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Знакомство с эволюционной генетикой (Гаевский)

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gZ jbk Djb\u_ jZaebqZxlky bkoh^gufb agZq_gbyfb qZklhlu Zee_ey Z.

Fh^_ev hl[hjZ ]_ghlbih\ gZ mjh\g_ h^gh]h ehdmkZ jZkiheh-

`_ggh]h \ O o jhfhkhf_

<hkihevam_fky ih^oh^hf dhlhjuc [ue ijbf_g_g ijb fh^_ebjh\Zgbb hl[hjZ gZ mjh\g_ h^gh]h Zmlhkhfgh]h ehdmkZ Hlebqb_ fh^_eb hl[hjZ gZ mjh\g_ h^gh]h ehdmkZ jZkiheh`_ggh]h \ O ojhfhkhf_ aZdexqZ_lky ij_`^_ \k_]h \ lhf qlh ihimeypby jZa-

^_e_gZ gZ ^\_ km[ihimeypbb kZfph\ b kZfhd DZ`^Zy bf__l k\hb qZklhlu ]_ghlbih\ kh k\hbfb dhwnnbpb_glZfb ijbkihkh[e_gghklb

41

 

 

KZfdb

 

 

KZfpu

 

 

 

 

 

 

 

 

=_ghlbiu

::

:Z

ZZ

KmffZ

AY

aY

Kmf

 

 

 

 

 

 

 

fZ

 

 

 

 

 

 

 

 

QZklhlu

x

y

z

1

g

j

1

 

 

 

 

 

 

 

 

Ijbkihkh[e_gghklv

W0

W1

W2

-

W3

W4

 

 

 

 

 

 

 

 

 

Hlghkbl. \deZ^

xW

yW1

zW2

<1

gW

jW4

<1

 

0

 

 

 

3

 

 

 

 

 

 

 

 

 

 

 

D fhf_glm hieh^hl\hj_gby qZklhlu ]_ghlbih\ m kZfhdIb kZfph\PkhklZ\yl

 

pAA

=

 

 

 

xW0

;

 

 

pAY

=

 

 

 

 

gW3

 

 

;

 

 

 

 

 

 

 

xW0

+ yW1 + zW2

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

gW3 + jW4

 

pAa

=

 

 

 

yW0

;

 

 

paY

=

 

 

 

jW4

 

;

 

 

 

 

 

 

 

xW0

+ yW1 + zW2

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

gW3 + jW4

 

paa

=

 

 

 

zW0

 

;

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

xW0

+ yW1 + zW2

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

QZklhlu khhl\_lkl\mxsbo Zee_e_c \ km[ihimeypbyo jh^bl_e_c

 

pA( f ) = pAA + 0.5pAa

 

 

pA( m) = pAY

 

 

 

 

 

 

 

 

 

 

 

 

 

pa( f ) = paa + 0.5pAa

 

 

pa( m) = paY

 

 

 

 

 

 

 

 

 

 

 

 

 

 

QZklhlu ]_ghlbih\ h[jZah\Z\rboky ihke_ hieh^hl\hj_gby ab]hl k mq_lhf ihem-

q_gguo \ujZ`_gbc jZ\gu

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

p (1)

= p ( f ) p (m) =

 

 

 

 

(xW0

+ 0.5 yW1 ) • gW3

 

(58)

 

 

 

 

 

 

 

 

 

 

 

 

+ yW1 + zW2 )(gW3 + jW4 )

 

 

 

 

 

 

 

AA

A

A

 

(xW0

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

pAa(1) = p A( f ) pa(m) + pa( f ) pA(m) =

 

 

 

 

 

 

 

 

=

(xW0

+ 0.5 yW1 ) jW4

+ (zW2

+ 0.5 yW1 )gW3

 

(59)

 

 

 

 

 

 

 

 

(xW0 + yW1 + zW2 )(gW3 + jW4 )

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

p (1)

= p ( f )

p

( m)

=

 

 

 

(zW2 + 0.5 yW1 ) • jW4

 

(60)

 

 

 

 

 

 

 

 

a

 

(xW0 + yW1 + zW2 )(gW3 + jW4 )

 

 

 

 

 

 

 

 

aa

a

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

p (1)

 

= p

( f )

=

 

 

xW0 + 0.5yW1

 

(61)

 

 

 

 

 

 

 

 

 

 

 

 

AY

 

 

 

 

A

 

 

xW0 + yW1 + zW2

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

p (1)

= p

( f )

=

 

 

 

zW2 + 0.5W1

 

(62)

 

 

 

 

 

 

 

 

 

 

 

 

xW0 + yW2 + zW2

 

 

 

 

 

 

 

 

 

 

 

 

aY

 

 

 

a

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

42

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

(66 )

>_ckl\b_ hl[hjZ dZd b ^ey fh^_eb Zmlhkhfgh]h ehdmkZ fh`gh ijhke_^blv gZ ijhly`_gbb jy^Z ihdhe_gbc hp_gb\Zy ^ey dZ`^h]h ihdhe_gby qZklhlu ]_ghlbih\ \ fhf_gl h[jZah\Zgby ab]hl eb[h hij_^_eblv baf_g_gb_ qZklhl Zee_e_c : b Z aZ h^bg wlZi hl[hjZ \ km[ihimeypbyo kZfhd b kZfph\ b \ ihimeypbb \ p_ehf

GZijbf_j baf_g_gb_ qZklhlu Zee_ey Z ^ey jZkkfhlj_ggh]h \ur_ kemqZy fh`gh hij_^_eblv ba \ujZ`_gbc

^ey kZfhd ∆ pa( f ) = paa(1) + 0.5pAa(1) (z + 0.5y)

(63)

^ey kZfph\ ∆ pa( m) = paY(1) j

 

 

 

 

(64)

^ey ihimeypbb

 

 

 

 

 

pa( f + m) =

2

( paa(1) + 0.5pAa(1)

z 0.5y) +

1

 

( paY(1) j)

(65)

 

 

3

3

 

 

Ijb g_h[oh^bfhklb ^Zggu_ \ujZ`_gby fh]ml [ulv ijb\_^_gu d \b^m

p = f (x, y, z, j, m,W0 ,W1 ,W2 ,W3 ,W4 )

Mqblu\Zy ]jhfha^dhklv ihemqZ_fuo \ujZ`_gbc ^ey bo h[jZ[hldb `_eZl_evgh bkihev- ah\Zlv j_kmjku i_jkhgZevguo dhfivxl_jh\

= _g _l b q_kd b c ] j ma

=_g_lbq_kdbc ]jma ieZlZ aZ hl[hj ij_^klZ\ey_l kh[hc g_ba[_`gu_ ihl_jb ihim- eypbb \ua\Zggu_ jZaebqbyfb \ Z^ZilZpbhgghc p_gghklb __ n_ghlbih\ ]_ghlbih\

Ihl_jb fh]ml \ujZ`Zlvky \ nbabq_kdhc ]b[_eb f_g__ ijbkihkh[e_gguo hkh[_c beb \ ebr_gbb bo \hafh`ghklb hklZ\blv ihlhfkl\h ]_g_lbq_kdZy ]b[_ev

Hp_gdm ]_g_lbq_kdh]h ]jmaZ == ^_ckl\mxs_]h gZ h^gh ihdhe_gb_ fh`gh ^Zlv gZ hk- gh\_ ke_^mxs_]h \ujZ`_gby Ws ]^_:VihdZau\Z_l kj_^gxx ijbkihkh[e_gghklv ihimeypbb Bkihevamy dhwnnbpb_glu hl[hjZ b jZ\gh\_kgu_ qZklhlu ]_ghlbih\ ihemqZ_f

== p2 s0 + 2 pqs1 + q 2 s2

<_ebqbgZ ]_g_lbq_kdh]h ]jmaZ aZ\bkbl hl mkeh\bc hl[hjZ b baf_gy_lky \ jy^m ihdhe_- gbc gZ dhlhju_ ^_ckl\m_l hl[hj GZ jbk ihdZaZgu \ZjbZglu ijhy\e_gby ]_g_lbq_- kdh]h ]jmaZ ijb hl[hj_ ijhlb\ ^hfbgZglguo n_ghlbih\ JZkq_lu \uiheg_gu gZ hkgh-

\Zgbb mkeh\by qlh ^h gZqZeZ ^_ckl\by hl[hjZ qZklhlu ]_ghlbih\ \ ihimeypbb [ueb jZ\gu ::S2=0.36; :ZST ZZT2 <b^gh qlh k ihgb`_gb_f ijbkihkh[- e_gghklb ]_ghlbih\ :: b :Z m\_ebq_gb_f bgl_gkb\ghklb hl[hjZ \hajZklZ_l gZqZev-

43

gZy \_ebqbgZ ]_g_lbq_kdh]h ]jmaZ h^gZdh mf_gvrZ_lky qbkeh ihdhe_gbc g_h[oh^bfuo ^ey aZ\_jr_gby hl[hjZ

 

 

 

 

 

 

 

:

:

:

:

 

 

 

 

 

 

 

 

 

 

 

: :

 

]jma

 

 

 

 

 

 

 

 

 

 

kdbc

 

 

 

 

 

lbqg

 

 

 

 

 

=_

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Ihdhe_gb_

Jbk >bgZfbdb ]_g_lbq_kdh]h ]jmaZ \ g_kdhevdbo ihdhe_gbyo \h \j_fy hl[hjZ ijhlb\ ^hfbgZglguo n_ghlbih\ ijb jZaebqguo mjh\gyo ijbkihkh[e_gghklb

Ijb hl[hj_ \ ihevam ]_l_jhab]hl kemqZc k\_jo^hfbgbjh\Zgby ihimeypby fh`_l gZoh^blvky \ khklhygbb mklhcqb\h]h jZ\gh\_kby JZ\gh\_kgu_ qZklhlu hij_^_eyxlky khhlghr_gb_f dhwnnbpb_glh\ hl[hjZV0 bV2):

p =

 

s

2

; q =

 

s0

 

.

 

 

 

 

 

s0 + s2

s0

+ s2

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

=_g_lbq_kdbc ]jma jZ\_g

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

s2

 

2

 

 

 

s0

2

 

 

s0 s2

 

==

 

 

 

 

 

s0

 

 

 

 

s2

=

 

(67)

 

 

 

 

 

 

 

 

+ s

 

 

+

+ s2

 

s0 + s2

 

s0

2

 

 

s0

 

 

 

 

>ey kjZ\g_gby jZaebqguo \ZjbZglh\ hl[hjZ djhf_ \_ebqbgu ]_g_lbq_kdh]h ]jmaZ fh`gh bkihevah\Zlv lZdhc iZjZf_lj dZd ieZlZ aZ hl[hj dhlhjuc ij_^klZ\ey_l ]_g_lb- q_kdmx ]b[_ev hkh[_c gZ ijhly`_gbb \k_o ihdhe_gbc ihdZ ^_ckl\m_l gZijZ\e_gguc

hl[hj

 

 

IH 3

(pi2 s0 + 2 pi qi s1 + qi2 s2 )

(68)

 

 

 

< ij_^klZ\e_gguo gZ jbk \ZjbZglZo h[sZy ieZlZ aZ hl[hj khklZ\beZ ijb

: ijb : b ijb : H[jZsZ_l gZ k_[y \gbfZgb_ lh qlh kgb`_gb_ bgl_gkb\ghklb hl[hjZ mf_gvrZ_l ]_g_lbq_kdbc ]jma gZ h^gh ihdhe_gb_ jbk

gh m\_ebqb\Z_l h[smx ieZlm aZ hl[hj

44

q(1- s)/(1 - qs).

Q Z k l h l g h a Z \ b k b f u c h l [ h j

>ey jZkkfhlj_gguo \ur_ \ZjbZglh\ hl[hjZ h[sbf y\ey_lky lh qlh Z^ZilZpbhg- gZy p_gghklv ]_ghlbih\ Z ke_^h\Zl_evgh b bgl_gkb\ghklv hl[hjZ hklZxlky ihklhyg- gufb Wlh kms_kl\_ggh mijhsZ_l fh^_ev hl[hjZ gh g_ iha\hey_l ijb[ebablvky d hib- kZgbx j_Zevghc kblmZpbb dh]^Z bgl_gkb\ghklv b agZd hl[hjZ f_gyxlky \ khhl\_lkl\bb k qZklhlhc hl[bjZ_fh]h ]_ghlbiZ < dZq_kl\_ \hafh`gh]h \ZjbZglZ fh`gh ij_^iheh-

`blv qlh Z^ZilZpbhggZy p_gghklv ]_ghlbiZ ihgb`Z_lky k jhklhf qZklhlu \klj_qZ_fh- klb \ ihimeypbb LZdZy kblmZpby kha^Z_l \ ihimeypbb [eZ]hijbylgmx h[klZgh\dm ^ey j_^dbo ]_ghlbih\

KhihklZ\blv h[uqguc b qZklhlgh aZ\bkbfuc \ZjbZglu hl[hjZ fh`gh \ jZfdZo m`_ bkihevah\Zgghc h^ghehdmkghc fh^_eb MjZ\g_gb_ hibku\Zxs__ baf_g_gb_ qZklh- lu Zee_ey gZijbf_j Zee_ey Z aZ h^gh ihdhe_gb_ ijb ^_ckl\bb qZklhlgh aZ\bkbfh]h hl[hjZ fh`gh ihemqblv ba mjZ\g_gby aZf_gb\ nbdkbjh\Zggu_ agZq_gby dhwnnb- pb_glh\ hl[hjZV0 , s1 bV2 gZ nmgdpbhgZevgh aZ\bkysb_ hl qZklhlu ]_ghlbiZV0 = f(p2), s1 = I STbV2 = f(q2):

∆ q = pq{q[f(2pq)-f(q2)]+p[f(p2)-f(2pq)]}/Ws,,

(69)

]^_ Ws = 1 - [p2•f(p2)+2pq•f(2rq)+q2•f(q2)]

(70)

 

>ey ijbf_jZ fh`gh jZkkfhlj_lv kemqZc dh]^Z dhwnnbpb_glu hl[hjZ ijyfh ijh- ihjpbhgZevgu qZklhlZf V0=p2, s1=2pq, s2=q2 dhwnnbpb_gl ijhihjpbhgZevghklb \ayl jZ\guf < wlhf kemqZ_ ijbT gZklmiZ_l mklhcqb\h_ jZ\gh\_kb_

< ijbjh^_ h^gbf ba kemqZ_\ qZklhlgh aZ\bkbfh]h hl[hjZ fh`_l [ulv baf_g_gb_ bgl_gkb\ghklb ih_^Zgby ilbpZfb kt_^h[guo gZk_dhfuo ih^jZ`Zxsbo k\hbfb \g_r- gbfb ijbagZdZfb y^h\bluf gZk_dhfuf ^jm]h]h \b^Z < h[s_f kemqZ_ qZklhlgh aZ\bkbfuc hl[hj fh`_l [ulv khklZ\guf we_f_glhf klZ[bebabjmxs_]h hl[hjZ

H l [ h j ] Z f _ l

Fh`_l hdZaZlvky qlh ]Zf_lu h[eZ^Zxl ^bnn_j_gpbZevghc kihkh[ghklvx d \u-

`b\Zgbx beb dZdbfb lh ^jm]bfb jZaebqbyfb \ebyxsbfb gZ bo kihkh[ghklv mqZkl\h-

\Zlv \ hieh^hl\hj_gbb < wlhf kemqZ_ gZqbgZ_l ^_ckl\h\Zlv _kl_kl\_gguc hl[hj gZ mjh\g_ ]Zf_l JZkkfhljbf ke_^mxsbc ijbf_j \ dhlhjhf ihimeypby ih hij_^_e_gghfm ehdmkm h[jZam_l ]Zf_lu Z1 b Z2 k qZklhlZfbSbTImklv Z1 emqr_ ijbkihkh[e_gu q_f Z2 Lh]^Z bo Z^ZilZpbhggZy p_gghklv fh`_l [ulv ijbgylZ aZ _^bgbpm Ijbkihkh[e_g- ghklv Z2 f_gvr_ _^bgbpu gZ \_ebqbgmVhij_^_e_ggmx dZd dhwnnbpb_gl hl[hjZ Wn- n_dlb\guc mqZkl\mxsbc \ jZafgh`_gbb nhg^ ]Zf_l ihke_ hl[hjZ hij_^_ey_lky ijh-

45

ba\_^_gb_f qZklhlu ]Zf_lu gZ \_ebqbgm hlghkbl_evghc Z^ZilZpbhgghc p_gghklb Z1)

p•1 Z2) q•(1 - s)

H[sbc nhg^ ]Zf_l hij_^_ey_lky dZd kmffZ

p + q( 1 - s ) = p + q - qs = 1 - qs

Hlghkbl_evgu_ ^heb qZklhlu ]Zf_l ihke_ hl[hjZ jZ\gu khhl\_lkl\_ggh

^ey Z1 p/(1 - TV^ey Z2

J_amevlZl hl[hjZ ij_^klZ\ey_l kh[hc baf_g_gb_ qZklhl jZkkfZljb\Z_fuo ]Zf_l Baf_g_gb_ qZklhlu ]Zf_lu Z2 aZ h^bg wlZi hl[hjZ fh`gh hij_^_eblv ke_^mxsbf h[jZ- ahf

q1 - qo = q(1- s)/(1- qs) - q = - sq(1- q)/(1- qs) (71)

bebT1 - qo = - pqs/(1 - qs)

(72)

Ihkdhevdm qZklhlu ]Zf_l \ kmff_ ^Zxl _^bgbpm baf_g_gb_ qZklhlu ]Zf_luD2 \ i_j\hf ihdhe_gbb [m^_l lZdbf `_ dZd ^ey ]Zf_lu Z1 lhevdh k ijhlb\hiheh`guf agZ- dhf

Kh\f_klgh_ ^_ckl\b_ w\hexpbhgguo nZdlhjh\

JZkkfhlj_ggu_ \ur_ ijhp_kku kihkh[gu_ baf_gblv ]_g_lbq_kdbc khklZ\ ihim- eypbb b \ua\Zlv w\hexpbhggu_ baf_g_gby \ _kl_kl\_gguo mkeh\byo qj_a\uqZcgh j_^- dh ^_ckl\mxl \ qbklhf \b^_ GZ ijhly`_gbb h^gh]h ihdhe_gby ihimeypby ih^\_j]Z_lky ^_ckl\bx fmlZpbhggh]h ijhp_kkZ ^j_cnZ ]_gh\ _kl_kl\_ggh]h hl[hjZ b fb]jZpbb >bnn_j_gpbjh\Zlv baf_g_gby ih ijbjh^_ \uau\Zxsbo bo ijhp_kkh\ ljm^gZy b qZklh g_jZaj_rbfZy aZ^ZqZ H^bg ba ih^oh^h\ d j_r_gbx wlhc ijh[e_fu aZdexqZ_lky \ hi- j_^_e_gbb mkeh\bc ijb dhlhjuo j_rZxsbf hdZau\Z_lky h^bg beb \ djZcg_f kemqZ_ ^\Z w\hexpbhgguo nZdlhjZ

< jZ[hlZo :cZeZ> @ =jZglZ> @ ijb\_^_gu khhlghr_gby iha\heyxsb_ hp_gblv kh\f_klgh_ ^_ckl\b_ ^j_cnZ ]_gh\ b lZdbo nZdlhjh\ dZd fmlZpbb hl[hj b fb]jZpbb Bgl_gkb\ghklv dZ`^h]h ba gbo fh`gh \ujZablv ihkj_^kl\hf dhwnnbpb_glh\ u, s, m.

Ih :cZeZ ^j_cn ]_gh\ [m^_l hkgh\guf nZdlhjhf dh]^Z ijhba\_^_gb_ •N•xi ]^_ xi h^bg ba mihfygmluo dhwnnbpb_glh\ 1 wnn_dlb\guc jZaf_j ihimeypbb Dh]^Z \_ebqbgZ \ujZ`_gby [ebadZ beb [hevr_ _^bgbpu fmlZpbb hl[hj beb fb]jZpby klZgh-

\ylky j_rZxsbfb =jZgl [6] hij_^_ebe mkeh\by ^ey ij_bfms_kl\_ggh]h ^_ckl\by ^j_cnZ ]_gh\ ijb \_ebqbgZo \ujZ`_gby •N•xiZ ^ey bo kh\f_klgh]h ^_ckl\by bg- l_j\Ze \ dhlhjhf \_ebqbgZ wnn_dlb\ghc qbke_gghklb ihimeypbb[i≤ N ≤ 0.5 xi.

46

>jm]bf ih^oh^hf fh`_l [ulv ihkljh_gb_ g_keh`guo h^ghehdmkguo fZl_fZlb- q_kdbo fh^_e_c oZjZdl_jbamxsbo baf_g_gb_ qZklhlu dhgljhebjm_fh]h Zee_ey aZ h^gh ihdhe_gb_ Ijb wlhf lj_[m_lky ^hims_gb_ qlh ^_ckl\b_ w\hexpbhgguo nZdlhjh\ Z^-

^blb\gh GZijbf_j ijb khq_lZgbb hl[hjZ b fmlZpbhggh]h ijhp_kkZ h[s__ baf_g_gb_ qZklhlu Zee_ey jZ\gh kmff_ baf_g_gbc qZklhl \ dZ`^hf hl^_evghf ijhp_kk_

∆ qh[s = ∆ qhl[ + ∆ qfml

(73)

∆ qh[s.= pq[q(W2-W1) + p(W1- Wo)]/Ws + pu - qv,

(74)

]^_XbYkdhjhklb ijyfh]h b h[jZlgh]h fmlbjh\Zgby

Ihkljh_gb_ ]jZnbq_kdbo aZ\bkbfhkl_c ∆ qh[s hl T iha\hey_l \ jy^_ kemqZ_\ \u- y\blv ^hihegbl_evgu_ lhqdb jZ\gh\_kby beb gZclb baf_g_gby iheh`_gby ^ey m`_ km- s_kl\mxsbo

=_g_lbq_kdZy ^bnn_j_gpbZpby \ ijhp_kk_ \b^hh[jZah\Zgby

H^gbf ba ijhy\e_gbc ijhp_kkZ w\hexpbb y\ey_lky \gmljb b f_`\b^h\Zy baf_g- qb\hklv JZaebqby \ n_ghlbibq_kdbo ijbagZdZo hkgh\Zgu gZ kms_kl\mxsbo ]_ghlbib- q_kdbo jZaebqbyo Ihke_^gb_ h[gZjm`b\Zxl k_[y dh]^Z m^Z_lky ihemqblv \lhjh_ ihdh- e_gb_ ihlhfdh\ ijb kdj_sb\Zgbb [ebadhjh^kl\_gguo \b^h\ H^gZdh \ jZfdZo f_g^_- e_\kdhc ]_g_lbdb g_\hafh`gh hl\_lblv gZ \hijhk dZdh\Z kl_i_gv ]_g_lbq_kdbo jZaeb- qbc f_`^m ihimeypbyfb h^gh]h \b^Z beb f_`^m jZaebqgufb \b^Zfb K ^jm]hc klhjh- gu hl\_lb\ gZ wlhl \hijhk \ jZfdZo fhghnbe_lbq_kdhc dhgp_ipbb hj]Zgbaf_gghc w\hexpbb fh`gh g_ lhevdh mklZgh\blv kl_i_gv jh^kl\Z gh b kjZ\gblv w\hexpbhggu_ jZkklhygby hl^_eyxsb_ ij_^klZ\bl_e_c ^\mo jZaebqguo \b^h\ ^jm] hl ^jm]Z b hl bo

h[s_]h ij_^dZ

< gZklhys__ \j_fy kms_kl\m_l g_kdhevdh f_lh^h\ ^ey hp_gdb ]_g_lbq_kdhc ^bn- n_j_gpbZpbb ^\mo ihimeypbc Fh`gh mihfygmlv f_lh^ hp_gdb lh`^_kl\_gghklb ]_gh- fh\ b [he__ ih^jh[gh hklZgh\blvky gZ f_lh^_ ZgZebaZ we_dljhnhj_lbq_kdbo k\hckl\ [_edh\ IheZ]Zxl qlh dZ`^hfm Zee_ex \ ehdmk_ khhl\_lkl\m_l k\hy bahnhjfZ [_edZ hlebqZxsZyky hl ^jm]hc bahnhjfu k\h_c ih^\b`ghklvx \ lhdhijh\h^ys_f ]_e_

FZkZlhrb G_c [1] ij_^eh`be ^bnn_j_gpbjh\Zlv ^\_ ihimeypbb gZ hkgh\_ we_d- ljhnhj_lbq_kdh]h ZgZebaZ [_edh\ < dZq_kl\_ oZjZdl_jbklbd ]_g_lbq_kdhc ^bnn_j_g- pbZpbb ij_^eh`_gh ^\Z iZjZf_ljZ ]_g_lbq_kdh_ koh^kl\h , dZd ^hey kljmdlmjguo ]_- gh\ dhlhju_ b^_glbqgu \ h[_bo ihimeypbyo b ]_g_lbq_kdh_ jZkklhygb_'dZd kj_^g__ qbkeh aZf_g Zee_e_c \ dZ`^hf ehdmk_

47

JZkkfhljbf ijbf_j hp_gdb ]_g_lbq_kdhc ^bnn_j_gpbZpbb q_luj_o \b^h\ ijbfZ- lh\ lZ[e gZ hkgh\_ ZgZebaZ ehdmkh\ dh^bjmxsbo [_edb djh\b < klhe[pZo ^ey dZ`^h]h \b^Z ijb\_^_gu pbnjh\u_ bg^_dku Zee_e_c b l ^ < kemqZyo dh]^Z ehdmk ihebfhjn_g \ kdh[dZo mdZaZgu khhl\_lkl\mxsb_ qZklhlu Zee_e_c

LZ[ebpZ

 

Ehdmk

RbfiZga_

=hjbeeZ

=b[[hg

;Z[mbg

 

 

1

2

3

4

5

1.

Ak

96

98 (0.20)

92

96

 

 

 

 

100 (0.80)

 

 

2.

Alb

100

100

100

99

3.

Aph

100

100

100

100

4.

Cer

100

98

98

102

5.

Dia

100

85 (0.67)

100 (0.67)

95 (0.88)

 

 

 

 

95 (0.33)

108 (0.33)

100 (0.12)

6.

Est-A

100

101

102

96

7.

Est-B

100

100

102

95 (0.17)

 

 

 

 

 

 

96 (0.08)

 

 

 

 

 

 

103 (0.75)

8.

G6pd

100

100

102

102

9.

Got

100

100

96

96 (0.14)

 

 

 

 

 

 

100 (0.86)

10.

Hb

100

100

100

100 (0.92)

 

 

 

 

 

 

102 (0.08)

11.

Hpt

105

107

107

107

12.

Icd

96

100

100

100 (0.94)

 

 

 

 

 

 

107 (0.06)

13.

Lap

100

100

100

100

14.

Ldh-A

96

96

96

96

15.

Ldh-B

100

100

100

100

16.

Mdh

100

100

93 (0.62)

106

 

 

 

 

 

100 (0.38)

 

17.

6-Pgd

97

97 (0.15)

94

94

 

 

 

 

105 (0.85)

 

 

18.

Pgm-

96 (0.12)

100

100

94

1

 

 

100 (0.88)

 

 

 

19.

Pgm-

100

96

100

102

2

 

 

 

 

 

 

>ey jZkq_lZ ]_g_lbq_kdh]h koh^kl\Z ^\mo ihimeypbc ih h^ghfm ehdmkm bkihevamxl ke_-

^mxs__ \ujZ`_gb_

I k

=

ai bi

,

(75)

ai2

 

 

bi2

 

]^_ ai b bi qZklhlu Zee_e_c i - ehdmkZ \ ihimeypbyo : b <

48

< dZq_kl\_ ijbf_jZ jZkkfhljbf ihimeypbb rbfiZga_ b ]hjbeeu Ehdmk Alb (2) fhghfhjn_g b ij_^klZ\e_g h^bgZdh\uf Zee_e_f k bg^_dkhf QZklhlu wlh]h Zee_ey

a100 b b100 jZ\gu _^bgbp_ b ihdZaZl_ev ]_g_lbq_kdh]h koh^kl\Z jZ\_g

I

2 =

1

1

= 1

 

12

 

 

 

 

+12

 

Ehdmk&HU\ dZ`^hc ihimeypbb fhghfhjn_g gh ij_^klZ\e_g jZaebqgufb Zee_eyfb

k bg^_dkZfb b IhdZaZl_ev ]_g_lbq_kdh]h koh^kl\Z jZ\_g

I 4 =

(10) + (0

1)

= 0

(12 + 02 )(02 +12 )

 

 

Ehdmk3JG\ ihimeypbb rbfiZga_ fhghfhjn_g b ij_^klZ\e_g Zee_e_f k bg^_d- khf \ ihimeypbb ]hjbeeu ihebfhjn_g Zee_eb k bg^_dkZfb b < wlhf kem- qZ_ ihdZaZl_ev ]_g_lbq_kdh]h koh^kl\Z jZ\_g

I17 =

 

(10.15) + (0 0.85)

= 0.15 = 0.174

 

 

 

(12 + 02 )(0.152 + 0.852 )

 

 

0.863

 

 

 

 

Kl_i_gv ]_g_lbq_kdhc ^bnn_j_gpbZpbb ^\mo ihimeypbc hp_gb\Zxl ih g_kdhevdbf eh-

dmkZf >ey wlh]h fh`gh \hkihevah\Zlvky nhjfmehc dhlhjmx ij_^eh`be F G_c [1]:

 

 

 

 

 

 

 

I =

 

I ab

 

 

 

(76)

 

 

 

 

 

 

 

Ia Ib

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

]^_ Iab kj_^gyy kmffZ ijhba\_^_gbc qZklhl h^ghbf_gguo Zee_e_c

 

 

 

 

 

 

 

 

 

 

 

N

Q

 

 

 

 

 

 

 

 

 

 

 

 

 

∑∑

DL EL

 

 

 

 

 

 

, D

=

 

 

 

 

,

(77)

 

 

 

 

 

N

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Ia kj_^gyy kmffZ d\Z^jZlh\ qZklhl \ i_j\hc ihimeypbb

 

 

 

 

 

 

 

 

 

 

 

k

n

(ai2 )

 

 

 

 

 

 

 

 

 

Ia

=

1

 

1

 

 

,

 

 

 

(78)

 

 

 

 

 

k

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Ib lh`_ ^ey \lhjhc ihimeypbb

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

k

n

(bi2 )

 

 

 

 

 

 

 

 

 

Ib

=

1

1

 

.

 

 

 

(79)

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

k

 

 

 

 

< jZkkfhlj_gghf ijbf_j_

 

 

 

 

 

 

 

 

 

 

 

 

 

Iab

=

1 + 0 + 0.15

= 0.383

Ia

=

1 + 1 + 1 +

= 1

 

 

 

 

 

 

 

3

 

 

 

 

 

 

 

 

3

 

 

 

 

 

 

Ib =

1 + 1 + 0.863

= 0.954

I =

0.383

 

 

= 0.392

 

 

1 ×

0.954

 

 

 

3

 

 

 

 

 

 

 

 

 

 

 

 

49

 

 

 

 

 

 

 

 

 

 

 

 

 

Hl ihdZaZl_ey ]_g_lbq_kdh]h koh^kl\Z fh`gh i_j_clb d ihdZaZl_ex ]_g_lbq_- kdh]h jZkklhygby'jZ\gh]h D = − ln I < gZr_f ijbf_j_' OQ Wlh ha- gZqZ_l qlh aZ \j_fy jZa^_evghc w\hexpbb \ lj_o jZkkfhlj_gguo ehdmkZo \ kj_^g_f ijhbahreh aZf_gu beb

J_amevlZlu hij_^_e_gby ]_g_lbq_kdh]h koh^kl\Z,b ]_g_lbq_kdh]h jZkklhygby' f_`^m ij_^klZ\e_ggufb \ur_ ijbfZlZfb ^Zgu \ lZ[e <_ebqbgu,b'ihdZaZgu gZ^ b ih^ ^bZ]hgZevx khhl\_lkl\_ggh

LZ[ebpZ

 

RbfiZga_

=hjbeeZ

=b[[hg

;Z[mbg

RbfiZga_

 

0.60000

0.4844

0.3764

=hjbeeZ

0.5108

 

0.5756

0.4466

=b[[hg

0.7248

0.5523

 

0.5050

;Z[mbg

0.9771

0.8061

0.6833

 

J_amevlZlu ZgZebaZ fh`gh ij_^klZ\blv ]jZnbq_kdb jZkiheh`b\ \b^u ijbfZlh\ \ m]eZo ibjZfb^u ^ebgu j_[_j dhlhjhc jZ\gu ]_g_lbq_kdhfm jZkklhygbx jbk

=b[[hg

=hjbeeZ

 

;Z[mbg

RbfiZga_

Jbk JZkij_^_e_gb_ \ \hh[jZ`Z_fhf ijhkljZgkl\_ \b^h\ \ khhl\_lkl\bb k f_`\b^h\ufb ]_g_lbq_kdbfb jZkklhygbyfb

H^gZdh dh]^Z qbkeh kjZ\gb\Z_fuo ihimeypbhgguo kbkl_f klZgh\blky [hevr_

]jZnbq_kdh_ j_r_gb_ l_jy_l kfuke lZd dZd ^ey1\b^h\ lj_[m_lky©N-1»-f_jgh_ ijh- kljZgkl\h

R bfiZga_

=hjbeeZ

=b[[hg

;Z[mbg

Jbk =bihl_lbq_kdh_ nbeh]_g_lbq_kdh_ ^j_\h g_dhlhjuo ijbfZlh\

50

Dh]^Z bamqZ_fu_ \b^u w\hexpbhggh k\yaZgu bo jh^hkeh\gmx hij_^_eyxl ke_^mxsbf h[jZahf W\hexpbhggh ijh^\bgmluc \b^ \ gZr_f ijbf_j_ wlh rbfiZga_ ihf_sZxl gZ \_jrbgm Gb`_ jZkiheZ]Zxl \b^ ]_g_lbq_kdb gZb[he__ ijb[eb`_gguc d i_j\hfm ]hjbeeZ AZl_f hij_^_eyxl kj_^g__ jZkklhygb_ f_`^m wlbfb ^\mfy \b^Zfb b ]b[[hghf b f_`^m l_fb `_ ^\mfy \b^Zfb b [Z[mbghf

(0.9771+0.8061)/2=0.8916. KjZ\g_gb_ kj_^gbo \_ebqbg ihdZau\Z_l qlh ]b[[hg, ih \b^bfhfm, [eb`_ d rbfiZga_ b ]hjbee_ q_f [Z[mbg Ke_^h\Zl_evgh gZb[he__ ijZ\^hih^h[gZ ko_fZ nbeh]_gbb ihdZaZggZy gZ jbk

Kibkhd ebl_jZlmju

1.:cZeZ N <\_^_gb_ \ ihimeypbhggmx b w\hexpbhggmx ]_g_lbdm F Fbj

230 k

2.J_cf_jk B N Hkgh\gu_ [bheh]bq_kdb_ ihgylby b l_jfbgu±F Ijhk\_s_gb_

k =jZgl < W\hexpbhgguc ijhp_kk F Fbj k

3.DZc^Zgh\ E A =_g_lbdZ ihimeypbc F <ukrZy rdheZ k

4.DbfmjZ F Fhe_dmeyjgZy w\hexpby l_hjby g_cljZevghklb F Fbj k

5.K_\_jph\ : K Hkgh\u l_hjbb w\hexpbb F Ba \h F=M k

6.=jZgl < W\hexpbhgguc ijhp_kk F Fbj k

51

Ij_^f_lguc mdZaZl_ev

 

Z^ZilZpbhggZy p_gghklv ]_ghlbiZ......

34

]Zevlhgh\kdb_ ijbagZdb.........................

5

]_g_lbdh Z\lhfZlbq_kdb_ ijhp_kku....

23

]_g_lbq_kdZy ^bnn_j_gpbZpby............

47

=_g_lbq_kdbc ]jma.................................

43

]_g_lbq_kdh_ koh^kl\h..........................

48

=_ghlbibq_kdh_ jZaghh[jZab_...............

5

]_l_jhab]hlghklv.....................................

8

]_l_jhab]hlghklv fZdkbfZevgZy

 

h`b^Z_fZy.............................................

9

]_l_jhab]hlghklv ihebfhjnguo ehdmkh\

...............................................................

8

]_l_jhab]hlghklv l_hj_lbq_kdb

 

h`b^Z_fZy.............................................

9

]jmiiu djh\b...........................................

5

^bgZfbdZ ]_g_lbq_kdh]h ]jmaZ.............

44

^j_cn ]_gh\............................................

23

_kl_kl\_gguc hl[hj ..............................

34

bg[jb^bg]..........................................

7, 25

Dhebq_kl\h ]_ghlbih\............................

7

dhwnnbpb_gl bg[jb^bg]Z ....................

26

dhwnnbpb_gl hl[hjZ.............................

34

djbl_jbc ihebfhjnghklb ......................

8

fZdkbfZevgZy ]_l_jhab]hlghklv

 

ihimeypbb...........................................

13

f_g^_e_\kdb_ ijbagZdb ..........................

5

fb]jZpby.................................................

31

fgh`_kl\_ggu_ Zee_eb ...........................

7

FmlZpbb .................................................

28

hl[hj ]Zf_l.............................................

45

iZgfbdkby ..............................................

15

iZgfbdlbq_kdZy ihimeypby..................

14

ieZlZ aZ hl[hj ..................................

43, 44

ihebfhjnghklv........................................

8

ihebnZdlhjbZevgu_ ]_ghlbiu.............

18

ihlhd ]_gh\.............................................

31

ijbagZdb kp_ie_ggu_ k ihehf ............

16

ijbkihkh[e_gghklv ...............................

34

iml_\hc ZgZeba.......................................

27

jZkij_^_e_gb_ ImZkkhgZ.................

13, 29

j_r_ldZ I_gg_lZ....................................

36

l_fi fmlbjh\Zgby..................................

30

ojhfhkhfgZy i_j_kljhcdZ ....................

28

qZklhlZ Zee_ey..........................................

6

q Z k l h l g h a Z \ b k b f u c h l [ h j ...45

w\hexpbhggh_ jZkklhygb_....................

47

wnn_dl [mluehqgh]h ]hjeurdZ ........

25

wnn_dl hkgh\Zl_ey .............................

25

wnn_dlb\gZy qbke_gghklv ihimeypbb.24

52

=Z_\kdbc GbdheZc :e_dkZg^jh\bq

AgZdhfkl\h k w\hexpbhgghc ]_g_lbdhc

Ebp_gaby EJ‹ hl

J_^Zdlhj : : GZabfh\Z

Dhjj_dlhj L ? ;Zklju]bgZ

Ih^ibkZgh \ i_qZlv 12.09.2002

LbjZ`bjm_lky gZ we_dljhgguo ghkbl_eyo AZdZa 79

>ZlZ \uoh^Z 16.09.2002

:^j_k \ Internet: www.lan.krasu.ru/studies/editions.asp

Hl^_e bgnhjfZpbhgguo j_kmjkh\ mijZ\e_gby bgnhjfZlbaZpbb DjZk=M

] DjZkghyjkd ij K\h[h^guc Zm^ e-mail: info@lan.krasu.ru

Ba^Zl_evkdbc p_glj DjZkghyjkdh]h ]hkm^Zjkl\_ggh]h mgb\_jkbl_lZ

] DjZkghyjkd ij K\h[h^guc , e-mail: rio@lan.krasu.ru

53