Документ взят из кэша поисковой машины. Адрес оригинального документа : http://www.sevin.ru/dissertations/gidrobiol/87.pdf
Дата изменения: Thu Feb 16 16:09:10 2012
Дата индексирования: Tue Oct 2 13:26:22 2012
Кодировка:

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jl lK b \ [ b j k d b c h, k _ Z j^ m ` g Z y jb b ,k _e , ^ i _ e y ,^ vsym d Z b g , p _ s m d e _e_gb kl l__j ,f Zk _ j _ [ j y g u c b a h e jh d Z j, i i _ k d Z j v a, v ] h e v y g i ,y , \ v x g g Z e b f j r, h d m, gk m ^ Z , ,_ v, ees , _a B a m q _ g b _ b a [ b j Z _ f h c b JhZ ^ h g gZ y n h ,j _ _ v k b g , p dm` q Z d mZ dZ l _ e v g h b a [ b j Z _ f h c l _ f i _ j Z lZ j j ,uk vmj, i i e h l \ ,Zh d m g v ju[ jZagh]h \hajZklZ ees , _a H i j _ ^ _ e _ g b _ k m l h q g u o b k J Z ^ h g` g Z o n h ,j _ _ v k b g , p _a m guy Z hkh[_gghkl_c l _ j f h b a [ b j Z d Z jyZ k v j, i i e h l \ ,Zh d m g v g b , dm ju[ jZagh]h \hajZklZ J b _ ` bo ees , _a B a m q _ g b _ w d h e h ] h n b a b h e h ] Z ^ q m kgdZ y n h ,j _ _ v k b g , p j Z j, v b i h \ _ ^ _ g q _ k d b o h k h [ _ g g h k l b c kj_ j k [ j y g u c d, Z d Z k i i _ Z_ hdm ij_^_e_gby ju[ \ l_jfh]jZ^b_gl_ gv ees , _a H i j _ ^ _ e _ g b _ \ _ j o g _ c k m [ e _Jl Z ^ e v` g Z c n h ,j _ _ v k b g , p Zm hy k _ j Z j, v b e _ l Z e v g h c l _ f i _ j Z l m j u b m _ j u [ j y g u c d, Z d Z k i i j Z a g h ] h \ h a j Z k l Z \ j Z a g u _ h d mhggvu k_a d , , B k k e _ ^ h \ Z g b _ i h \ _ ^ _ g b y -b Jj y ik i j _ Z l _ j e y ^ v[ _ k l _ j e _ Z mr, k , , ^ _ e _ g b y j u [ \ _ k l _ k l \ _ g g-u og _,p k e_ e h ] e Z a d Zq _ o h g v i m [h h d m, gk m ^ Z [d j r , g Z e b f v ,_ \ b y o i h e _ \ u _ ^ Z g )g u _ ( =e Z\Z 3. Hk h[_ gg hk l b jZk i j_^_ e_ g by b i h\_ ^_g by ju[ jZ ag hc w d he h]bb

\ l_ jf h] jZ ^b _ gl gu o m k e h \b y o Ih\_^_ gb_ h^gZ ba ke h`g_ crb o nhjf ijhy \e_gby ` bag_^_y -l_ev \hlgu ,o h[_ki_ qb\Zxs Zy ieZk lbqghk lv ih h lghr_gb x d [uk ljuf b kj_^u :dmklb q_kdb_ . , hilbq_k db_ , ]b^jh^ bgZfb q_kdb_ b we_dlj bq_kdb_ ih , hk \_s_g ghk lv , l_fi_ jZlmjZ b k h ^_j `Z gb_ dbkehj h ^Z kha ^Z xl g_h ^g hj ^u b g_h[ o h^bf hkl v \u[(hjZ lZk ,h\ Ijh 1978; IZ\eh ,\ 1979; =bjkZ1981; Deyr , lhjbg 2). JZk ij_ ^_ e_ gb_ dhjf h \uo h j ]Zgbaf h \ b nhjfbj mxs b o \ , 198 , o bs gb dh \ \h ^h_f_ kbk l_f u ljbhl lZd `_ g_h ^ ghjh(^ gh gl_cn _e v 1987; Fb, jhnZ FZ , 1980, o __ 2006; F b o __ \ b ., ^j ,\ 2010). I hw lhfm ijb h p_g d_ jZkij _ ^_e_ g by b i ju[ \ _kl_ kl\_g guo b bkdmkk l\_g guo l_jfh]jZ ^b_ g lguo m ke h\by l u \ Zl v \e b ygb _ [b hl b q _kd bo b Z [bh lb q _kd bo n Zd l hj h \ . < ijbjh^guo b wdki_jbf_glZevguo mkeh\byo ju[u kZfhijh j Z x l h i j _ ^ _ e _ g g m x l _ f i _ j Z l m j igj m y \ e a h g m_ Z d p b x l _ j f h b a [ b j , h x yy j b e b l _ j f h i j _ n _ j _ g ^ m ,f 1947; B \ e _,\ E _ c a _ j h, \ b q ). I j _ ^ i h e Z ] Z ,x l (Fry Z 1960 q l h a h g Z b a [ b j Z _ f h c l _ f i _ j Z l m j u i j _ ^ k l Z \ e y _ l k h [ h c- h [ e nbabheh]bq_kdh]h hilbfmfZ jhklZ b iblZgby ihcdbehl_jfgu g u o ` b \ h l g u( o \ e _,\ E _ c a _ j h,\ b q 0). D f h f _ g l m g Z q Z e Z g Z r b- o b k B 196 \Zgbc hiulu ih bamq_gbx ih\_^_gby b jZkij_^_e_gby j- [ \ l u g u o m k e h, \ bdyZ d i j Z \ ,b e j h \ h ^ b e b k v \ d h j h l d h f \ j _ f _ g g h f b g l _ o ih f b g m ,l uZ k ). W d k i _ j b f _ g l u [ h e v r _ c i j h ^ h e ` b l _ e v g h k l b [ u e b _ q u (De Vlaming, 1971; Re utter, Herdendorf, 1974; Co utant, 1975; C rawshaw, 1975).

10 Ijb wlhf h [u qgh bkke_ ^ h \Zebk v , h [blZxsb_ \b ^u

\ ij_kgu o \h ^Z o K R: D Z g Z, ^ uZ g g u _ ^ e y j u [ i j _ k g u o \ h ^ h _ f h \ J h k k b b i j Z d l b q _ k d b h ^ eb . Hkh[_gghklb ih\_^_gby b jZkij_^_e_gby ju[ \ ijhp_kk_ l_ gby \ wdki_jbf_glZevguo mkeh\byo . < djZldh\j_f_gguo (2-12 q ) h iulZ o gZ i j b f _ j _ f h e h ^ b d Z j i h \ u o b h d m g _ \ u o \ b ,^ h q \ l h h l _ f a Z jgZ l m j g u c i dZ i_ h \ u [ h j g Z q b g Z _ l k y \ k d h j _ i h k e _ i h k Z ^ d b j u [ \ w d k i _ j b f _ .g l Z e G_ijh ^ he `bl_e vguc 0.5-1 q , i_j bh ^ hagZdhfe_gby k gh \hc h [k l , hdheh k f _ g y _ l k y i _ j _ f _ s _ g b _ f \ ^ j m ] b _ h l k _ d b ] j Z ^ b _ gK g hhce _ l d b Z . l _] mkl e _ s ,Z] m k l _ ,j u Z j i Z e h l \ u b h d m g y i h k e _ Z d d e b f Z p b b d e _ l g _ d ,i 18-20є K q _ j _ a q Z k d h g p _ g l j b j h \ Z e b k v \ h l k _ d Z o k l K f Z _ - Z l m 21-23є _, i a Z j l_f k l_fi_jZl25-29є K 0.05). L Z d h c f _ l h ^ b q _ k d b c i h ^ o h ^ i j b g y l \ j Z [ h l Z o g Z \ k _- o l b gu o m k lZghebg_c gu o ]hjbahg lZevgu o : \hd , \_jlbdZevgu o ]h \uo^ \ mo dZ , djm , f_jgu o bolbhljhgZ ouley, 1977; Reynolds, Casterlin, 1977 Z < gZrb o (McCa ). ^ebl_evgu o hi ulZ o ^ ey hij_ ^_ e_gby ah g 3- oH k m lh qguc djb l_jb u BL ijb gyl k l Z [ b e v g h ] h \ u,[ hdj h ] ^ Z \ l _ q _ g b _ l j _ o i h k e _ ^ h \ Z l _ e v g u o ^ g _ c h Z lh\_jgu_ jZaebqby kj_^g_kmlhqguo agZq_gbc BL hlkmlkl\h\Z .

11 ^jm ]bo k_f_ ckl \ h lf_ q_gu [he__ gba db_ agZ q_gby 26єBL m hdmgy ^h : hl H K 14-

17.0є K m jZ ^ m `ghc nhj_ eb .b gZebfZ AgZ q_gby H BL hleb qZ xlky m l_ ieh [ b \u o \b ^h \ , ju [ jZagbpZ - b oheh ^hex Z HBL m ju[ \gmljb k_f_ckl\Z fh`_l [ulv kms_kl\_gghc beb g . K _ ] h e _ l d b s m d b b gh dq b ggZ x s b _ o b s g b q Z l v \ d h gbpf_ xel _ li Zb , Zm y , _ j f _ j g h l Z d h c ` _ m j h \ _, gdvZ H B L k _ ] h e _ l d b ^ j m ] b o , \ b b l\ Zj x s[ b o k y db i^h u ahhieZgd lhghf ]b_ \b ^u dZji h \uo \u [bjZ xl h q_g v [eba db_ a . Fgh HBL k o h ^gu_ k H BL hdmg_ \uo \b ^h. \ i _ l oe ]hevpZ Barbatulus barbatulus, \ hlebqb_ hl \vxgZ , ba[bjZxl 15.1±0.2є K fheh^v , \ k _ e _ g p _ \ [ u q d Z ] h e hNeogobius kessleri b [ u q d Z p m p Proterorhinus \ZqZ bdZ marmoratus 22-23є K ( <_j [bp dbc b 2005). K_ ]he _ldb kl_jey ^ b ., ^j Acipenser ruthenus ih m jh \gx H BL 2є K [ebadb fheh ^b kb [bjk dh ]hKfbj 23.6±0. ( hk_ljZ gh\ 2009). GZb[he __ [ukljh \u[hj (2-4 BL ) ijh bk o h^be m kZ fuo l_ie , H kml ex[b\uo ju[ М dZjiZ , jhlZ LZ [e bpZ AgZ q_gby HBL m ju [ . 2. . g, k_j_[jygh]h b ahehlh Z jm]b_ bkke_^h\Zggu_ \b 26.5 - 28.0 0+, 1+ Kbg_p ^hklb]Zeb ahgu HBL lhe 26.5 - 27.5 0+ Mde_cdZ g Z 6-9- _ k m .l d Ibh \ _ ^ _ g b _ 26 - 27 0+, 1+ E_s f h e h ^ b b k k e _ ^ h \ Z g g- o \ u 25.5 - 26.5 0+, 1+ =mkl_jZ ^h\ \ ]jZ^b_gl_ l_fi_jZlmj 25.5 - 27.0 0+, 1+ Ya v b f _ e h d Z d h [ s b _, ql Zl u _jd 23 - 26 0+, 1+ Iehl\Z b h k h [ _ g g .h k lgbh ] b _ d Z j F 20 - 21 2+ I_kdZjv i h \ u _ i j b \ u [ h j _ l _ f-i _ j Z 16.5 - 17.0 3+ =hevyg lmju ^_j`Zebkv lhevd JhlZg 0+ 26.5 - 28.0 ]jmiihc H l ^_evgu_ hkh [b . 25.0 - 26.5 0+, 1+ J_ qghc hd mgv hdmgy , ]hevygZ b - jhlZ , _jrZ 24.0 - 25.5 0+, 1+ ?jr gZ qZklh aZlZb\Zebkv \ 22 - 26 0+, 1+ Km^Zd h l k _ d h \_ j b h ^ b q _ k d b f _ g y y ,i
l_fi_jZl m ju h ljZ `Zxl b o l_iehe x [b \h klv b ^ jm ]b_ hkh .[_ gghk l

12 sboky jy^hf fhjnheh]bq_kdbo b nbabheh]h [bhobfbq_kdbo

oZjZdl_jb

G bdhevkdbc, 1980; Ih^ ^m[gu c 1971; Kljm dlm jZ , 1971 , ..., 1991; IZ\ eh\ b ., ^j 2001, 2007; Kl he [mgh\ ; Kl he [mgh\ , 2005 , =_jZ kbfh\ 8). LZd Zy ^bnn_j_ g , 200 pbZpby iha\hey_l hkh[yf h^gh]h \b^Z [he__ wnn_dlb\gh hk\Z dhgdj_l gh ]h \h ,h_fZ _ ]h Zj_Z eZ .\ J Z [hlu ih b am q_gbx l_ fi_jZ ^ b \k p _ehf guo j_Zdpbc ju[ jZaebqguo \gmljb\b^h\uo ]jmii d fhf_gl- gZ m [hlu ijZdlb q_kd b hlkmlkl \h \Z eb .

13 jhlh\ h]h h l\_jk lby b nhjfZ l_eZ (ju[ lhe[ mgh\ K , =_jZkbf ,h\ 2008), i h fg_g b x

^jm ]b o \ \hajZ k 4 ]h ^Z b k lZjr_ ij b ^ ebg_ l_eZ ( EZg ,]_ 67; DZkvy l_ 15-16 kf 19 gh,\ B axfh \ 1997). , < Z ` g h h l f _ l, bq l h \ _ k l _ k l \ _ g g u o m k e h \ b y o g Z m q Z k h d\ _ j- \ h ^ h v , i l ^ Zo ` _ g g u o l _ i e h \ h f m a Z ] j y, ah ^ _ g b Z a f _ j g u _ h k h [ b e _ s Z , b Zid ` h l \ u gghj x l e_ nhjf bj m xl l_ieho he h^hex[ b \u_ ]jmiibjh \db m kdZk :kljZm -b ( :kljZ , 1971; k d Z k b j [ b p d ,Z1984; < b j [ b p d ,Z1988). < j Z [ h l Z o i h h i j _ ^ _ e _ g b x H B ,< k k h l ^ _ e v g u o \ b ^ h \ j u [ \ i j _ ^ _ e Z o , Z ^ jg_hZaegZ q g u o j _ a m e v l Z l h \ - g _ i h Z g h(Norris, 1963; Ga rside, Morrison, 1977; Hall et a l.,1978; Melisky et al., 1980; Calhoun et al., 1982; Fangue et al., 2009).

14

= e Z \ Z 4. Nhjf bj h\Z gb_ jZk ij_ ^_e _g by b i h\_ ^_g b y ju [ \ l_ jf h _glguo mkeh\byo \ hglh]_g_a_ ju[ I jhp_kk bg^b \b^mZevgh ]h jZa \blby `b\hlgu o jZa^_ey_ lky gZ ihke_^h\Z l_evg u_ i_jbh ^u hglh ]_g_aZ , hleb qZxs b_ky ki_pbnb qghklvx hlghr_gb gbafZ b kj_ ^u \_jph1934; JZkk ( K_ ,\ , 1948; Dju `Zgh \kdbc , 1949; >jy ]bg1961; , Gbdhevkdbc . ]_ [mZ ^a_ , 2001) b ih \_ ^_gby \ eh ,\ ( IZ 1979; =bjkZ1981) iha \hey_l iheg__ hk \_lblv jZaeb qgu_ i_ jbh ^u . hglh , Ij_ ^ih eh ` _gb_ h , lhhf ba [bjZ _fZy l _fi_j Zlm jZ m fheh ^ b b \aj ql jZaebqgZ [ueh \ ukd ZaZgh _s_ erguson, 1958). Wlh gZreh k\h _ ih 1958 ] . (F \ l \ _ j ` ^ _ g b _ \ d j Z l d h k j h q g u o h i u l Z o g Z f h e h ^ b j Z (Garg de, Taih, j _ e b ^ m ` sih c n t 1958), h i u l u g Z i e h l \ _ ^ Z e b i j h l b \ h i h e h ` g u _ (Alabasterv Downing, j_ame , lZlu 1958). Ijhl b \hj _ qb \u _ ^Zggu _ ih em q_g u b \ dj Zldh \j _f_gg uo hiu ghd jZgg_c fheh ^ b b g_iheh \ haj(_euo ygdbg McCauley, Read, , LZlv j u,[ 1972; 1973; Otto et al., 1976; Ulvestad, Zar, 1977). Lh evdh ij bf_g_ gb_ [hev rbo e guo ]jZ ^b_gl guo m klZgh \hd b bo lbhljh gh \ iha \h ebeh hij _ ^_e k l Z j r 2-4 e _ (Barans, Tubb, 1974; Reutter, Herdendorf, 1974; Reynolds,Casterlin, _ l 1978b; Ehrlich et al., 1979; Medvick, Miller, 1979). G _ k f h l j y g Zl hl hb k k e -_ ^ h ,q \ Z g b y i j h \ h ^ b e b k v k f _ ` g u o h l j _ a d Z o h g,l h h_^ lg\ _ ja` ^Z e h k v i - _ ^ 2-3 \ i] _Z j iheh`_gb_ N_j]xkhgZ h \u[hj_ \ajhkeufb ju[Zfb [he__ gba H B .L > e y h d h g q Z l _ e v g h ] h j _ r _ g b y w l h ] h \ h i j h k Z g Z f b i j -h \ _ ^ _ lu, ho \Zlu \Zx sb_ \ k_ i _jbh ^u bg ^ b \b ^ m Zevgh ]h .j Za \blb y ju [ L_ jf hba[ bj Zg b_ m e bqb g hd b f Ze vdhg g \ Z y ju[d p b y g Z ] j Z .

15 h q_ \b ^gh , ^jm ]bo l_ieh ex [b \uo \b ^ h , j u [ Z q_gby BL ebrv i hke_hd \ ag 8-9 kml

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j b f _ g l Z f h ] m l h l j Z ` Z l v m j h \ _ g_vg b _B b j Z k i j _ ^ _ e _ g- b _ f .I ^ HL k _ ] h e _ l d h \ b ] h ^ h)\ b\ dm k e h \ b y o l _ j f h ] j Z ^ b _ g l Z k e _ ^ m- _ l l _ h\ _ j g h k l y f b h l j Z ` Z _ l l _ i e h e x [ Z \ h k ldv` _ j uj [ m ] b _ h k h [ _ g g h k l , b lZ ^ .] b b

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16

_ g l g u o m k e h \ .b y h l j _ [ g h k l v \ [ h e _ _ g b a d h c l _ f i _ j Z l m j _ (Ho- k h a Io ^ey

kanson, 1977; Dhr _e_\ , 1984) b h [bl Zgb_ \ ijb ^hgguo kehyo \h ^u - j y^h gh \gufb h [t_dlZ fb iblZg by fh ]m l [ulv hkgh \ gufb ij b qbg Zfb g b a d h c l _ f i _ j .Z l m g u k b l _ e v g h g b a d b _ a g Z q _ g b y H B L m \ k Hlj h ]jmii jZ ^m `ghc nhj_e b hljZ ` Zxl o heh ^hex [b \h klv \b ^ Z b m keh gby \ _kl_kl\_gghc kj_^_ . b g Z f b d Z h d h g q Z l _ e v g h b a [ b j Z _ f h c l _ f i _ - 17° K ^ e y a h e h l h ] h d Z j j Z l m j u m a h e h l h ] h d Z Z Zm ` y h c n h j _ e b k ,y16° K ^ e y e _ 5°ZK b (1), j j ^ k g (2), s e_sZ kbgpZ b iehl \u (3), (4) (5). ^ey jZ ^m `ghc nhj_e b AZ o Zjh1955; = m eb , \Z ^ h \ , 1978; L b l Z j, _ \ 1980; : l e Z..., 2002). k

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J b k . 4. D Z j l Z ko _ f Z j Z k i j _ ^ _ e _ g b y d j m i g uo ( : ) b f_e dbo w o hf br_g_ c ( ;) ( hl ju[ ) gZ mqZkld_ j .

18 I h e m q _ g g u _ ^ Z g g u _ o h j h r h k h ] e Z k m x l k y k h [ s b f b i j _ ^ k l Z \ e _ g- y f b b

g Z f b d _ l _ f i _ j Z l m j g u o l j _ [ h \ Z g btter, jHerdendorf, 1974; Beitinger, (Reu c u [ Fitzpatrick, 1979; E Z i d b g b 1981; B \ Z g h ,\ ZE Z i d ,b 1982; K \ b j k d ,b c Z-i ., ^ j g E dbg1987) b ^ hihe gyxl ij_ ^k lZ \e_ gby h [ baf_g_ gbyo H BL k m \_ , j Z k l(Z _ a g Z q b l _ e v g h _ m \ _ e b2-5є K _ l g Z l k l \ b _ b a f _ g _ g b_ g b _ g q_gb h km , , kgb` c [he__ q_f 10є K GZ ijbf_j_ \b ^h \ ju [ [ueh i hdZaZgh hilbf ZevgZy gZ ). 49 , qlh l _ f i _ j Z l m j Z ` b a g _ ^ _ y l _ e v g h k l b j u [ i h e h ` b l _ e v g h d h j j- _ e b j l _ e v g h b a [ b j Z _ f h c l _ f i _ j Z l ng,j 19c (Jobli m h 81). > Z g g h _ k h h l \ _ l k l \ b-_ i h ey_l jZkp_gb\Zlv HBL dZd djbl_jbc hilbfZevghklb mke j u [ . L Z d b f h [ j Z a\ hh g l h ] _ g _ a _ f Z d k b f Z e v g u _ a g Z q _ g b y H B L , f \ fZevdh \uc i_jbh ^ k_ ]he_l dh \ b ]h ^ h \bdh \ ju [ q_ ]h hgb kgb, `Zxl ,m , ihke_ klZ [bebabj m ykv \ i_ j bh ^ iheh. \haj_e hkl b < h a j Z k l g u _ h k h [ _ g g h k l b j Z k i j _ ^ _ e _ g b y b i h \ _ ^ _ g b y- j u [ gu o l_ jf h ]j Z ^b _ gl g u o m k e h\ by o . =b^jhZdmklbq_kdb_ bkke_^h\Zgby \ jZchg_ _ k l _ k l \ _ g g u o l _ j f h ] j Z ^ b _ g l g u o m k e h \ b c m D h(k h j v d k dkhdch _J W = l hf h\ = \ h ^ h o j Z g b e b sk_ i h k e _ ^ m x s b f h l e h \ h f j u [ \ u y \ b e b j y ^ h k h [ _-g g h k ) i j _ ^ _ e _ g b y b i h \ _ ^ _ g b y g _ l h e, vg h b h\ a jhh^ b e u o h( k b .k_ c M q Z dh f e k j h [ 4). kldb g_ ihkj_^k l\_g gh]h k[jhkZ ih^h] j_ lu o \h^ bkihev a mxl ky \ d ih \ gZ ] m eZ ^ey [h evr bgk l \Z l_ ieh ex [b \u o \bk_ \ ]hju [ dh \ ^h \bdh \ b ^ h e_l , ]h g_iheh \ haj_ eu o hkh [_c e_s ,Z] mkl_ju , iehl \u , mde_cd b . Ijbe_ ]Zxs b_ m qZkldb f_gvr bf \ebygb_f l_jfZ evgu o khkgh \ gh_ f_k lh ^ey gZ ] m eZ- iheh [ jhkh \ eu o hk h[_c [he vrbgkl \Z \b^h\ i h^ \ha^ _ck l\b_f ih\ur_ gghc l_f i_j .M ijhbkoh ^b l kf_s_ g b_ kjhdh ,\ dhlhju c aZdZ g qb \Z_ 1-3 g_ ^_eb g_ j_klZ lky gZ jZg vr_MklZgh \e_gZ \_k_ggy y fb ]jZ pby iheh \ha j_eh ] h mxs_ ] h . , bkiheva e_sZ ih^h]j_ lu_ \h^u e br v ^ey [he__ jZgg_ .h =jZ^b_g llZ ] g_j_k l_fi_ jZ lmju - \ ^bZ iZahg_ 25-28 ^h 15-20є K h[ m k eZ\eb \Z_ l \hajZk lgmx ^bnn _j_ gpbZ pb hl ^_e_ gby l_ ieh ex [ b \u o \ b ^h \ ju [ \ jZch g_ k [jhkZ ih ^ .h ]j_ luo \h ^

= e Z \ Z 5. Km l hq gu_ b k_ a hgg u_ j blf u jZk i j_^_ e_ g by b i h\_ ^_ g l_ jf h] jZ ^ b_ gl gu o m k e h \b y o . Ijhp_kku `bag_^_yl_evghklb ju[ j Z a f g h ` _ g bj _ k, l j Z a \ b l ,b _ b l Z g,b _ ,h i j Z k i j _ ^ _ e _, g ibh_\ _ ^ _ g b i h ^ \ _ j ` _ g u k m l h q g h c b k _ a h g g h c Gjb l f b _ ( -b dhevkdbc Miller, 1979; RZlm gh\k ,dbc , 1974; 1980). Ohjhrh b a\_ klgu i jbf_j kml h qguo \_jlbdZevguo b ]hjbahgl Zevg uo \hfb ]jZpbc ju [Zf [h , iha eyx sbo e _ _ w n n _ d l b \ g h b k i h e v a h \ Z l v i b s _( \ A _ kj,k 1971; k u t, 1971b; u m _ _ jm j Bret k Ih ^ ^m [gu c FZebgbg , , 1988; IZ \ eh \ b ., 2007). L_ fi_jZlm jgu _ Z ^ZilZpbb ^j \ i h e g h c f _ j _ h l j Z ` Z x l w l h l \ Z ` g u c Z k i _ d l ` b a g _ ^ f b ^ l- e v g (R_y _ Gb _evk_g :eZ [Zkl_ Ee hc ^ 1984; Ha_jgxd Z h ^gZdh kml h qgu _ , 1982; ,j , , 2000 ), k _ a h g g u _ j b l f u j Z k i j _ ^ _ e _ g b y b i h \ _ ^ _ g b y i j _ k g h \ h ^ g u- j u [ o _glguo mkeh\byo g_^hklZlhqgh .bkke_^h\Zgu K m l h q g Z y ^ b g Z f b d Z l _ j f h b a [ b j Z i g^ j b[ g y ba uq Z [ e Z kv \ i _ j b h ^ k h h h jm 1971 ih 1995 ] ] . (Medvick, M iller, 1979). GZb [he__ yjd bc ij bf_j k m lh qg jbh ^b qghk lb l_jfhj_ ] m eypb hggh ]h ihihh q_j_ ^ guc \u [hj ^g_f b \_ ^_gby ghqvx i jhlb \hihe h`g u o agZq_ gbc l_ fi_jZ lmju [he vr_jhsluf Micropteru salmoides b f Z e h j h l u f h d m g _ f Micropterus dolomieui, h [ b l Z x s b o \ h ^ g

19

l _ o ` _ \ h ^ h _ f Z (Reynolds, Casterlin, 19 78b). W l h l k e m q Z c j Z k k f Z l j b \ Z _ l o

ijbf_j jZkoh`^_gby l_jfZevguo gbr m kbfiZljbq_kdbo \b^h\ . M jy ^Z \b ^ h \ ju [ М s mdb fZkd bghg ]Z Esox masquinongy, ` _ e l h ] h Z f _ j b d Z g k d h ] h h d merca flavescens, k _ j _ [ j y g h ] h d Z j Zynolds, Casterlin, 1977b, P gy (Re k y 1979b,c) b ]h ^h \bdh \ f mdk m gZ us muksun ( K \ b j k d ,b c Z e l h g _ g Coregon < , 2002) agZ q_gby HBL \ ^ g_ \gh_ \ j_fy [ ueb \ur_ q_f gh qvx 4-7є K gZ , . G Z r b w d k i _ j b ,f _ j h \u ^ _ g g u _ \ m k e h \ b y o _ k l _ k l \ _ g g h ] h , n h l igl _ ihdZaZeb m jZ ^ m ` ghc nhj_eb gh qghc m jh \_gv ) H BL , qlh (16.3є K ij_ \urZe ^ g _ \ g h c g Z K K m s _ k l \ _ g g h c j Z a g b p u \ j _ Z d p b y o , f h d m, b y d Z 2.5є . eh^g iehl \u b smdb \ jZagh_ \j_ fy kmlhd g _ h [gZjm `_gh iha \heb . Wlb ^Zggu_ m keh \gh jZa ^_ eb lv 3 ju m gZu g_ bf_xsb o jZagb pu H BL ^g_f b ]j [ ii : , ba [bjZxs b o [ he__ \ukhdmx l_fi_jZl mj m ^ . g_f beb gh qv x O h j h r h b a \ _ k l g Z k i h k h [ g h k l v g _ d ,h l h j iuj o f ,_ jf^ h \ e h ^ b g _ j d b gZ b \b h Oncorhynchus nerka g _ lhevdh i jhy \ey lv kmlhqguc jb , gh ]j Zpbc lf fb b ij_ h ^ h e _ \ Z l v \ k q b l Z g g u _ f b g m l u l _ f i _ j Z l m j g u c b g l (Brett,e \ 10-15єC _ j \ Z 1971b). < i h e _ \ u o b k k e _ ^ h \ Z g b y o g Z f, bq l h h \d Z h k e u c e _ s \ J- u [ b g i aj aZgh kdhf b B\Zgvdh\kdhf \h^hojZgbebsZo \ e_lg__ \j_fy j_]me \ _ j l b d Z e v g u _ k m l h q g u _ , f i hj^ g b fbZ y k v h l ^ g Z g Z j Z -7k f ,h Z g b _ b] Zp b 1k l y h l ^ _ e v g u _ h k h^ hb18 f . J Z a g b p Z l _ f i _ j Z l m j u \ _ j o g b o k e h _ \ \ h ^ u [ e_lhf h[uqgh g_ ij_ \urZ_l H^g Zdh ^Z`_ wlh]h ^hk lZlhqgh 2-3єC. ^ey \h gh \_gby k m lh qgu o \_j lbdZevgu. o fb ] jZp bc K_ah gg Zy ^b gZfbd Z jZkij _^_e_g by b ih\ _^_gby f heh^ b . G Z ijbf_ j_ ju[ Zf_jbdZ gkdhc SiZebb fontinalis [ ueh ihd Za Zgh k _ah ggh _ i hgb ` _g b _ BL alvelinus ghy [j_ gZ K k i hke_ ^ mxsbf ih \ur _gb_f \ (Sullivan, Fisher, 1953). 3-5є fZ jl_ ;ebadb_ j_a me vl Z lu \ hk_gg _ abfg bc k_a hg i he m q_gu fh- jZ , gZ eh ^b e_ sZ b ]hj Rodeus sericeus (Garside, Tait, 1958; Zahn, 1963; Lenkiewicz, qZdZ 1964). < lh `_ \j_fy m dZf leuronectes platessa (Zahn, 1963) b wk lmZ jgh ]h P[Ze u [u qdZ Gillichthys mirabilis (De Vlaming, 1971) k_ah ggu o baf_g_ gbc BL g_ j m `_ gh< [ he vrbgk l \_ jZ [hl hi ulu [u e b g_ ijh ^h e `b , h ^gZ dh b b [ he__ . l_e vguf ^ebl_e vgu_ wdki_ jbf_ glu \uy\b eb hlkml kl\b_ k_a hgghc ^bgZf b gby H BL 7-gZ (Barans, Tubb, 1973; Reutter, Herdendorf, 1974). Ijb w lhf - bk 20єC ke_^h\Z ebk v g_ \k_ , k_ah gu Z \ \_k_gg_ hk _ggbo hiu lZo l_fi_jZ lmjZ \Zj vbjh \ZeZ h l20єC, qlh k gb `Z _l a gZ qbfhk lv ihe m q_ ggu o . j_a me vl 5 ^h < gZrbo wdki_jbf_glZo hp_gdZ k_ahgguo baf_g_gbc l_ jZagh]h \h ajZklZ i jh\h ^beZkv \ g_ijh ^he`bl q _evg uo (2-12 ) hiulZo \ hk_ggb i_jbh ^ ]h ,^ Z ^ebl_ev guo ( [he_ _ ihem) ]h ^ZklZg ^Zjl, guo ^he `bl_ evgh klvx b ijh 10-14 k m l .h L _ f i _ j Z l m j Z Z d d e b f Z p b b a b f h c k h a b Z geby_e_Z g h c d 1-4є K( k l f \ ), \ k b hk_gvx K ( \ _k_ggb _ b hk_ggb _ 8-12є ), \ e_lgbc k_ah g K ( e _lgb _ ju. [u 18-22є ) D j h f _ l ,h ] H B L m f h e h ^ b e _ s Z h i j _ ^ _ e y e b \ ^ b Z i Z a h g _ l - f i _ j h _ fZpbb 2hl 30є K \ e_lg bc b ab fgbc k_a hgu ^h . A g Z q _ g b y, \ B y \ e _ g g u _ \ d j Z l d h \ j _ f _ g g u o m i k _ Zho _ l d h \ , e _ s uL , h ul ] e iehl\u dZjiZ b ] m k l_ju vx [ ue b ,g q_f e_lhf i_ j \mx hq_j_^v w l , , hk_g b`_ .< k \ y a Z g h k \ e b y g b _ ,f i k _ a h g Z d m w d k i _ j b f _ g l u i j h \ h ^ b e b- k v \ hkdhev gu o mkeh \by o b hleb qZ ebk v ebrv l_f i_j Zl m jhc Zdde bfZpbb .

20 > e b l _ e v g u _ i h e m ] h ^ h \ u _ h i u l u i j h \ _ ^ _ g u d Z d g Z \ j uy[ Z u o g _ i h - j _ ^ , a lo k

k l \ _ g g h b a \ h ^ ,h _ Z Z b g Z Z d d e b f b j h \ Z g g u, o h k _ g _ l g _ c a b f g _ l fd d eg_c b l_fi_ jZlmj_ \ eZ [hjZ lhjgu o . mkeh \byo K_ ]he_ ldb e_sZ , Z ddebfbj h \Zgg u_ abfg_c l_fi_jZ lmj_ , \ ^_ dZ[j_ n_ \jZe_ b i_ j\hc iheh\ bg_ fZj lZ - ijb^_ ebkv oheh ^ ghc (1-3єgu ]j Z ^b_ g lZ bk 5). Ijb djZ ldh \j_f_ ggu o aZ oh ^Z ah K ) (j . k_dpb b k l_f i_jZ lmjhc h gb k h \_ jrZeb j_a db_i jb hkdb g b h ^g Z 13-20є K , [j w lhf ba j u[ g_ ih] b[eZ b \ l_q_-5 fbg gZ o h^ beZ h[jZ lguc< i h lebqb_ h l 0.5 gb_ . mlv hk_gg b o gb_ e_ sb i jb gZ ebqbb dhjfZ .g_ ihke_^ g _c ^_ dZ^_ f Z j , abf < ib lZebkv ju[u klZeb qZs_ ihy \eyl vky \ ahg_ l_f i_jZlmju 8-11є K Z i hlhf m re b \ , \h^m , ijh]j_ lmx 18-20є K K wlh]h f hf_g lZ b ^h k_j_^ bgu Zij_ ey e_s ba ^h . iZahg l_fi_ jZl mjuє K 13-24 . Ih \_ ^_ gb_ b jZkij_ ^_ e_gb_ k_ ]h e_ldh \ e_sZ h \Zggu o d hk_g , Z ddebfbj agZq_ gbyf l_f i_jZ [ueh ZgZeh ]bqg uf , lmju , lhevdh \ i_j 5-30 k mlhd ihke_ \u_ ihkZ ^ db hgb ijb ^_j `b \Ze bk v l_fi_ jZ .lmju 4-5є K L _ f i _ j Z l m°j K ,Z

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< j _ f y ] h ,^ Z _kyp f J b . 5. K _ a h g g Z y ^ b g Z f b d Z H B L m f h e h ^ b e _ s Z k ( Z b i e h l(\ u \ ^ e b l _- e v ) [) g u o h i u .l 1 l _ f i _ j Z l m j Z \ h ^ u \ \ h, ^ 2 f _ l _ j \ Z e b f h ^ Z B L m j u Zo h_bg . M wlhc ]j miiu lZd `_ aZnb dkbjh \Zg i_ ahgm \ lj_ lv _ c ^_dZ.^_A fZjlZ b h k _ g g b _ bfgb_ e_sZ\ l_ q_gb_ abfu b \ i_j \hc ih eh \bg_ , l Z d b \ u k h (16-22є K l _ f i _ j Z l m Zm i h k e _ ^ dmx ) ,j dhgp_g ljbjh \Zebkv lhevdh \ l_ieh K \h ^ (16-24є c).

XI XII

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21 E_lgb c be b Zd debfbjh \Zggu c d l_fi_ jZlmj_ e_sk ^_ dZ[jy i h Zij_ e 18-20є K ,

g_baf_ ggh \u[bjZe l_fi_ jZ lmjm h. l L_fi_jZ lmj m K e_s ba[_]Ze 14 ^h 25є K 8-11є . Ju[u , k e m qZ cgh \hr_^rb_ \ h lk_db k l_f1-5є K lmjhc b \ucl b i_jZ g_ fh]e , kZfh klhy l_evgh b l_evgh_ ij_[u \Zgb_ \ khk lhygbb r hdZ \_eh d ] dh . >e , h^gZ b[_eb i_j_g_k_ ggu_ \ l_ q_gb_ [eb `Zcr_ ]h qZ k Z \ (18-20є K hgb \hkklZ l_ie mx \h ^m ), gZ\e b\Z eb k \hx `b ag_^ _y l_e vghk lv . LZdbf h [jZa\ hf l_evgu o wdki_j bf_glZ o fheh ^ v e_sZ - b ie , ^eb e y _ l q _ l d m x k _ a h g g m x ^ b g Z f b d m b a [ b j Z _. f H c _lg v x _ \ juZ[ h m ji u hk _fi l j- h \ u r _ g g u o a g Z q _ g b c l _ f i _ j Z l m j u h k_ekZg[ e y _ \ lhkay [ g h \ e . _ lb fyh c ,\ hc h yA k h k _ g g b _ a g Z q _ g b y l _ f8-11єZ l m jlu m ] b \ Z x l , jZ d d e b f b j h \ Z g g u o b i_j K h i u[ e_lgbfb d abfgbf m keh \ byf `_ \j_fy ju [u , . < lh , Zdebfbjh \Zg gu_ -d hk gbf agZ q_gbyf l_fi_jZ lmju be b \ o hehe_s ieh l \Z beb \ l_i , mo h ^yl ( ^gu_ , ), e u _ i e h l \ ) m q Z k l d b ] j Z ^ b. _I g lj Z o h ^ f h e h ^ b w l b o \ b ^ h \ \ _ k g h c b a a ( Z __ l_fi_jZlmj \ e_lgb_ ^Z_l hkgh\Zgb_ ij_^iheZ]Zlv \ebygb_ nZdlhjh\ \ i_j \ m x h q_j_ nhlhi_jbh ^Z , , ^v . Hiu lu ijh ^ he ` bl_ evghk lvx k mlhd \uy \beb ^ \_ ]j mii :u mj u [ ^ g b o k_ 10-14 h ahggZy ^ bgZfb dZ H BL hlkml kl\ m _ l b eb m eZ[h \ujZ`_ gZ gZqbl_ , k ^j m]b o hgZ a gZ lZ[e 4). Fheh^ v kbg pZ ( . , ahehlh]h b k_j_[jy gh] h , dZjZk_ c _ c db b dZjiZmde , smdb i hkl h y g g h \u [ b j Z _ l \ ]jZ ^ b_g l_ \uk hdbc m j h \_ g v f l_f i_jZ lm , Z heh ^ v i_ey^b [ he__ L Z [ e b p 4. A g Z q _ g b y ( H ) m j u [ \ j Z a g u _ k . a h g u .Z KB L _ gbad .bc :gZeh ]b q

22 l_fi_ jZlmju khh l \_lk l \ mxsb_ a bfg_ c b e_lg_c , l_f[ bjZ ebk v , \u i_jZ lmj_ gZb

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Ijb fbgbfZ e vghc ©miblZg ghklb k h »( Z \lhjhc iheh \bgu 25 25 fZjlZ ih \lhj m- x ih eh \bgm bxey ] ^Z ), dh 20 20 ihjh] ij_djZs_gby i b l Z g b y g b6є_K `, 15 15 j Z k i j _ ^ _ e _ g b _b l_jfhj_]meypbhggh 10 10 ih\_^_gb_ fheh^b 55 e_sZ g_ aZ\bkbl l_fi_jZlmju Zdde 0 fZpbb Ju [u, Zdd . 0 1 2 3 4 5 6 7 8 9 1 0 1 1 12 1 3 1 4 1 5 16 12 16 0 2 4 6 8 10 14 ebfbjh\Zggu_ d l_fi_jZlmj_ gb`_ 30 30 6є K ih \k_c \b ^b , [ f h k l, b [ u e b k i h k h [ 25 25 g u \ i Z ^ Z l v \ k i ,y q d m i h k d h e v d m \ -l _ j f h 20 20 ]jZ^b_glguo mkeh \byo khkj_^hlZqb\Z 15 15 ebkv \ ahg_ kZf 11 gbadhc l_fi_jZlmj 10 10 22 ~1є K b gZ o h ^be bkv 33 iZkkb\ghf khklhy 55 44 gbb . Ijb l_fi_jZlmj_ 0 0 1 2 3 4 5 6 7 8 9 1 0 1 1 12 1 3 1 4 1 5 16 6 8 12 16 0 10 14 \ u r _ 14є K \ _ k l -_ k l < h a j Z k u [ , ] h^u jl \_gguo mkeh\byo \ukhdZ J b k 7. M j h \ g b H B L m e _ s Z . ( Z b hd m gy Zddebfbjh h l f _ q _ g Z ) ( [ ), dhjj_eypby f_`^m \ Z g g u o d l _ f i _ j Z l m j.1 k _ a h g 2 i j h f _ ` m _ e_lg, Z bc qbke_gghklvx - hl lh qguc3 \_k_gg_ hk_ 4 abfgbc , , ggbc . ^_evgu o ih dhe_gb dZjih\uo b hdmg_\uo \b^h\ ju[ b l_fi_jZlmjgufb mkeh\by (Broughton, Jones, 1978; Le Cren, 1958; Karjalainen et al., 1996). Ijb ZddebfZpbb d k_ahgguf agZq_gbyf l_fi_jZlmju m ju \uy \e_gu ZgZ eh ]b qgu_ m jh \gb l_jfh ba .[bjZgby ( jbk7).

24

?kl_kl\_ggu_ mkeh\by . DZd aZ jm[_`hf , lZd b \ Jhkkbb\_ ^_g jy ^ , ijh bkke_^h\Zgbc ih hij_^_e_gbx hkh[_gghkl_c k_ahgguo ,b `b ih\_^_gby b jZkij_^_e_gby ju[ \ _kl_kl\_gguo \h^h_fZo b \ j ih^h]j_ luo \h^ =JWK b ( :kljZ mkdZk Coutant, 1975; Fh j^moZc :WK , 1971; ;helh\kdh c , 1975; Gb dZ ghjh \ , 1977; Crowder, Magnuson, 1982 ; Spigarelli et al., 1982). < g Z k l h y s _ _ \ j _ f y \ J h k k b b l Z d b _ b k k e _ ^ h \ ,Z h lb^ _ e- v j _ d gyi gu_ jZ [hlu ijh \h ^y lky ebr v , \ Fhe ^ Z \bb b MdjZ bg_ Ebl \_ . G Z r b b k k e _ ^ h ,\ Zi g h \ y ^ _ g g u _ g Z J u [ b ,g = k d v d h \ k d h f b B \ Z -g v d h jb _ hjhf k d h, f \ h ^ h o j Z g b e b s Z o \ j Z c h g _ k [ j h k Z i h ^ h ] j _ l u o \ h ^ D h k - j h f k l dh\kdhc =JWK iha\hebeb \uy\blv k_ahggu_ hkh[_gghklb ih\_ ^ _ e _ g b y j u< abfgbc k_a hg h lf_ q_gh ijb \e_ q_gb_ ih ]jZ ^b _g lm l .[ fheh^b b g_ iheh\ haj_e u o ju[ \ hl_iey _fu_ mqZk ldb \hae_ - l_ie klZgpbc . <_kghc fhe h ^v ju [ lZd `_ ijb \e_dZ_ lky gZ m qZk ldb k [j hk \h ^ .
= e Z \ Z 6. d W h e h ] h n b a b h e h ] b q _ k d b _ b i h \ _ ^ _ g q _ k d b _ h k h [ _ g g h k l b j Z k ij_^_ e_ gby ju[ \ ]jZ ^b_ gl_ l_f i_jZ lm ju : g Z e b a m h k h [ _ g g h k l _ c i h \ _ ^ _ g b y b j Z k i j _ ^ _ e _ g b y j u [- \ ] j Z lmj ijb ^ _ckl \bb jZaeb qgu o Z [ bh lb q_kd bo b [bh lb q_kd b o n Zdl [hevrh_ dhe b q_kl \h jZ [h l lds, Casterlin, 1979a; Spotila et al., 1979; (Reyno Cherry, Cairns, 1982; Christiansen et al., 1997). F_oZgb af l_jfhj_ ] m eypbh g ih\_^_gby ju[ ^hklZlhqgh keh`_g b hkgh\Zg gZ \aZbfh^_ckl\b g _ j \ g h c k b k l _ f u k l _ j f h j _ p (Sullivan, f 1954; B \ e _,\ E _ c a _ j h,\ b q _ilhjZ b 1960). K l h q d b a j _ g b y n b a, b a g Z q] _ g b _ H B Ll h m k l Z g h \ h q g Z y l h heh bb w beb set-point), n h j f b j m x s Z y i h \ _ ^ _ g b _ b j Z k i j _ ^ _ e _ g b _ ` b \ h l g u o ] j Z ^ b _ g l g u o m k e (Crawshaw, 1977). < d Z ` ^ h f \ h a j Z k l _ b \ d Z ` ^ u c k _ h\byo k m s _ k l \ m _ l k \ h y m k l Z g h \ h q g Zkye _l ^ h q \ d Z _ e, v k \ h _ a g Z q _ g b _. H B , Zl b g h < h i j h k h , ld Z d j Z a e b q g u _ w d h e h ] b q _ k d b _ b n b a b h e h ] b q _ k d b _ hf baf_gy lv oZ jZdl_j guc ^ ey \b ^ Z ^ bZiZahg H BL m ij_k gh \h ^gu \ h a j Z k l g u o ] j m k il Z \ Z e k y h l d.j u l u f , hi

25

a y g ZY h P b 959; Javaid, Anderson, g_ ke [h , 1gg 1967a,b), Z ^ Zggu_ h l_jfhba [ bjZgbb \ ^ ebl_evgu o hiu lZ o hlk m . G Z f b i h d Z aq lZhg h m k e h \ b y o i _ j _ f _ g g h ] h j _ ` b f Z d h j f e _ g b y , \ f h ^ b n b p b j m _ l i h \ _ ^ _ g b _ b j Z k i j _ ^ _ e _ g b _ f h e h ^ b d Z j i-h \ u o ^ h \ j u [ \ l _ j f h ] j Z ^ b _ gZggu_ h l_jfhba[bjZgbb m ju[ \ i_jbh^ g_j_klZ ijZdlbq_kdb hlkmlkl\mxl . >ey \uykg_gby hkh[_gghkl_c l_jfhba[bjZgby \ ij_^g_j i j h \ _ ^ _ g u i j h ^ h e ` b l _ e v g u k m lh i u l u g Z g _ i h e h \ h a j _ e u o b a (12 _ ) h k h [ y o e _ s Z b ,i ekhhl ^ \ _ j ` Z \ r b o k y \ e Z [ h j Z l h j g u o m k e h \ -b y o u e _ g g u o b a \ h ^ h _ f Z \ w l h l ` _ h l j ._ a h d \ j _ f _ g b A g Z q _ g b y kB L h k l Z \ b f u _ k l _ f i _ j Z l m j Z f b g _ j _ k l Z b _ k l , hi ^ u , m g _ i h e h \ h a j _ e u o b i h e h \ h a j _ e u o h k h [ _ .c G[ u e h e jhZh-a jb_q g _ib \ ae e u _ e _, s h l e h \ e _ g g u _ \ \ h ^ h _ f _ k l _ f i _ j Z l m j h 8°C,h m ` _h d2- e h b c \ ^u gh Z h c ^ _ g v h i u l Z i _ j _ r e b \ l _ f19-22Z l m j m e h ] b q g h \ _ e b k _ - y b i _ j °C. : g Z [ e h \ h a j _ e u _ ,e _ sg Z d h i h 4- e _ k m l h d h i u l Z q Z k l v h k h [ _ c \ _ j h^ b ko l _ f i _ j Z l m j m C. G Z _ k m l d b \ k _ g _ i h e h \ h a j _ e u _ j u [ - k d 10-13° 10u j h \ Z e b k v \ h l k _ d Z o k l _ f i _ j C, l m j h h e h \ h a j _e u _h l k _ d Z o k 7-11° Z Z i c \ l _ f i _ j Z l m j h c C. I h e h \ h a j _ e Z y i e h l10-12- _ k m l d b \ u [ b j Z e 24-25° \Z gZ l _ f i _ j Z l m j m °C. ; h e _ _ d j m i g u _ h k h [ b i j h ^ \ b ] Z e b k v i h - ] j Z ^ b 24-27 i_jZlmju f_^e. gg__ _ ; b h o b f b q _ k d b c Z g Z e b ,a q l ihh d Z a Z g h r _ g b _ [ _ e d h \ k u \ h j h l d b khhl e m g _ i h e h \ h a j _ e u o b i h e h \ h a j _ e u o e _ s _ c j Z a e b q Z e h k-b k g _ , aZ v d e x q _ g b _ f k h h l g h ъ -_ b b y ] e h [ m e b gdhh\ l h j h _ m g _ i h e h \ h a - _ e u rg , j [ _ c [ u e h k ^ \ b g m l h \ i h e v a m M _ i hueoh \ h a j _ e u o k Z f h d h [ g Z j . i j\ h [ u q g h _ \ _ k _ g g _ _ i h \ u r _ g b _ d h g p _e g [ m je Zgp \ b \ bd j h \ b , m g b o . 2] hl b h ` _ h l f _ q _ g h g Z b [ h e v r _ _ k h ^ _ j ` Z g b _ Z e v [ m f b g Z < lhh[ s __] h .b ` \ j _ f y m i h e h \ h a j _ e u ,o h e e h _ c g g u o b a \ ,h \ ^ hk_j fZ \ g _ g b b k - ] j Z ^ b l_s\e_ Z _ g l g u f b h k h [ y f 10 g_Z v h i u l h [ g Z j m ` _ g h i h g b ` _ g g h _ k h ^ _ j ` Z b^g ,Z Z e v [ m f b g Z b i h \ u r _ g] eg h _ b g. h \ , q l h f h ` _ l [ u l v k \ y a Z g h k Z d h[me 2 a Z p b _ c ] h g Z ^ h h [ j Z a m x s b o . iWhdpk_i k k b f _ g l Z e v g Z y i e h l -Z i h j _j h\ \

26 l _ e y f [ _ e d h \ h ] h h [ f _ g Z [ u e Z i h q l b ] h l h \ Z d . _ jh_e mm_ g g u _ gI kl q

k \ b ^ _ l _ e v k l \ m x l h k h h l \ _ l B L \ bm j H kl b L ,Z 32 _ f i _ j Z l m°j K
28 24 20 16 12 0 32 28 24 20 16 12 0 32 28 24 20 16 12 0 5 10 15 20 25 30 \ 5 10 15 20 25 30 [ 5 10 15 20 25 30 Z

1 1 2 2

K mdb l J b k . 8. K j _ ^ g _ k m l h q g Z y b a [ b j Z _ f Z y l _ f i _ j Z lmjZ fheh ^ b e_s Z l \u( [ ) b hd mgy ( Z ieh ), (\) ijb jZaebqghf j_`bf_ dhjfe_gby i_jbh ^1 dhjf heb ]h o _ lu b ^Zn gby . 3 jZaZ kmldb dhjf hlk m lkl\ m _ l ,2 .

^Zggu u[ bo nbabheh]h [bhobfbq_ k h k l h y g bG _ i h e h \ h a j _ e u c .x e_, bajZkoh^h\Z\ s abfgb a Z i Z k u ` ,b k Zj _ f b l k y - \ l _ jl i e u _ \ h ^ u j Z g, v q r__f i h e h \ h a j _ e u _ h k M [ b i h -e h .h \ h a j _ e h ] h e _ s Z \ k e-m q Z _ l h \ g h k l b d g _ j _ k l m- g Z [ _ l k y m k l h c q b \ u c \ u [ -h j \ u k h d h c l _ f i _ j , l m jl u - l Z\ h km kl\b_ ]hlh\ghklb beb ih^ ^ysbo mkeh\bc ^ey g_j_k \ha\jZl \ [he__ ijhoeZ^g a h g. u I h e h \ h a j _ e Z y i e h l b a [ b j Z _ l l _ i e m x \ h- ^ m k j _ ,_ q _ f i h e h \ h a j _ e u c e _ s . < u [ h j i h g b ` _ g g h c l- _ f i _ j Z l m ,j u h \ k _ c \ _ j h y l g h k l b i , g_h[oh^bf ^ey khaj_\Zg ] h g Z ^ ( D h r _ e ,_ 1984). > _ -c \ k l \ b l _ e v gm Z f _ j b d Z g k d h ,h ` _ e l h ] h h d mPerca flavesgy cens \b ^Z, [ebadh ]h j_ qghf m h d m , xk h a j _ \ Z g b y i h e h \ u g i j h ^ m d l h \ g _ i j h b k o_hk^ b l ,e b m j h \ _ g v a b f g _ c -l _ f i lmju g_ himkdZ_lky 12°C (Hokanson, 1977). LZdbf h [ j Z a h fl _ f i _ j Z l m j Z C, , 9-13° g_ y\eyxsZyky hilbfZev ^ e y j h k l Z b i b, l Z g b y _ _ kdhj \k_]h g_h[oh^bfZ ^ey [he__ , w n n _ d l b \ g h ] h k h a j _ :Z g b y .\ ] j Z ^ b _ g l l _ f i _ j Z l m j u- \ u k l iZ_l \ dZq_kl\_ ^bj_dlb\gh n Z d l h, j g Z i j Z \ e y x s _ ] h j u [ Z d f _ k l Z f k h a j _ \ Z g b y- b h ghfm h [t_d lm iblZ gby . G_ k e m q Z c g h\ , _kl_kl\_gguo mkeh\b \ l _ _ fl h ] b Z ^ b _ g l g u o ej gj c \h l g _ j _ k l b \ r b _ k y h k h [ b e mo h^yl \ [he__ ijh o eZ^g m q Z k l d b \ h ^ h _ f Z ^ e y. g Z ] m

27
ih l_jfhba[bjZgbx m [hevguo ju[ [ueb ijh\_^_gu gZ k_j_[j , [hevr_jhlhf hdmg_ b ]he m [hf kheg_ q g bd_ Lepomis macrochirus c j Z a e b q g u f b \ h a [ m ^ b l _ e y f b a Z [ h e _ \ Z g b c s hydrophila, Escherichia coli, Staphylo Aeromona coccus (Reynolds. et al., 1976, 1978a,b; Kluger, 1979). I h d Z a Z glh [ h e v g u _ ,qh hkh[b ijhy\eyxl \ ]jZ^b_gl_ l_fi_jZlmju j_Zdpbx ih\_^_gq_kd М \ u [ h j [ h e _ _ \ u k h d h c l _ f i _ j Z l m j u i h k j Z \ g _ g b x 1978; B L (Kluger, k H P u] Z g b .,^ j1997). < g Z r b o h i u l Z o b a m q _ g h \ e b y g b _, \ a Z \[ Z e _ \ u o b c ua h gg Zg Aeromonas punctata b Saprolegnia sp. gZ HBL m fheh ^b dZjiZ \ l_jfh ]jZ g u o m k e h (\ b Z oe 5). l y [. H[s bc oZjZdl_j \u[hjZ H BL L Z [ e b p5. A g Z q _ g b y H B L Z (° K m a ^ hjh \uo a^hjh\u o b [he vguo ju[ - kms ) k l \ _ g g h g _ j Z a e bgb hiulZ Bgt_dpby kl_cM ju [ dhgljhevghc ]j m i . \ha[m^bl_ey 5-6 8-9 10 - 11 \u [hj H BL ijhbk o h ^be lhev gZ8-9 kmldb hiuM [hevgu o . lZ g_l 25.5-27.5 29.5-29.5 29.5-29.5 ju[ \u[hj ijhbkoh^be [uklj__ dhgljhev ) М aZ k mlhd 4-5 . JZagbpZ agZ q_g A. punctata, 29.0-31.5 32.3-32.8 32.8-32.4 ^h hiulZ HBL m a ^hjh \uo b [he vguo A. punctata, [ _ c \ p _ e h f k h k l Z 1.0- e Z \ey 29.8-30.3 30.5-32.1 32.1-31.8 gZ kmldb 5 3.1° K . Saprolegnia, >ey fheh^b e_sZ b iehl 30.0-31.0 32.4-32.0 33.0-32.3 gZ kmldb 5 ihdZaZgu ^hahaZ\bkbf- _ w u l u j _ Z d p b b i h \ _ ^ _ g q _ k d h c e b o h jhZ ^ e Z [ e _ g b _ k h \ j _ f _ g _ - i h k , __ kdb f ^_gby A.punctata, Z l Z d ` _ [ h e _ _ k e Z [ h _ i j h y \ e _ g b _ i j b i h \ l h j g h \ h a [ m ^ b l _.e ; h e v g u _ h k h [ b e _ s Z b i e h l \ u b a [ b j Z x l a g Z q _ g b y y gZ2-6 q K \ u r _ i h k j Z \ g _ g b x k h a . h > h \ u f b y j _ Z d p b y a Z n b d k b ^ j ZggZ ijb \k _o l_ fi_jZlm jZo Zdd ebfZpbb Ihlg m ^ b l _ e v g h _ a Z ^ _ j ` b \ Z 10 ^h 30 q K j b . g b _ f h e h ^ b d Z j i Z \ a h g _ H B L i h k e _ b g t _ ijb b b^bl d ]b [ _ eb A.punctata d p \h j u [, i j b \ h a f h ` g h k l b \ u [ h j Z [ h e _ _ \ u k h d h c l _ f i _ j Z l m j u j u [ u . < _ j h y l, gih \ _ ^ _ g q _ k d Z y e b o h j Z ^ d Z m k b e b \ Z _ l k h i j h l b \ e y _ f h k h k i h k h [ k l \ m _ l \ u ` b \ Z _ f h k .l b gj lu_[ j _ k gl h a h e h l h f m d Z j Z k x b d Z j B ,q h i_j_`b\Zgby [he_agb ijboh^blky moh^blv \ ahgm aZij_^_evgh jZl m ju l30-32 ^h 34-35 q K ] ^ _ b o k m s _ k l \ h ,\ Zi g b Z g b _ b j h k l y \ -g h m (h ), bl _ l _ g ,u Z \ h a f h ` g h k l v i h \ j _ ` ^ Z x s b o b e _ l Z e v g u o i h k e _ ^ k l \ b c \ _ k (Anitha et al., 2000).

28 jhi_il b ^h \ \ \h ^ beb q_j _a o jhg b q_ kdb \ `b \e_ggm x dZ gxex \ i _j \uc `_em ^h -

q_d fha ]Z ju [ i_ j _ ^ g Z q Zehf h iulZ k ^habjh \ dh cfdhfh[ _abgZH 10 [ ] kh, [v IE 50-100 f d ] h k h. [ v _ ] b k l j b j h \ Z e b l Z d ` _ ^ \ b ] Z l _ e v g m x b i h b-k d h \ J g h k l v b k k e _ ^ m _ f. u o j u [ D Z j Z k vd d e b f b j h \ Z g g u c d l _ f i _ j K lq _ j j__a ,Z 15є Z m 0.5-3 q \ u o h ^ b e \ a h g m , l_fi_jZlmj 25-28є K I h k e _ b g t _ d p b b I E H k d h j h k l v \ u o h ^ Z , g _ b . h ^ g Z d hh k l b ] g m \ ,^ 23-25є K j u [ u h k l Z \ Z e b k v \ w l h c l _,f k _ j\ _ jl rm jy - i _ , ihZ Z_ jbh^bq_kdb_ i_j_f_s_gby \ [he__ l_ieu_ beb oheh^gu_ hlk . Ijb bgt_dpbb [hf[_abgZ ijhbkoh^beZ aZ^_j`dZ ju[ \ hlk_dZo 16-17є K b e b r v k i 1.5k q l y u [ u \ u o h ^ b e b g Z k l Z [ b e v g u c m j h \ _- g v b m j f h c l _ f i _ j Z l 23-25є K mju . K l _ i _ g v \ e b y g b y [ h f [ _ a b g Z g Z l _ j f h b a [ b j Z g b _ m j u [ -f _ g y k l b h l l _ f i _ j Z l m j u Z d.d Iejb f Z p b b b f Z p b b d l _ f i _ j K l g Z _- x b Zdde 28є Z m j [ e ^ Z _ f h _ \ l _ q _ g b _ i h e m q Z k Z i h g b ` 2-3єbK h [ m kg Z \ e _ g h m \ _ e b q _ g _g _ BL eh ^ \ b ] Z l _ e v g h c Z d l b \ g h k lM j u [ j Z k _ cd e b f b j h \ Z g g u o d l _ f 15-j Z l m . b dZ , Zd i_ 16є K h l f _ q Z e h k v i h e g h _ h [ _ a ^ \ b ` b \ Z g b _ 1\ q , _i q h k e _ q _ ] h i j h-b k o h , l _gb_ ^ b e i h k l _ i _ g g u c \ u [ h j a h g u d H e bL Z p b y j u [ d l _ f i _ j Z l m j\ _- e b . : dB f 5є K m _ q b \ Z e Z \ j _ f y h [ _ a ^ \ b ` b \ Z g b y q , ^ h u [ u g Z o h ^ b e b k v \ k h k l h y g b- b h p 9 j gby bgh ] ^Z gZ ,[hdm ^ \ ahg m H BL [ue _s_ [he__ .f_ ^ e_gguf , \uoh < ^bZiZ ah g_ l_fi_jZlmju gb`_ hilbfZevguo agZq_gbc kgb`_gb_ BL m f _ g v r _ g b _ f ^ \ b ] Z l _ e v g h c Z. d l b \ g h k l b LZdbf h [jZa\ \_ ^_gb_ g_cjhi_ ilb ^h \ [ _abgZ b IE H b \h ^b l d , hf [hf ij a Z f _ ^ e _ g b x \ u [ h j Z l _ f i _, j Z lf m g u r _ g b x ^ \ b ] Z l _ e v g h c b i h - k d h m _j v b l b \ g h k l b j uh[ l j Z ` Z y g Z q Z e v g u _ k l Z ^ b b i _ j _ o h ^ Z j u. [ W l bb - jh [ b h , d ] i_ a me v lZ l u k \ b ^ _ l _ e v k l \ m x l h [ mk b e _ g b b j h e b g _ c j h i _ i l- ^ h \ \ b g_gby m ju [ ijb l_fi_jZl mjZ o gb `_ 5є K . Ijhy\e_gb_ jZaebqguo nhjf ih\_^_gby ju[ \ l_jfh]jZ^b_glguo \byo . L_jfhj_]meypbhggh_ ih\_^_gb_ y\ey_lky qZklvx ieZklbqghc ihebnmgd p b h g Z e v g h c k b k l _ f u i h \ _ ^ _ g q _ k d b o\ d Z xZZ l Z p b c j Z a g h h [ j Z a g u , e ^ q ixs_c k h q _ l Z g b y \ j h ` ^ _ g g u o b i j b h [ j _ l _ g g u o j _ Z d p b c b (n h j\ e h,i\ h \ _ ^ _ IZ f D Z k m ,f 1994; 1996). K m s _ k l \ m x l j Z a e b q g u _ n h j f u i h b_k^ _ g b yh \ Z yg \ ke_^ l _ e v k , h _ j j b l h j b Z e ,v k h p b Z e v g hh_j h g b l _ e v g h _ j h ^ m d l b \ g h b dl_ g_ , h[ , j_i , i_ s_\h_ b . ^ (j FZgl_cn , 1980, 1 987; Fbo__2006; Fb o __\ b ., ^ 2010). Fgh _e v ,\ j ] b _ n h j f u i h \ _ ^ _ g b y i j h y \ ,e d Z l k y j Z \ ,b \ e l _ k g h c k \ y a b ^ j m ] k. ^ j m ] yxd i h < aZ ^Z qb gZr_c jZ [hlu \oh ^beh ba m q_ gb_ jZaeb qgu o nhjf i , ijhy \eyxs b o ky \ l_jfh ]jZ ^b_glg u o . mkeh \by o B k k e _ ^ h \ Z l _ e v k d h _ i .h \ I h_i g b _\ m k e h \ b y l _ j f h ] j Z ^ b _ g l g h c k _^ Z\ ] h j b a h g l Z e v g u o m k ,l Z u [ h \ d \ Z g m ` ^ _ g u h j b _ g l b j h \ Z l v k y d Z d g jgu uo g h j h ^ g h k l v l _ f i _ j,Z ll Z d u b g Z d h g k l j m d l b \ g u _ h k h [ _ g g . k l b mj h W l h i j _ ^ i h e Z ] Z _ l a g Z d h f k l \ h k g h \ h , i h [ k e _ ^ g hx\ d h c \ u [ h j l h c c klZ m sb b e b b g h c l _ f i _ j. Z< h j\ u _ o h i u l Z o g Z q Z e v g u c i _ j b h ^ a g Z d h f k lm k q_gby ijhbk o h ^be h ^bgZdl_ \h . < h q_gb_ hl ihe m 3- o q ju [u ijb ^_j `b qZkZ ^h \ Z e b k v l _ f i _ j Z l m j u i j _ ^ \ Z j b l _ e v g h c. W d h e[ b f Z pjb b\ g h f _ j g- h _ i Z ld ueh Z \Zgb_ fheh ^ b ihh ^bgh qd_ beb, [he__ djm igu_ ju [u ih qlb g_ ^ \b \ ]j mii_ -

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j _ c l b \ k h k _ ^ g b _ h l k _ d b k [ h e _ _ o h e h ^ g h c b e b l. _ Ji Z h c blp Z i e ag _f l _ f i _ j Z l m j u \ k f _ ` g u o h l k _ d Z o 2-3є k. l Z \ Z y _ f j u [ u g Z q b g Z e b khK A el eZ \ Z b \ Z l v b [ h e _ _ ^ Z e v g b _ m q Z k l ,d lb_ f ji Z ^ jbZ lg l j Z \ h ^ u \ d h l h j-u o h ] _ _ mZ q Z e Z k v h l b k o h ^ 4-6єcK g b [ h e _ _i m k 4-6 q m ` _ i j h y \ e y e Z k v l _ g ^ _ g gh Z .K ly \ u [ h j Z h i j _ ^ _ e _ g g h c l _ f i _ j Z l m .j gD c b k o h ^ u i _ j \ u o k m l h d i j h h ahg m h [ m q _ g , y Z d i j Z \ ,b a Z \ _ j r Z e k yg h ] ^ Z j u [ u \ j _ f _ g g h h [ j Z a h \ u-\ Z e b bd eh .B kdhevdh f_edb o ]j mii beb3-5 hkh [_ cM fheh ^ b g_dhlhju o \b ^ 2-3 ih . h d m g_vj r, g Z e b f] h e v y g h l Z gb k k e _ ^ h \ Z l _ e v k d h _ i h \ _ ^ _ g b _ -q Z s _ , , ,j ) ^beh bg ^b \b ^ m Zevg h , m fheh ^ b e_sZ \u ] m kl_ju , iehl , , kbgpZ М \ ]jmii_ . GZb [ h e _ _ l _ i e h e x [ b( \ d Z j, i k _ j _ [ j y g u c b a h e h l h ,c j d ZZ Zbk b h e h ^ h- e x u_ hl jg o ) [ b \ u _ ( j Z ^ m ` g Z y n h )j _ u v k j Z \ g b l _ e v g h [ u a Z j h m l d b b e b g )_ k d h je[u (kl k \u [bjZeb ahg m . H BL ^_ kem, qZ _iZ ^Z y \ hlk_db k l_fi_jZ lmjhc < jy ih \ 25-32є K , j u [ u k h \ _ j r Z e b j _ a d b _ , [ \ juhi k u ] b \ Z y b a \ h ^ W l b h k h [ b l _ j y e b h j b jdb .u l Z p b x \ i j h k l j Z,g g _ \ _ h ] e b b a [ _ ` Z l v o h e h ^ h \ h ] h b e b l _ i ,e b \ h ] h r kl f h\ , dhg_ qghf bl h ]_]b [Z eb , ih . L_jjblhjbZevgh_ ih\_^_gb_ \ ] j Z ^ b _ g l _ l _ f i _ j Zl m j u g Z [ e x ^ Z e h k v k j Z \ g b l_evgh j_^dh, \ hiulZ o gZ k_]he_ ldZ o e_sZ b hdmgy ihdZaZ gh . LZd , qlh _^bg bqgu_ hkh [b beb g_k dhe vdh j u [ aZg bfZe b h lk_d k H BL b g_. Ih- k dZ im iuldb hk lZe vgu o ju [ \hc lb \ w lm ah g m j_rb l_e vgh ij_k_d ^hfbgZg lZfhgb \ul_kgye b ijbr _e vp_\ , khh lk_dZ ,b ih djZc g_c ,b ba ijh\h `^Zy o f_j_ gZ hlk_ dZ \gba ih ] jZ ^b_g lm l_fi _jZlmju ^gb c k aZg bfZ_f ,2 . H lk_d hk_ ,k ^hfbgb j mxsbf b hkh,[y f b lhy ggh dhglj heb, jh \Zeky ihk qlh [ u khi_ jgb db g_ ih iZ ^Zeb ^Z `_ \ g_M qb lu \Zyqlh jZ agbpZ l_fi_jZ lmju f_ ` ^ m khk_ ^g bf . ]h , dZfb [ueZ K l_fi_ jZlmjg uc [Zj v_j f_ ` ^m ^hf bgZ glgufb hkh [yf 2-3є , lZevgufb k_]he _ldZfb k hk lZ\eye h dheh 4-6є K . > j m ] h c i j b f_ lj _ j f h b a [ b j Z g b _ m k _ ] h e _ l d h \ j Z ^ m ` g h c n h j _ e b l_ l_fi_jZl mju ^h 24є K H^bgh qgZ y hkh[v ba[bjZeZ ahg mK H BL 8 hl . 14-16є . ? k e b \ h i u l _ i j b k m l k l \ h \ Z e b ,l jkb_ ] j u [ _ l h d ^ h f b g Z g l a Z g b f Z e w heu \ p_gljZevghc qZklb ]jZ^b_glZ b \kyq_kdb ij_iylkl\h\Ze ijh g _ _ ^ \ m o h k l Z. e D g u^ Z y b a h k l Z \ r b o k y j u [ i u l Z e Z k v i j h g - d g m vZ` o b g m g h h ^ g Z [ u e Z \ u l _ k g _ g Z \ h l k _ d b k 9-10є K_ ^ jj Z l ] m h c є K , l _ f i , m Z y 20-21 , j m k e h \ b y y \ g h g _ [ e Z ] h i j b y l g u _ ^ e y j h k l W l b ij b l[ u gi b y q _ j _ ^ g h .Z u Z hh \ i e u \ Z e b \ h l k _ d k h i l b f Z e v g h c l _ f i _ j Z l m j h c ,b h kghZkd_h^ g b c k ^ - g_ f _ ^ e _ g g h b a ] h g y e b k v g Z i j _ ` g _ _ f _ k l h k _ ] h e _ l? k e b ^ \ h fl b g Z g . dhf u _k g_gb_ h ^ghc ba ju [ gZ dhjhldb c ijhf_ `mlhd \j_f_g b hk \h [h ` ^Z , b ^ j m ] Z y b a j u [ k h i _ j g b p \ i e u ,\ ZkeeZ ^ h \ g _ _ g _ f _ ^ e _ g g h _ b a ] g Z _ \ Zeh b a w l h ] h h.l k _ d Z y k b l m Z p b y i j h ^ h e ` Z e Z k v \ 10 _ q _ gl b _ \ h .i_uol Z LZdZ l km hd k L _ f i _ j Z l m j g u c [ Z j v _ j f _ ` ^ m ^ h f b g Z g l g h c h k h [ v x b h k l Z-e v g u 4 6є K B g l _ j _ k gl h l _ j j b l h j b Z e v g h _ i h \ _ ^ _ g b _ m l _ i e h e x [ b \ - o b . ,q h u [b \u o \b ^h \ hlf_ qZehk v \ ^bZiZ ahg_ l_ fi_jZl m ju : m e_sZ b , [ebadhc d H BL hdmgy 24-26є K m n hj_eb ijb , 14-16є K . I b s _ \ h _ i h \ _ ^ _ g b _ aZ ^Z qm ^Zggh ] h jZa ^_ eZ \oh ^beh ba dZq_ gb .< , md hlj_Z]bjmxl ju[u gZ i_j_ghk dhjfZ \ hlk_db k [he__ gbadhc -

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\ ^ b Z i Z a h g _ k d h j h k l _ c g Z ] j _ \ Zh 50 ^ q uK hq l \ a g Z q b l _ e v g h c f _ j _ a 4 ^ \h k y l h l k _ aFhbggZb f Z e v g u _ a g Z q _ g b y i h d Z a Z l _ e y m \ k _ o -b a m q . f _ q _ g u a b f hk \ b ^ _ l _ e v k l \ m y h k g b ` _ g b b l _ j f h m k l h. c E _hl k lf b j ,c qb\ h agZq_gby DLF fZdkbfZevgu b \ f_gvr_c kl_i_gb aZ\bkyl h \h ^u . Kh \f_klgh_ i jbf_g_gb_ f_lh ^h \ OEF b DLF i ha \hey_ l b k k e _ ^ h \ Z l v l _ j f h Z ^ Z i l Z p b h g g u c i h l i_j g p w Z h f j O [ E h \ k _ c - \ _ , b b l e u, i F j h y l g h k lhbl j Z ` Z _ l \ b ^ h \ h c f Z d k b f, m f D L n b a b h e h ] b q _ k d m-x i e , Z E LF klb qghk lv hj ]ZgbafZ ju [ . < e b y g b _ k _ a h g Z g Z m j h \ _ g v < E L i h ^ l \ _ j ` ^ _ g h \ w d k- i _ j b e_ldZ o j hlZgZ k bkihe vah \Zgb_f kdh jhk ^h 10° gZ ]j_ \Z. 7). 0.2 l_c K q ( lZ [e hl F Z d k b f Z e v g u _ a g Z q _ g b y D L F b E L h lf fb_gq _ g Z e e _ luh_f a g Z q _ g b , bf u vg a b f h ( j < 0.05). G Z b [ h e v r Z y l _ j f h m k l h c q b \ h k l v m k l Z g h \ e _ g Z i j c k d h j h k l b g Z 0.2-0.4° K ,q f b g b f Z e v g Z i j b k d h j h k l b g 5-10° \ K .q ]j_\Z y Z]j_ Z AgZ q_gby DLF gb `_ 1-2° K gZ agZ q_gbx D LF b EL jh lZg gZ EL . Ih [ e b a h d d l Z d b f l _ i e h e x [ b \ u f w \ j b l _ j f, gd Z d \ bkZ fj _ [ j y g u c d Z j Z uf ^_ dZji .

34

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35 G Z i j b f _ j _ f h e h ^ b e ,_ i e h l \ u b i _ e y ^ b i h d Z a Z g h be b y gl b -_g h sZ 5- \ k m h q

] h ] h e h ^ Z g b y g Z m j h \ _ g v r h d h \ h c b e _ l Z e . g h cd hl \ _ fyi _ lj_ f l m v Rh Z -Z i_jZl m jZ m ku luo hkh [_ c e_sZ 8, m lZ \beZ29.5 b m i_ey ^ b 31. khk ie hl \u 28.1° K e_lZevgZ y6, 31.7 b 29.2° K k hhl\_lkl \_ggh , 34. . M ]heh^ gu o ju[ l_jfh m k l h c q b \ h k l v :g r h_ h \ Z y l _ f i _ j Z l m j Z k 28.2, Z \ b7 Z 25.7° K e _ b` d h k l 23. e b ,lZevgZy 28.6 b 27.1° K m e_s Z l \u b i_ey ^b khhl \_lkl \_ggh 31.8, , ieh . < e _lgb c k_ahg ijb kdhjhk lb 8-10є K \Z m ^ k_ ]he_ ldh \ hdmgy gZ ]j_ q \h u qZevghc klZ^bb aZ[he_\Zgby kZijhe_]gbhahf \uy\e_gu [he__ DLF b EL b 35.7є K ih kjZ \g_gbx k h a ^ hjh \ufb (35.8 [ b 36.2є K M (34 ) hkh yfb ). k _ ] h e _ l d h \ d Z j i Z i j b a Z [ h e _ \ Z g b b d j Z k g m o h c ^ Z ` _ \ a h-g _ l l b f m f (ZH B L K g Z [ e x ^ Z e k y0% h l o h ^ L h e v d h j _ Z d p b y i h \ _ ^ _ g q _ 30є ) 10 . e b o h j Z ^ d b i j b \ h a f h ` g h k l b i _ j _ f _ s _ g b y \ a h g m l _ f i _ j-Z l m j f Z e v g h c i h a \ h e y e Z j u [ Z f i _ j _ ` b l v .a Z [ h e _ \ Z g b _ B g l _ j _ k gl h i j b l _ f i _ j Z l m j _ i \ jh_^ uu r Z x s _ c d j b l b q _ k d b c m j h \ ,q h , \ 30 q K \ i j _ k g h \ h ^ g u o \ h ^ h _ f Z o k j _ ^ g _ c i h e h k u J h k k b b g Z [ e x ^ Z , g b _ i b l Z g b y b j h, km Z _ j uv[ r _ g b _ \ b ^ h \ h ] h j Z a g ,h Z [ jlZ adb` _ k m s _ k l lf g hZy \_ggh_ mom^r_gb_ dZq_kl\Z \h^u b khdjZs_gb_ qbkeZ dhjfh Fhj^moZ c ;hel h\kdhc1975). < ^ bZiZahg_ l_fi_jZl mju38hl m , 1974, 30 ^h qK k _ j _ [ j y g h ] h d Z j Z k y j Z a \ b \ Z _ l k y ,l _ ihelh \ j u c r y \ d y _ l k y ] _ g- h l h d d hhc h e guf ihkdhevdm ijb\h^bl d m\_ebq_gbx qZklhlu ojhfhkhfguo , h ^ g h p _ i h q _ q g u f i h \ j _ ` ^ _ g b y f (AnitDa et al., 2000). B f _ g g h \ ^ b Z i Z a h g >Gh wlbo agZq_gbc l_fi_jZlmju jZkiheh`_gu mjh\gb DLF b O l_iehe x [b \uo \ b ^h \ ju [ . K d h j h k l v g Z ] j _ \ Z \ h ^ u \ e b y _ l g _ l h e v d h g Z l _ j f ,h g hk lbh c q Z \ h m gb g _ d h l h j u _ n b a b h e h ] b q _ k d b . FnZm g d p Z b v g Z y Z d l b \ g h k l v i b s _ dkbfb e l _ e v g u o d Z j [ h ] b ^ j Z a m d Z j , Z e b l duZ b Z h d m g y h l f _ q _ g Z i j b k d ii h \ j ky g Z ] j _ \ Z \ h ^ 0.04 q K , q i h a \ h e y x s _ c j u [ Z f m k i _ r g h Z d d e b f b j - \ Z l v u h \ u r _ g b x l _ f i _ j Z l m j u \ h \ k _. E _ a h f uh l f _ q _ g i h k e _ ^ h \ Z l _ e v g u c k l hg m j h \ g y n _ j f _ g l Z l b \ g h c Z d l b \ g h k l b k m \ _ e b q _ g b _ f k d4- j h k l h 50° K , q \ ^ j m ] b _ k _ a, hh k h [ _ g g h h k _ g v x ,b g a b e x ^ c _ l k y j _ a d-h _ k gu Z[ fh Z `_gb_ Zdlb \ghk lb dZj .[hM ^ jZa ^ b dZjiZ ]b fheh , hdmgy b ieh l \u \uy \e_gu f_g_gby Zdlb\ghklb Zp_lbeohebgwkl_jZau b kh^_j`Zgby n j Z d p b b [ _ e d Z \ , f hb ^ ] _ l _ e v k l \ m x s b _ h i h \ j _ ` ^ Z x s _ f ^ _ c k l \ b b k\ a _ dbo kdhjhkl_c gZ]j_ \Z \h^u . K h h l g h r _ g b _ w d h e h ] h n b a b h e h ] b q _ k d h ] ,h h d h hq Z il l_ e b fh m b f Z g vg a [bjZ_fhc b \_jog_c e_lZevghc l_fi_jZlmju m jZaguo ih wdheh]bb \b^h\ ju[ . : g Z e b a k h h l g h r _ g b y l _ f i _ j Z l m j u w d h e h ] h n b a b h e h ] b q _ k d h ] h h i l b f m f Z , h d h g q Z l _ e v g h b a [ b j Z _ f h c b \ _ j o g _ c e _ l Z e v g 49c \ b ^ _ f\ i f h Z l m j h l h _jj k d b o b i j _ k g h \ h ^ g u o j u [ \ u y \ b e l _ k g m x \ a Z b f h k \ y a v (Job-b o i wl ling, 1981). G Z f b g Z h k g h \ _ k h [ k l \ _ g g u o j _ a m e v l Z l h \ b ^ Z-g g u o ^h\Zl_e_c ling, 1981; :eZ[Zkl_j (Job , Eehc^1984; =heh \Zgh\ b., ^j , 1997) ij_^ijbgylZ ihiuldZ \uy\blv khhlghr_gb_ wdheh]h nbabheh]bq_k l _ f i _ j Z l m j g u o l j _ [ h \ Z g b c j u [ ? \ j h i _ c k d h c .q Z k l b J h k k b b

36 G Z i j b f _ j11 \ b ^ h \ j u [ m k l Z g h \ ,e q _ h ha g Z q _ g b y l _ f i _ j Z l m j u g _ lg

i j Z d l b q _ k d b k h \ i Z ^ Z x l k h i l b f Z e v g h c l _ f i _ j Z l mljZ [ c 9). f [ j b ( h .e w L Z [ e b p9. L _ f i _ j Z l m j g u _ o Z j Z d l _ j b k l b d b j u [ \ g Z q Z e _ h g l Z L_
H i lb f Z e vg Z y l_f i_jZ lmjZ j hklZ m k_] he _ldh\ b ] h^h\bd h\ k m (~ gZ 10° K \ur_ . A gZ q_ gby h ilbfZe vghc l_f i_jZlmju jhk lZ b H BL ) ijZdlb q_kdb kh \iZ ^Z xl q_ gby O EF \ur_ H 10-12єgZ m f heh ^ b l_ieh . AgZ BL K ex [b \uo \b ^h \ b gZ є K m oheh ^h ex [ b \uo \b ^ h.\ Ijb[ wlhf l_ fi_jZ lmj 12-15 ju \_j og_c ]jZg bp u ` bag_ ^_ y l_e vghkl b m b kke_ ^ h \Z ggu o \b ^h^h \Z jv 28 \ 41є K h ljZ `Zy jZagbp m \ l_jfh mklhc qb FZdkbfZ evgu_ a gZ q_gby , . \h klb j u [ i_jZlmj guo oZjZ dl_jbk lbd \uy \ e_g u hlhdZ jiZ , ahe m ]h dZjZky , g_kd he v b j hlZgZ dh gb `_ m fheh ^ b ^j m ] b o dZ jih \uo \b ^h \_s_ hdmgym s m db b kb [bjk , b gb `_ ]h hk _l jZ , fbgbfZ e vgu_ i_ey ^ nhj_ eb b gZe bfZ m ,b . G Z i j b f _ j _ f h e h ^ b j u [ j Z a e b q g u o w d h e h ] b q _ k d b o ] j, mlih m k qi agZ q_gby O EF \ur _, Z m dZj iZ b iehl \u \ur_ b agZ q_gb c EL DLF ( lZ [e 10). . GZb[he__ \ukhdb_ agZq_gby DLF b EL ijb e_lg_c l_fi_jZ [ueb m dZj iZ b, m mdb \u b hdmgy agZ q_gby D LF .[ueb -\ jZ \gu s iehl

37

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JZg__ [ueh ij_^eh`_gh g_kdhevdh \ZjbZglh\ jZa^_e_gby j l _ f i _ j Z l m j _ g _ j _j k l] b ,g1973), k h q _ l Z g b x l _ f i _ j Z l m j u g _ j _ k l Z b (> y Z \ u ` b \ Z g b y ( G b d Z g h, j h \ 1976) b e b i h l _ f i _ j Z l m j g u f g b r Z f h [ b (Magnuso n et al., 1979; Cherry, Ca irns, 1982). >ey jZa^_ e_gbyh[blZxsbo \ , ju[ i j _ k g h \ h ^ g u o \ h ^ h _ f Z o k _ \ _ j h a Z i ,Z iZ J h g h r _ g b x d l _ f i- _ j Z l ^ h hlkkbb g h f m n Z d l h j m g Z f b i j _ ^ e h ` _ g h b k i h e v a h \ Z g b _ lH _ o O h d Z a Z j B ,L i E F b l _ f i _ j Z l m j g h ] h ^ b Z i Z a h g Z ` b a g _. ^ _ y k _ e v g h k l bb b k w l b f f h < l hhl\_l \ \u^_eblv q_luj_ ]jmiiu ju[ : 1. G Z b [ h e _ _ l _ i e h e x [ b \Z j, i k _ j _ [ j y g u c b a h e h l h ,c j h l Z gAkgvZ : d u_ dZ.Z j q_gby
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< ijhp_kk_ w\hexpbhggh]h jZa\blby b hk\h_gby jZaghh[jZaguo wdheh]bq_ kdbo gbr m ju[ \ujZ[hlZehkv g_kdhevdh hkgh\guo nhjf l_f l Z p b b d l b \ g u c \ u [ h j h i j _ ^ _ e _ g g h c l _ f i _ j Z l m jkg\ y a Z a h g u k- h i l b .: , hc gghc f Z e v g h c k d h j h k l v x ,j Zbal\ b lgbbyy b ,j h j h ^ Z f h g k l j b j h \ Z g g Z i j b f _ i Z ikl _ [he__ 300 \ b ^ h \ j u [ j Z a g u o l Z d k h g h f b q _ k d b ob l] jZm isib o \ j Z a e b q g u o , h[ x ] _ h ] j Z n b q _ k d b o r b j( h l \ Z _,\ 1958; Fry, 1971; Jobling, 1981). L _ f i _ j Z l ,m j m Beo dhlhjmx ju[u ba[bjZxl \kdhj_ ihke_ ij_^hklZ\e_gby bf \ha [h ^ gh ]h \u [hjZ gZau \Z xl ba [bjZ_ fhc l_fi_jZl m j hc lhevdh - \ ^eb , . H ^gZdh l_evguo wdki_jbf_glZo kmlhd m^Z_lky hij_^_eblv hdhgqZl 2-3 \ u r _ [bjZ_fmx l_fi_j. lmjm Z K m s _ k l \ m _ l i j _ ^ i h e h ` q _ h bb_f _ g g h h d h g q Z l _ e v g h b a [ b j Z _ f Z y , lg l m j Z m j u [ k \ y a Z g Z k l _ f i _ j Z l m j h c \ h a f h ` g h ] h i j. hL Z, dg h -^ b g b b k o Z` _ [ h e _ _ \ u k h d b _ h d h g q Z l _ e v g h b a [ b j Z _ f u (~ 28-31°_K Z ljm jb j u [ _ l_fi j k _^ u , ) h [ b l Z x s b o \ \ h ^ h _ f Z o [ h j _ Z e v g h c a ,h g u f _ q h k k b b k Z d Z jgi Z hl J _gu m ( a Z ),Z k _ j _ [ j y g h ] h b h [ u d g h \ _ g g h ] h d Z j Z k _ c b ] h,e q h h_ rk h ] e jZ k m g Z k y l\ db hlZ _l j Z c h g h f i j h b k o h ` ^ _ g b _ f w l b o ] hb ^ < h k l h q g Z y ). a G y k e m q Z c g h (X \ h\ : b_ , \u [hj ba [bjZ_ fhc l_fi_jZl mju ijhbk oh ^bl ih qlb h ^bgZdh-\ h m eh^gu ,oa^hjh\u o b [ hevgu o bj u[ e vdh ih ^ hk lb`_gbb ahg u H BL , lh _ l k y j Z a g b p Z n b a b h e h ] h [ b h o b f b q _ k d h ] h b b f f m g h .e h ] b- q Z k d Ag _ q _ g b y H B L h l j Z ` Z x l l _ f i _ j Z l m j g, ud h lm k u _ \ b y h l \ _ l k l \ m x-l l _ f _ hj eh kh jZl m jguf ihlj_ [ghk lyf ju [ \ dhgdj_ lguc \ k_ahg e_gguc i_ jbh , h ij_ ^_ ]h ^Z h g l h ] _ g ,_ a jZ h i j _ ^ _ e _ g g h f n b a b h e h ] b q _ k d h f k h k l h y gD h e _ j ] Z ib . bb-h [ Z g b y H, Bg _ i j _ \ u r Z x s b -6 q K ih ^ l \_j ` ^ Z xl \Z `ghk lv Zk lZlb q ghk L 2_ , l_fi_jZlmjguo mkeh\bc kms_kl\h\Zgby ^ey hilbfbaZpbb jhk klb ib lZgby ( DhgklZg lbgh., 1991; A ^Zgh \b2001). <_jogb_ b g b `gb_ e \ b ^j ,q l Z e v g u _ l _ f i _ j Z l m j u o Z j Z d l _ j b a m x l l h l ^ b Z i Z a h g j h _ f-ih aj Z , \ dhlh l f \ _ f h ` g h k m s _ k l \ h \ Z g b _ \ b ^ Z \ j Z a e b q g u o m q Z kL d Z o f Z h _ Z, e Z f . l Zdb j[j ah h d h g q Z l _ e v g h b a [ b j Z _ f Z y l _ f i _ j Z l m j Z h l j Z ` Z _ l w \ h e- p b h x n b a b h e h ] b q _ k d b c h i l b f m f k m s _ k l \ ,h \ Z g e _ l Zb ^ v g Z y l _ f i j Z l m by \ eZ _ ]jZgbp m \hafh ` gh ]h h [ blZgby . K Z f h i j h b a \ h e v g u c \ u [ h j l _ f i _ j Z l m j u \ ^ e b l _ e v g u o , wj d- k i _ j ih o h ^ys bc \ ^ \Z wlZ i_j_oh ^g uc b wlZ i klZ [bevgh o ZjZd l_j_g ^ey iZ , ]h \u [ hjZ \k_o bk ke_ ^ h \Zggu o \b .^ hM ju [ - b o h eh ^hex [b \uo \b ^h \ ju [ mjh \_g \ l_ieh H B L b o Z j Z d l _ j i h \ _ ^ _ g b y \ ] j Z ^ b _ g l _ l _ f i _ j .Z Am Zu b l _ e v g Z y l gj q jZaebqg Z f i e b l m ^ Z d h e _ [ Z g b c i h d Z a Z l _ e y ^ Z ` _ \ i j _ ^ _ e Z o h ^ g h ] h- k _ f \ h \ Z g b _ \ g m l j b \ b ^ h \ u o j Z a e b q b c H B L m d Z a u \ Z x l g Z r b j h- d b c k h [ e _ g b c d k m s _ k l \ h \ Z g b x \ h i l b f Z e v g u o b i _ k k b f>Zbenvng_u o . j _ g p b j h \ d Z j u [ h ^ g h ] h \ b ^ Z g Z bjambi ji Z x s b _ j Z a g u _ l _ f i,_ j Z l ,][ u fh`_l m\_ebqb\Zlv ljhnbq_kdbc Zj_Ze b kihkh[kl\h\Zlv - \_e m ghklb ihi m eyp bc . G Z b [ h e _ _ \ Z ` g u f b n Z d, l h ij jZ f^ _ e y x s b f b i h \ _ ^ _ g b _ b j Z k i j _ ^ _ e h _b j u [ \ l _ j f h ] j Z ^ b _ g l g u o m k eyh\ ebyyxol k y \ h a j Z k l b k _ aK gi h] h ^ \ -e _ , \ .h yZ gb_f \hafh`ghklb hkms_kl\eylv gZijZ\e_ggh_ ^\b`_gb_ -ebqb

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40 \ b y o k \ h _ ] h j h ^ Z d h f i j h f b k k l _ f i _ j Z l m j g u o l j _ [ h \ Z g b c j u [ b

^ j m ] b o n Z d l , jl h \ d b o d Z d k h ^ _ j ` Z g b _ d b k e h jghZZ b\ q b\ h ^ _b ^ h-k l m i hZ , ^e _ g h k l v d h j] f Zj h ^ b g Z f b q _ k d b _ m ,k e jhb\ b y l k l \ b _ o b s ,g i Z jhZ a b-l Z j , b^ i km bd \ g u o b ^ j m ] b o a Z [ h e. _ I\ Z e h ` c l _ e v g h _ ^ _ c k l \ b _ \ u k h d h c l _ f i _ j Z hgb b j h k l j u [ \ a h g Z o i h ^ h ] j _ l u o \ h ^ \ j y ^ _ k e m q Z _ \ f h ` _ l g b \ _-e b j h lb \guf \ebygb_f ^j m ]b o n Zdlhjh \ . > e y \ u y k g _ g b y \ h i j h k Z h k l _ i _ g b \ e b y g b y d j Z l d h k j- q g u o h g u o b a f _ g _ g b c d, eZ f lZ l d ` _ k m s _ k l \ m x s b o a h g l _ j f Z e v g h ] h a b ZZ g Z i j h p _ k k u l _ f i _ j Z l m j g u o, Z _Z i[ l Z pbbf c i h k l h y g g u c f h g b l h g ^ h oh^ l_jfhj_]meypbhggh]h ih\_^_gby b l_jfhmklhcqb\hklb g_ lhev \ b ^ h \ , g h b \ b ^ h \ \ k _ e _. g < _ j\ h y l, grhb j h d h _ j Z k i j h k l j Z g _ g b _ - ] h e h p_ jhlZgZ b lxev db ella cultriventris, f h ` _ l [ u l v k \ y a Z g h k b a f _ g _ g b _ f Clupeon jy^Z n Zdlhjh\ , \ lhf qbke_ b l_fi_jZl m j u \h^ghc kj_^ u . L _ j f h j _ ] m e y p b h g g h _ i h \ _ ^ _ g b _ o Z j Z d l _ j g h ^ e y i j _ ^ k l -Z \ b l _ k h g ,h \ h l i j h k l _ c r b o ^ h f e _ d h i b l K Z s b o j h b a \ h e v g u c \ u [ h j -h i j _ . Zxfhi e _ g g h c l _ f i _ j Z l m j u \ u y \ e _ g m j u [ j Z a g u o l Z d k h ghh[ b l Z_xk d b , fbq s b o g Z j Z a e b q g u o ] _ h ] j Z nr bh q l_Z o d b o\ Z g l Z j d l b q _ k d b o b Z j d l b q _ bj k , m f _ j _ g g u o b l j h i b q _ k d. b J _ Z h g Z y l _ j f h b a [ b j Z g b y i j h y \ e y _ l k o adpb o j Z a g u o w d h e h ] b q _ k d , o Z ] a g h ] ih \ h a j Zh ll Z b q b g d b ^ h \ a j h k e- h c h b j jmi (k e [ b ), \ h \ k _ k _ a h g u. H q ^ _ \ b ^ g , l _ j f h j _ ] m e y p b h g g h _ i h \ _ ^ _ g b _ y \ ]h Z h \jh`^_gghc j_Zdpb_c b fh`_l [ulv hlg_k_gh d gZb[he__ h[s Z ^Z ilZpbyf \h ^gu o `b \ hlguo . L _ j f h j _ ] m e y p b h g g h _ i h \ _ ^ _ gkbe_h ` j u [ c i j h p, _\ d e x q Z x s b c l -_ f gu kk i_jZl m jg mx Z ddebfZpb x h ^ys bc gZ nhg_ ^j m ] b o n hjf ih-\_ ^ _g , ijh bk o bk k e _ ^ h \ Z l _ e v k d h ] jhj b l h j b Z e v g h ] h b i b s _ \ h ] h . h \ _ ^ _ gub, y\ g m l j-b , l_ i

41

3. A g Z q _ g b y h d h g q Z l _ e v g h b a [ b j Z _ f h c l _ f i _ j Z l m j u - \ h g ex [b \u o b o heh ^he x [b \uo \b ^h \ baf _gyxlky [ ghkl v d l_ jfhba [ b . Kihkh g b x i h y \ e y _ l k y g Z e b q b g h q g u o w l Z i H o h j ZZ l\ b leb y h b a [ b j Z _ . Zd gq a _ vg l_fi_jZlmjZ ^hklb]Z_l fZdkbfZevguo agZq_gbc m k_]he_ldh , b \ ^ Zevg_cr_f x \_g bevgu o b iheh \h aj_eu o hkh [ _c 2-14° K ,m k gb `Zxlky . 4. < u y \ e _ g u k m l h q g u _ b k _ a h g g u _ b a f _ g _ g b ,y o lZ jjZ h b _ j[ b b Z _ fdl a j \ujZ`_gghklv dhlhjuo kms_kl\_ggh jZaebqZxlky m ij_kgh\h j u [ j Z a g h ] h \ b ^ Z b \ h .a M kkllZ g h \ e _ g_u m j h \ g y l _ j f h b a [ b j Z g g b c jZ Z 4: e_l by (18-28є K i_j_ o h^guc -16є K khhl \_lkl\ mxsbc l_fi_jZ lmjghf m i ), (11 ), jhklZ b jZa \blby ju [ , \_k_gg_ hk_(5-8є K b abf gbc K ggbc ) (1-3є ). 5. J Z k i j _ ^ _ e _ g b _ b l _ j f h j _ ] m e y p b h g g h _ i h \ _ ^ _ g b _ j-u [ a Z ] _ g g u o b w g ^ h ] _ g g u o n Zl d lfh j h \ Z l m j u Z d d e,b f Z p b bb y d h j f Z : _ i_j gZebq , k l _ i _ g b i h e h \ h a j ,_ a Z khleb_ \ Z ,g b c c j h i _ i l b (h[ h f [ _ a b). M \ _ e b q _ g b _ eh[ g_ ^\ g ^ b Z i Z a h g Z b a [ b j Z _ f u o l _ f i 2-14є l m j j b Zh l k m l k l \ b b d h j f Z i j b \ h _jZK i g j Z k r b j _ g b x h [ e Z k l b h [ b l Z g b y f hu [ a [ b j Z g b _ j _ Z e b a m _ l k y g Z . L_j j b ]bo nhjf ih \_ ^_gby bkke_ ^ h \Zl_e vkdh ] h , l_jjblhjbZe vgh ]h b ibs _ \ h ]h . 6 < _ j o g y y e _ l Z e v g Z y l _ f io_Z ZZ mlj_ j b a m x s Z y a h g m w , jj ld Z n b a b h e h ] b q _ k d h ] h i _ k km f m f Z Z e v g u o i j _ k g h \ h ^ g u o j u [ g Z o h , b [hj_ ^ b Z i Z a h g _28h ^ lh 41є K H g Z i h \ u r Z _ l k y \ e b q b g h q g h f , i^ h k b b ] ^ Z _ l . _jl h _ f Z d k b f m f Z \ f Z e v d h \ h f b k g b ` Z _ l k y m g _ i h e h \ h a j _ e u o. b i GZb[hevrb_ agZq_gby ojhgbq_kdh]h e_lZevgh]h fZdkbfmfZ l _ j f b q _ k d h ] h f Z d k b f m f Z h l f _ q _ g u m f h e ,h fb gj b f Zee_vl gu a b ^ b u[ h f _fhc =heh^Zgb_ b aZ[he_\Zgby ihgb`Zxl l_fi_jZlmjgmx, mklh . Z i h \ u r _ g g Z y k d h j h k l v g Z ] j _ \ Z \ h ^ u b a f _ g y _ l g _ l h e v d , glh j f h h_ b nbabheh]h [bhobfbq_kdbc klZlmk hj]ZgbafZ . 7. F _ ` ^ m h i l b f Z e v g h c l _ f i _ j Z l m, j h c hjg q Z l Z e v g h b a [ b j Z _ f h d hk _ \_jog_c e_lZevghc l_fi_jZlmjhc ju[ kms_kl\m_l hij_^_e_gg . L _ f i _ j Z l m j g u c h i l b f m f f h e h ^ b g\ u r , q _ f m \ a j h k e u o Z u a -g Z 3-10° K Z _ ,j[ q_gb_ \_j o g_c e_ lZevghc l_fi_jZl mju m 8-17° K \ur_]b hi-lb l_ o b ^j m o gZ f Z e v g u o a g Z.q _ G b c h k g h \ Z g b b h d h g q Z l _ e v g h b a [ b j Z _ f h c l gZ , ojhgbq_kdh]h e_lZevgh]h fZdkbfmfZ b l_fi_jZlmjgh]h ^bZ l_evghklb ij_kgh\h^gu_ ju[u k_\_jh aZiZ^Z Jhkkbb jZa^_ey ]jmiiu ih hlghr_g bx d l_fi_ jZ lmjghf m n Zdlhj m . 8. A Z d h g h f _ j g h k l b l _ j f h b a [ b j Z g b y m j u [ i j h y \ e y x l k y \ e Z [ h j Z l h j g u o b i j b j h ^ g u o m k e h \ b y oZ c h g _ l _ j f Z e v g h ] h a Z ] j y a g _ g b y k m s _ k .< j lgZy ^bn n_j_ gpbZpby i hi m eypb c l_i eh ex [b \u o ,\b ^ h \ \b db b . K_ ]he _ldb^h ]h g _ i h e h \ h a j _ e u _ h k h [ b i j b \ e _ d Z x l k y \ j Z c h g u k [ j h k Z l _,i e u o \ a j h k e u _ j u [ u b k i h e v a m x l w l b j Z c h g u ^ e y [ h e _ . I j Z g gq _ gh g _ hem _] g u _ ^ Z g g u _ i h a \ h e y x l i j h ] g h a b j h \ Z l v i h \ _ ^ _ g b _ b j Z k i j -_ ^ _ e _ f h ] j Z ^ b _ g l g u o m k e h \ b y Z d ` _ h p _ g b l v w d h e h ] b q _ k d b c j b -k d \ , Z lo fZevgu o k [jhk h \ ZghfZe vguo i h ]h ^guo y \ e_gby o b ^he ]h \j_f _ , ijb f _ g _ g b y o d e.b f Z l Z

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1. E Z i d b g < < ., K \bj kdbc :., Fh e h \Zg h \ < . Dh \ .gu o .J . K 240243. . 3. K \ b j k d b c : F ., =heh \Z gh \ < . D

43

14. = h e h \ Z g h \ Z B E., = heh \Z gh\ < . D< e b y g b _ Z [ b h l b q _ k d bo n Z dl h j h \ ( l_f i_jZlmjZ , ly `_eu_ f_ lZeeu Zdl b \ghk lv dZj [ h ]b ^ jZa h [t_d lh \ i b , jG ) gZ b o lbhn Z]// K ., =heh\Z gh\Z B KjZ \gbl_e vguc ZgZe ba h .K qZl_ev gh ba [bjZ_f hc b \_jog_c e_lZ evgh c l_f i_jZ lmju m- fheh ^h \ ij_ kgh \h ^g u o j // [ Ljm ^ u DZj_e J: G K_j Wdki_ jbf eh ]by 012. u . GP . . . [bh .2 < 3. ( \ i_ q Zlb ) Im [ebdZp bb \ ^j m]bo ba^ : Zgbyo

21. I h ^ ^ m [ g u c : = ., =h e h \ Zgh \ <., DZ i d b g < < E . I h \_ ^_g b_ ju [ \ l_jfh ]jZ^b_g lgu o m ke h\by o \ aZ\bkbf hklb hl ;bhe \g mlj \h^ : B gn hjf // k _ahgZ . ]h^Z . . [xe . E., 1976. < 30. K 4143. . 22. = h e h \Z gh\ < D ., ;ZaZjh \ F a[_]Z_fu_ l_f i_jZ lm ju m fheh^b // ;b he \ gmlj \ h^ : Bg nhj. f[xe . E., 1981. < 59. K 4547. . . . 24. = h e h \Z gh\ < D . L _ j f h j _ ] m e yl h j g h _ i h \ _ ^ _ g b _ ] b ^ j h [ b h gl h \ // ;bheh ]by \gmlj \h^ : Bgnhjf [x e . E., 1987. < 73. K 5660. . . . 25. I h ^ ^ m [ g u c K :., = h e h \Zg h \ < .,D ;Z aZjh\ F B ., Dm^j ydh\ .K < < ebygb_ l_jfh ] b ^jh ^ bgZf b q_k dbo m ke h \bc \ ah g_ k [jhkgu o jZk oh ^h \ D hkl gZ jZ kij_ ^ _ e_gb_ Wg_j k ljhbl_ evk1995. < 6. K 3841. // j u [ . ] . l \h . 26. = h e h \Z gh\ < D . L_f i_jZ lmjgu_ ghjf u `bag_^_y l_ev ghk lb ij_kg h \h^gu o ju [ // Jb[g_ ]hkih ^Zjkl\h ). 2009. K ., = h e h \Zg h \ < . D< _ jo g b _ e _l Z ev g u _ l _ f i _ j Zl m j u b l _ j f h ij_n_j_ g ^ m f fh eh ^b s mdb ius L. // Jb [ g_ ]hk ih ^Zjk ( Db_ \ 2009. Esox luc l \h ). < ui 66. K 6570. . . 28. I h ^ ^ m [ g u c : = ., = h e h\Zg h\ < .,DEZi dbg .< K_ ahg gZy ^b gZfb < dZ b jZ_fu o l_fi_jZ lmj L_hj_ lb q_kd b_ Zki_d lu ju [ h o hay ckl \_ -gguo // ju [ \Zgbc \h ^ h ojZgb: ebs B;< < : G KKK., 1978.

44

29. E Z i d b g < < ., = heh \Z gh \ < .,DK \bjk dbc :., F Khdh eh \ . L_jfhZ ^ Z <: ilZ pbhggu_ o Z jZdl_j bklbd b e_sZ (L) Ju [ bgk dh ]h \h ^h ojZg beb sZ Abramis brama // Klj mdl m jZ e hdZe vghc i hi meypb b m ij_k [ gh \h. d u 0. K 3785. . Ju bgk ^g o ju [ 199 . 30. = h e h \ Z g h \ < D ., hd e 2jhk kh\_s « Ih \_^_gb_ j ». ;hjh .d 6. K 1640. . . u[ 199 . 33. F Z e b g b g E D ., ;Z aZjh \ F B e h \Zgh \ < ., DE b g g b d < > ., =h . ju [ . 2- ] h hf . Ju[b gkd : « i_qZ lb 2005. K 5981. », . 39. K f b j g h \ : D ., = h e h \Zg h \ < .,D K\bjkd bc : F . <_jogb_ e_ lZevgu_ l_ jZlmju e_sZ \u b h dmgy \ jZae b qgu_ i_j bh ^u /hglh ]_g_aZ , iehl / : dl m Ze vgu_ ijh [e_f u wdhe h ]b q_kd hc nba bhe o bf bb b ]_g_ lbdb ` b . FZl_. j , [ bh h ]bb \hl gu o F_ ` ^ m gZj gZ m. qg . dhgnKZjZ gkd ^ \h Fhj ^ h \ mg ,lZ 05. K 224228. . : Ba . 20 . 40. = h e h \Z gh\ < D . L _ f i _ j Z l m j g u c h il b f m f b l _ f i _ j Zl m j g u _ ] j Z g b p u ` ba g_^_y l_evg hklb hk_ ljh\u o \b^h:d\Zd mevl mjZ hk_ ljh\u o ju[ // « \ ju[ : >hk lb`_ gby b i_j ki_d lb \ u jZaIV F_ ` ^ mgZj G Z m iq jZdl dhgn : k ljZoZgFZ ». \blby . .. ., . vl_ .j ^hd e F .: B a^ \h

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