973. CA:0 . ., >AB>;0:5 . ., 565=0@C . !, >@>30O . .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 1980. 02845=:> . .. >76CE8= . !.   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973, A. 331  332. 02K4>2 . .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 1980. >=F>20 3. !, @83>@5=:> . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 19730. >=F>20 3. !, @83>@5=:> . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 19736. >@>30= . .  0B5@80;K ?> <5E0=87<0< 2708<>459AB28O H8- 165 B>284=>9 65;57K A 3=?>B0;0<8G5A:>9 >1;0ABLN.   := " 8?>B0;0<>-M=4>:@HHK5 2708<>>B=>H5=8O. 8H8=52, 196M" B. 106, A. 70 90. 0E0@>2 . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K 8H8=52, (B88=F0, 1973. K1:>25F . /-   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. ;L8= . ., >7=5A5=A:89 . !.   :=.: !B@5AA 8 53> ?0B>35=5-B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. 0;8=8=0 . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. 0;N6=K9 . ., >@>O= . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. 0AA8;L . .   :=.: @>1;5<0 38AB>35<MB8G5A:8E 10@L5@>2.  19650. 0AA8;L . .   :=.: $878>;>38O 8 ?0B>;>38O M=4>:@8==>9 A8AB5<K. 0B5@80;K I AJ5740 M=4>:@8=>;>3>2 #!! . %0@L:>2 19656. 0AA8;L . ., 0B;8=0 -. (.   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 1978, A. 230. 0AA8;L -.   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 1980. 0E0=0 . !. >@B8:>-28AF5@0;KHO @53C;OF8O DC=:F89 I8B>284=>9 65;57K. 8H8=52, 0@BO >;4>25=OA:0, 1960. 0E0=0 . !.   :=.: III :>=D5@5=F8O ?> 2>?@>A0< M;5:B@>D878>;>388. 852, 1960. 0E0=0 . !, 0=47N: -. .  &8B8@>20=> ?> 0E0=0 . !.  8?>B0;0<=G5A:85 A8=4@><K. 8H8=52, 0@BO >;4>25=OA:0, 1961. 0E0=0 . !.   :=.: 0B>D878>;>38O 38?>B0;0<CA0. 8H8=52, 0@BO >;4>25=OA:M, 1961. 0E0=0 . !.   :=.: 0B>D878>;>38O 38?>B0;0<CA0. 8H8=52, 0@BO >;4>25=OA:M, 1961. 0E0=0 . !, 0BC;>20 .   &8B8@>20=> ?> 0E0=0 . !.  8?>B0;0<=G5A:85 A8=4@><K. 8H8=52, 0@BO >;4>25=OA:M, 1962. 0E0=0 . !, !5@3552 . ., !8<0H:52HA .-.   :=.: VII :>=D5@5=F8O => ?@>1;5<0< D878>;>388. CB08A8, 1965. 0E0=0 . !. 8?>B0;0<=G5A:85 A8=4@><K. 8H8=52, 0@BO >;4>25=OA:M, 1965. 0E0=0 . !, 0?>?>@B . .  #G5=K5 70?8A:8 8H8=52A:>3> C=825@A8B5B0, 1967, B. 87, ! 9. 0E0=0 . !, 8:0 . .   :=.: @>1;5<K D878>;>388 38?>B0;0<CA0. 852, 1968, ! 2. 0E0=0 . !.  #G5=K5 70?8A:8, 1969, B. 106, A. 3 30. 0E0=0 . !, 0?>?>@B . .   :=.: A5A>N7=K9 18>E8<8G5A:89 AJ574. "0H:5=B, 1969. 0E0=0 . !.   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K-8H8=52, (B88=F0, 1973. 0E0=0 . !, 5;L=8: . ., >1C . .   :=.: :BC0;L=K5 ?@>1;5<K AB@5AA0. 8H8=52, (B88=F0, 1978. 0E0=0 . !, C= "EF 3>: 0=   :=.: 0CG=0O :>=D5@5=F8O ?@>D5AA>@A:>-?@5?>4020B5;LA:>3> A>AB020 8H8=52A:>3> C=825@A8B5B0, 1978. 0E0=0 . !, 5;L=8: . . >;L 38?>B0;0<CA0 2 @53C;OF88 3>-<5>AB070 M=4>:@8==KE 8 >1<5==KE ?@>F5AA>2. 8H8=52, (B88=F0, 1978. 0E0=0 . !.   :=.: 8AB>35<0B=G5A:85 10@L5@K. ., 1978. 0E0=0 . !, 8=L.  ;8O=85 ?>;>2KE 3>@<>=>2 ?0 18>M;5:B@8G5A:CN 0:B82=>ABL ;8<18:>-38=>B0;0<8G55:?E AB@C:BC@.  0CG=0O A5AA8O 3>AC==25@A8B5B0, 8H8=52, 1978. 0E0=0 . !, C;L15@5 . .  &8B8@>20=> ?> 0E0=0 . !. >;L 38?>B0;0<CA0 2 3><5'>AB075 M=4>:@8==KE 8 >1<5==KE ?@>F5AA>2. 8H8=52, (B88=F0, 1978. 0E0=0 . !, >E>20 . 11.  &8B8@>20=> ?> 0E0=0 . !. >;L 38?>B0;0<CA0 2 3><5>AB075 M=4>:@8==KE 8 >1<5==KE ?@>F5AA>2. 8H8=52, (B88=F0, 1978. 0E0=0 . !, $0< "E8 !CM=   :=.: 0CG=0O :>=D5@5=F8O ?@>D5AA>@A:>-?@5?>4020B5;LA:>3> A>AB020 8H8=52A:>3> C=825@A8B5B0, 1979. 5==>= . $878>;>38O M<>F89. ., 1927. 8@8;;>2 . ., 8 !. ., .@35=A . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. 8@8;;>2 . ., 'C28;8=0 . ., C@8;5=:> . .   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 1978. >@=520 . ., %08B . .   :=.: >?@>AK D878>;>388 2535B0B82=>9 =5@2=>9 A8AB5<K 8 <>765G:0. @520=, 74.  @<!! , 1964. >@=520 . .   :=.: "578AK 4>:;04>2 II A5A>N7=>9 :>=D5@5=F88 ?> 2>?@>A0< D878>;>388 2535B0B82=>9 =5@2=>9 A8AB5<K. @520=, 1966. >@=520 . .  $87=>;. 6C@=0;. 1969, 55, I, A. 93 98. >ABN: . $.   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B=8=F0, 1973. >HB>O=F X. !. A=>2K A@02=8B5;L=>9 D878>;>388. !@02=8B5;L=0O D878>;>38O =5@2=>9 A8AB5<K, B. 2. ., 74.  !!! , 1957, A. 16 28. @5?5 . .   :=.: XIV !>25I0=85 => ?@>1;5<5 2KAH59 =5@2=>9 45OB5;L=>AB8. . ., 1951, A. 10 14. @KH5=L . $ >;?0:>2 . ., ":0G .. .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973, A. 271 272. C7=5F>20 -. /(.   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973, A. 215 216. C;038= . ., >20;52 . ., @82>@CG:> . ., (5@5<5BL52A:0O !. .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 1980, A. 143 156. K@35 . -   :=.: "578AK A5A>N7=>3> A8<?>78C<0. 8H8=52, 1978, A. 29. C3>25F . ., 0<F520 . X. 8 4@.   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 1978. 0:0@>2 . ..  #A?5E8 D87=>;. =0C:, 1978, B. 9, ! 4, A. 82 87. 0:>2552 . .   :=.: !B@5AA 8 040?B0F8O. 8H8=52. (B88=F0, 19780, A. 118 119. 0:>2552 . .   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 19786. 0<0;K30 . .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 1980. 167 0@8= . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. 0@BK=5=:> $. .   :=.: >2>5 > 3>@<>=0E 8 <5E0=87<5 F-f 459AB28O. 852, 0C:>20 4C<:0, 1977. 0B59 . . AA;54>20=85 @5D;5:A>2 C @K1. 2B>@5D. :0=4. 48A ., 1960. 0B@>A>2 . . 0BAE>;0<==K =5@2=>9 A8AB5<K ?@8 1>;52>< A8=4@><5. 2B>@5D. :0=4. 48A. L2>2,' 1981. 55@A>= $. 3., 02;>20 . ., 815@ . ., 0@:>2A:0O . . _  :=.: !@5AA 8 040?B0F8O.  8H8=52, (B?===0, 1978, A. 183 184. 55@A>= $. 3. 40?B0F8O, AB@5AA, ?@>D8;0:B8:0. ., 0C:0, 1981. 5;L=8: . ., @820O . ., 3C5= 3>: %>9.   :=.: $878>;>38G5A:85 8 :;8=8G5A:85 ?@>1;5<K 040?B0F88 >@30=87<0 G5;>25:0 8 682>B=KE : 38?>:A88, 38?5@B5@<88 8 38?>48=0<88 I =5A?5F8D8G5A:85 A@54AB20 2>AAB0=>2;5=8O. ., 1978. 5;L=8: . ., 0E0=0 . !, >1C . .   :=.: "578AK 4>:;. XXVIII 564C=0@>4=>3> :>=3@5AA0 D878>;>3>2. C40?5HB, 1980, A. 578. 5;L=8: . ., >1C . .   :=.: "578AK 4>:;. -3> A5A>N7=>3> AJ5740 M=4>:@8=>;>3>2. ., 1980. 5;L=8: .  0;0489 . !, 3C5= !C0= 8=L.   :=.: $0C=0, M:>;>38O = D878>;>38O 682>B=KE. 8H8=52, (B88=F0, 1980. 5;L=8: . ., 5;5C . .  !>AB>O=85 A=<?0B>-04@5?0;>2>9 A8AB5<K C ABC45=B>2 2 ?5@8>4 8E 040?B0F88 : #C 2 7028A8<>AB8 >B B8?>;>38G5A:8E >A>15==>AB59. 8H8=52, (B88=F0, 1980, A. 98. 5=LH8:>2 . ., >;LH0:>20 ". .   :=.: !B@5AA 8 040?B0F8O. 8H8=52, 1978, A. 35 36. 5=LH8:>2 . ., 5@578= $. ., >;LH0:>20 ". ., ":0G52 . .  :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 1978. 8BNH>2 . ., 0:8F:0O . ., (0;O?8=0 . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973, !. 31 34. OA=8:>2 . ., >A52 . ., &>75@5=:> . ., >@4>2A:0O . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. 0C<5=:> . . !5@>B>=8= 8 <5;0B>=88 2 @53C;OF88 M=4>:@8==>9 A8AB5<K. >2>A818@A:, 1975. 0C<5=:> . ., (8H:8=0  ..  @>1;5<K M=4>:@8=>;>388, 1979, 25, 3, A. 49. ?0@8= . . 87=L, 55 ?@8@>40, ?@>8AE>645=85 8 @0728B85. ., 1968. @15;8 . . 5:F88 ?> D878>;>388 =5@2=>9 A8AB5<K. ., 1935. @;>2 . ., 0A8;L520 . ., 070@C : . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. AB0H:>2 . ., @;>20 ". .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K, 8H8=52, (B88=F0, 1973. 0;0489 . !.   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 1978. 0=:>00 . !.   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. 168 5B@>2 . .  ;8=8G5A:0O <548F8=0, 1976, B. 54, ? 11, A. 12- 19. >3>4052 . .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B8==F0, 1980, A. 168 173. @5H5BN: . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B?8=F0, 1973. @>1C . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973, A. 163 164. >1C . .   :=.: "578AK 4>:;. V A>25I0=8O ?> ?@>1;5<5 8-AB>35<0B8G5A:85 10@L5@K, ?>A2OI. 100-;5B8N A> 4=O @>645=8O 0:04. . !. (B5@=0. ., 1978. >75= . . A=>2K M=4>:@8=>;>388. ., KAH0O H:>;0, 1980. CA8= . /-   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B=88F0, 1973. !028=0 . ., 0=:>20 . !, /:>2;520 . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B8==F0, 1973. !028=0 . ., >4K<>2 . ., 0=:>20 . !, 0BC=O= . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. !5;L5 .  G5@:8 >1 040?B0F8>==>< A8=4@><5. ., 54387, 1960. !5;L5 .   :=.: >2>5 > 3>@<>=0E 8 <5E0=87<5 8E 459AB28O. 852, 0C:>20 4C<:0, 1977. !5@>20 . ., 0=35<0: . ., >B0?>20 . ., '5;L=0O . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. !B5?0=>20 !.   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B=HH0, 1973. !C40:>2 ,- -   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. !C40:>2 - .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 1980. "8E>=>20 . .   :=.: !B@5AA 8 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, 1973. ">=:>3;0A . ., $C@4C9 $. ., %0940@;8C !. X.   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 1980. "C@0:C;>2 /. X.   :=.: >2>5 > 3>@<>=0E 8 <5E0=87<5 8E 459AB28O. 852, 0C:>20 4C<:0, 1977. "C@?055 ". . 5480B>@=0O DC=:F8O 0F5B8;E>;8=0 8 ?@8@>40 E>-;8=>@5F5?B>@>?. ., 74.  !!! , 1962. $09B5;L15@3 . ., ;04:89 ". ., !5<8: . ., #40;>5 10. $.   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 1978. $5;L415@3 . $0@<0:>;>38G5A:89 ?>4E>4 : 87CG5=8N <>730. ., 0C:0, 1971. $C@4C9 $. ., 010@M . ., 5;>CA ". .   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 1980, A. 210 221. %0940@;8C !. X.   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B88=F0, 1980. '07>2 . ., A8=G5:>2 . . -?8D87. ., 1974, A. 15. '5@=>:0= . $.   :=.: 5@2=K5 8 M=4>:@8==K5 <5E0=87<K AB@5AA0. 8H8=52, (B8==F0, 1980, A. 235 245. (0;O?8=0 . ., '0?0D8;>20 . .   :=.: !B@5AA 8 040?B0F8O. 8H8=52, (B88=F0, 1978. 169 (0D@0= . 1, ;8A>2 . ., )5@18=0 . .   :=.: !B@5AA 53> ?0B>35=5B8G5A:85 <5E0=87<K. 8H8=52, (B88=F0, i! 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Hormone Res., 1965, 21, 623. Sutherland E. W.  Science, 1972, 177, 401 408. Ungersledt U.  Acta Phvsiol. Scand., 1971, suppl. 367, 1. Welch B. L., Welch A.  Fed. Proceed., 1973, 32, 3, 275. Verney F. B.  Proceed. Royal Soc, 1947, 135.  1. 2545=85............ 3 2. -2>;NF8O B5>@88 AB@5AA0....... 7 3. ;0AA8D8:0F8O AB@5AA>2........ 11 4. >@D>DC=:F8>=0;L=K5 87<5=5=8O >@30=>2 8 :@>28 ?@8 AB@5AA5 . ......., . . . 14 5. 8>;>38G5A:85 AB@5AAK........ 17 !. $878>;>38G5A:85 AB@5AAK........ 26 6.1. 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