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i hc khoa hc t nhin - i hc quc gia h ni Khoa sinh hc - b mn di truyn hc

DI TRUYN HC qun th & tin ha

INH ON lONG

Di truyn hc c s

Ni dung
Mt s nh ngha S di truyn trong qun th Cc yu t nh hng n trng thi cn bng di truyn ca qun th Di truyn hc qun th v s hnh thnh cc loi

Tin ha hc phn t
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Ni dung
Mt s nh ngha S di truyn trong qun th Cc yu t nh hng n trng thi cn bng di truyn ca qun th Di truyn hc qun th v s hnh thnh cc loi

Tin ha hc phn t
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Mt s nh ngha Theo di truyn hc, loi (species) l tp hp cc c th sinh vt thc s hoc c tim nng giao phi vi nhau. Cc thnh vin ca mt loi cng chia x vn gen chung. Cc loi khc nhau trong t nhin khng c kh nng trao i vt cht di truyn vi nhau. Nh vy, s ngn cch cc loi ch yu da trn tiu chun s ngn cch giao phi (ngn cch di truyn/ngn cch sinh sn?). Vn gen (gen pool) l tp hp ton b cc thng tin di truyn, tc l b y cc alen ca tt c cc gen c trong mt qun th hay mt loi ti thi im xc nh.
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Mt s nh ngha Qun th (population) l tp hp cc c th trong cng mt loi c cng mi trng sinh cnh, tn ti trong cng mt thi im nht nh, c tim nng giao phi vi nhau. Tn s alen (alen frequency, i khi gi tt l tn s gen ) l s bn sao ca mt alen chia cho tng s bn sao ca tt c cc alen c trong qun th. Di truyn hc qun th (population genetics) l mt chuyn ngnh ca di truyn hc chuyn nghin cu v tn s cc kiu gen v tn s alen qua cc th h.

Ni dung
Mt s nh ngha S di truyn trong qun th Cc yu t nh hng n trng thi cn bng di truyn ca qun th Di truyn hc qun th v s hnh thnh cc loi

Tin ha hc phn t
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S di truyn trong qun th


i vi qun th t th Xt v tn s kiu gen, qu trnh t th phn ( thc vt) l qu trnh ng hp t ha. Qua tng th h, tn s c th ng hp t tng dn, ngc li s d hp t gim dn. Hin tng tng t cng xy ra i vi ng vt khi xy ra hin tng giao phi gn (ni phi). i vi ngi l hn nhn cn huyt.
I II III IV V VI VII VIII Aa Aa AA Aa Aa aa AA AA Aa aa AA aa AA aa AA aa

AA

aa aa

S di truyn trong qun th


i vi qun th ngu phi

Trn c s ca c ch gim phn, to giao t v th tinh c th d dng nhn thy rng i vi cc qun th ngu phi t l mt kiu gen no s l tch tn s cc alen tng ng. Qun th c coi l cn bng khi tn s cc alen v kiu gen trong qun th duy tr n nh qua cc th h. Nm 1908, Hardy & Weiberg pht hin ra cng thc v tn s kiu gen ca mt qun th ngu phi cn bng lun l: Tn s kiu gen: p2[A1A1] : 2pq[A1A2] : q2[A2A2], Trong , p v q ln lt l tn s hai alen A1v A2 (gi s locut ch c 2 alen), ngha l p + q = 1.
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S di truyn trong qun th


i vi qun th ngu phi

C th d dng nhn thy cng thc Hardy-Weinberg thc cht l nh thc Newton Tn s kiu gen: (pA1 + q A2)2, vi p + q = 1. Cc qun th ngu phi c tn s cc kiu gen v alen khng ph hp vi cng thc Hardy Weinberg l cc qun th khng cn bng. Tuy vy, nu hin tng ngu phi xy ra hon ton v qun th ln, chi sau 1 th h duy nht, qun th khng cn bng s chuyn v trng thi cn bng.

S di truyn trong qun th


i vi qun th ngu phi

i vi cc loct nhiu hn hai alen, cng thc Hardy Weinberg c m rng theo nguyn tc biu thc Newton, v d: tn s kiu gen = (pA1 + q A2+ rA3)2, vi p + q + r = 1 (vi locut c 3 alen); hoc (pA1 + qA2+ rA3 + sA4), vi p + q + r + s = 1 (vi locut c 4 alen), v.v Cc qun th ngu phi c tn s cc kiu gen v alen khng ph hp vi cng thc Hardy Weinberg l cc qun th khng cn bng. Tuy vy, nu hin tng ngu phi xy ra hon ton v qun th ln, chi sau 1 th h duy nht, qun th khng cn bng s chuyn v trng thi cn bng.

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S di truyn trong qun th


i vi qun th ngu phi

i vi qun th ni phi mt phn, cng thc HardyWeinberg c hiu chnh l: (p2 + fpq) (A1A1) + (2pq - 2fpq) (A1A2) + (q2 + fpq) (A2A2) Trong , f l tn s c th ni phi trong qun th. Trong thc t, f c tnh bng: [1 (tn s d hp t quan st / tn s d hp t l thuyt)]

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ng dng ca nh lut hardy-weinberg Xc nh tn s cc alen t tn s cc kiu hnh

Nu hai alen sinh ra ba kiu hnh khc bit, tn s alen tnh bng tn s ng hp t + 1/2 d hp t. Nu qun th trng thi cn bng, tn s alen ln tnh bng cn bc hai tn s kiu hnh ln. Nu qun th trng thi cn bng, tn s alen ln lin kt NST X bng s c th c biu hin tnh trng / tng s c th c.

Qun th trng thi cn bng, nhng tn s ng hp t cao hn l thuyt l do ni phi (ni phi lm thay i tn s kiu gen nhng khng lm thay i tn s alen). C th xc nh tn s ni phi t s d hp t quan st v s d hp t l thuyt.
T vn di truyn
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Ni dung
Mt s nh ngha S di truyn trong qun th Cc yu t nh hng n trng thi cn bng di truyn ca qun th Di truyn hc qun th v s hnh thnh cc loi

Tin ha hc phn t
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Cc yu t nh hng trng thi cn bng di truyn t bin. mi th h, vn gen ca qun th thng c b sung thm bi nhng t bin mi. S nh hng ca s lng t bin n t l cc kiu gen v alen trong qun th gi l p lc t bin. p lc t bin (mc nh hng) lin quan n s t bin thun (mi) v nghch (phc hi) hnh thnh.

Chn lc. c 3 phng thc chn lc.


Chn lc bnh n Cc c th trc khi chn lc Qu trinh thch ng Cc c th sau khi chn lc 14 Chn lc nh hng Chn lc tch ly

Cc yu t nh hng trng thi cn bng di truyn


Chim s Passer domesticus b cht trong nhng trn bo tuyt d di New York thng c cnh di hn hoc ngn hn so vi gi tr trung bnh. Trc y, tr s sinh c trng lng cch xa 3,6 kg thng b cht hoc b tt bnh ngay khi sinh hoc sau khi sinh v chng hoc qu to gy kh sinh , hoc qu b sng st. loi chim s Geospiza fortis qun o Galapagos, vo ma kh c v nhng cy thn tho nh thng sinh trng km, nhng nhng cy c r su v ht to vn kt qu. Chn lc u tin nhng con chim s c m to, c kh nng n c nhng ht ln v chng li nhng con chim c m nh, ch n c nhng ht c.

V d chn lc bnh n:

V d chn lc nh hng:

V d chn lc tch ly:

Chim ct mi Accipiter nisus c kch thc ln gp i chim trng, chuyn bt nhng con mi ln, cn chim trng th ngc li. Chn lc tch ly c th l mt c ch tin ha thnh hai loi mi, nhng cn c c ch cch ly trc giao phi ngn cn s giao phi gia chng c th din ra.

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Cc yu t nh hng trng thi cn bng di truyn Kch thc qun th. Kch thc qun th cng nh, h s ni phi cng ln, lm tng cc th ng hp t, v ngc li.

Nhp c. Mang n cc kiu gen t mt vn gen khc, gy ra sai lch. Nu cc kiu gen khc nhau c xu hng di c hoc nhp c khc nhau th mc sai lch s din ra theo mt chiu hng nht nh . Giao phi chn la. V d: s cc oan mt dng h dn n s hn nhn gia nhng ngi trong h hng, chng hn 33% ca bnh alkapton niu l do c cp b m l anh em h hng ly nhau. Nu theo xu hng ngc li, s lm tng s c th d hp t. Lc dng di truyn. Do c hi di truyn thnh cng ca cc alen nht nh t th h ny sang th h khc c th khc nhau, gy sai khc v tn s kiu gen so vi mong i.
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Cc lc lm thay i tn s alen trong qun th


t bin

T l t bin nhn chung l thp, song khc nhau gia cc locut v gia cc loi khc nhau. Ngoi kh nng t bin t pht, nhiu yu t mi trng c tc ng trc tip lm tng tn s t bin (ha cht, chiu x, cc tc nhn ly nhim ). t bin l ngun gc tn cng ca mi bin d mi. t bin c th trung tnh, c hi (phn ln) hoc c li (t) v vic chng c c duy tr khng ph thuc vo cc iu kin c th ca mi trng.
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Cc lc lm thay i tn s alen trong qun th


t bin

Phn ln cc gen c th t bin xui v ngc. t bin ngc thng xuyn xy ra vi tn s thp hn so vi t bin xui.

S thay i tn s alen qua mi th h:

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Cc lc lm thay i tn s alen trong qun th


Sai lc di truyn

nh lut Hardy-Weiberg gi thit cc qun th c kch thc ln, giao phi hon ton ngu nhin. Cc qun th trong thc t thng c kch c khng phi lun lun n nh; tuy vy, chng cng ln cc yu t ngu nhin ch gy tc ng nh n cc tn s alen. Mi s thay i tn s alen ca qun th bi cc s kin ngu nhin (t xut) c gi l s sai lc di truyn ngu nhin. S thay i tn s alen do cc s kin ngu nhin c vai tr quan trng trong tin ha cc qun th nh.

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Cc lc lm thay i tn s alen trong qun th


Sai lc di truyn

S bin ng ngu nhin so vi l thuyt gi l li ly mu, ging nh khi chng ta tung ng xu 4 ln thu c 3 mt xp : 1 mt nga, thm ch c 4 mt u nga. Khi s mu cng nh, li ly mu cng ln.

Sai lc di truyn l ngu nhin, nn chng ta khng th d on c s thay i tn s alen. Tuy vy, do kch thc qun th nh hng trc tip n li ly mu. Nn chng ta d on c mc nh hng (cng ) ca s sai lc di truyn, qua kch thc qun th hu hiu (Ne).

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Cc lc lm thay i tn s alen trong qun th


Sai lc di truyn

Nu t l c/ci trong qun th l nh nhau v mi c th c sc sinh sn tng ng th kch thc qun th hu hiu ng bng tn s ca c th tui sinh sn ca qun th, (Ne = 2 Nf/m) Nhng nu s c th c/ci khng bng nhau th kch thc qun th hu hiu (Ne) bng:

Vi: Nf = s c th ci trong qun th tham gia sinh sn. Nm = s c th c trong qun th tham gia sinh sn.
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Cc lc lm thay i tn s alen trong qun th


Sai lc di truyn

S cn bng gia t bin v sai lc di truyn ngu nhin

Trong : H l t l s cc th d hp t; : t l t bin trung tnh Ne: kch thc qun th hu hiu

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Cc lc lm thay i tn s alen trong qun th


Di nhp c

Tn s alen A (p) qun th y sau khi mt s c th di c n t qun th x v chim tn s m qun th y mi l:

Trong , px v py l tn s alen A tng ng ti cc qun th x v y, m l t l c th t qun th x trong qun th y mi. S thay i tn s alen qun th y sau khi c s nhp c:

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Cc lc lm thay i tn s alen trong qun th


Chn lc t nhin

Chn lc t nhin khng c ngha l u tranh sinh tn ngha quan trng nht ca chn lc t nhin l s gen tng i ng gp vo cc th h tng lai ca qun th.

S sng st n thun l khng d, m kh nng truyn gen qua cc th h c quyt nh bi kh nng sinh sn.

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Cc lc lm thay i tn s alen trong qun th


Chn lc t nhin

Chn lc t nhin c o bi mc thch nghi, (k hiu l W) phn nh kh nng sinh sn tng i ca mt kiu gen.

W = 1 vi kiu gen sinh sn nhiu th h con nht

H s chn lc (s = 1 - w) l i lng o mc chn lc tng i chng li mt kiu gen nht nh.

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Cc lc lm thay i tn s alen trong qun th


Chn lc t nhin

nh hng ca chn lc n tn s alen. Chn lc chng li tnh trng ln (s> 0 vi aa).

Chn lc chng li tnh trng ln trit (s = 1)

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Cc lc lm thay i tn s alen trong qun th


Chn lc t nhin

Chn lc chng li tnh trng tri (s >0 vi AA v Aa)

Chn lc chng li d hp t (s> 0 vi Aa)

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Cc lc lm thay i tn s alen trong qun th


Chn lc t nhin

Chn lc u tin d hp t (s > 0 vi AA v aa)

Cng thc chung:

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Cc lc lm thay i tn s alen trong qun th


, , ,
Chn lc t nhin

Kt hp gia t bin v chn lc: V tn s alen ln gy hi thng him

Nu xy ra t bin

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Cc lc lm thay i tn s alen trong qun th


Chn lc t nhin

Nu alen ln gy cht trng thi ng hp t (s = 1)

i vi alen tri, tn s trng thi cn bng l

Lu , do tc ng ca chn lc cc alen tri gy hi thng him hn nhiu so vi cc alen ln gy hi.

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Mt s cng thc tnh h s tng ng v hi quy

Cng thc tnh h s tng ng

Cng thc hi quy cho hm

y = ax + b

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Ni dung
Mt s nh ngha S di truyn trong qun th Cc yu t nh hng n trng thi cn bng di truyn ca qun th Di truyn hc qun th v s hnh thnh cc loi

Tin ha hc phn t
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di truyn hc qun th v s hnh thnh cc loi

Ngay t nhng thi k s khai ca x hi loi ngi, mt cu hi c t ra l: chng ta t u n? chng ta l ai? chng ta s i n u?

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Hc thuyt tin ha ca acuyn (1859) Trong cun The Origin of Species (1859), Darwin cho rng s bin i vn gen trong loi do s cnh tranh gia cc c th. Cc dng bin d c li s c tn ti v duy tr, cn cc dng bin d khng c li s b mt i. Qu trnh nh vy din ra theo thi gian v c gi l qu trnh chn lc t nhin. Trong qu trnh chn lc t nhin, nu c tnh c trng ca mt loi b bin i ln th s l c s hnh thnh loi mi.
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thuyt hnh thnh loi ngi theo ha thch Aegyptopithecus (mt dng ng vt ging kh) l t tin ca vn, vn ngi v ngi xut hin cch y khong 30 40 triu nm Dryopithecus (t tin ca vn ngi v ngi) sng cch y khong 22 32 triu nm bt u tp hp thnh by, n trn ng c. Hominid (t tin ca loi ngi) xut hin khong 19 triu nm trc . Australopithecus l dng ngi tin s xut hin cch y khong 4 triu nm trc, i bng hai chn thng ng, bt u bit s dng cng c. Nh vy, vng phn b m rng nhanh chng.
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thuyt hnh thnh loi ngi theo ha thch T Australopithecus, khong 2 triu nm trc hnh thnh Homo habilis (l nhnh sau ny pht trin thnh ngi hin i). Homo habilis sau tin ha thnh Homo erectus, s dng cng c thnh tho v t chc x hi mc cao hn (phc tp hn). Homo sapiens (ngi hin i) tch hnh thnh loi mi t Homo erectus vo khong 30.000 n 50.000 nm trc. Nh vy, theo nghin cu ha thch, ngi hin i hnh thnh cch y khong 40.000 nm. Ha thch ngi ging ht chng ta ngy nay c nht tm thy c tui 5300 nm.
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thuyt hnh thnh loi ngi theo ha thch

Ha thch ngi c 5300 nm (Oetzi) c tm thy dy ni Alp (gia A v Italia), c c im ging ht ngi chng ta hin nay, l ha thch ging chng ta c nht tm thy.
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Loi v s hnh thnh loi

Cc loi khc nhau b ngn cn v kh nng giao phi, c th do cc c ch cch ly, c phn thnh hai nhm (theo Dobzhansky): (1) Cch ly trc giao phi, v (2) Cch ly sau giao phi.
Cch ly trc giao phi

Cch ly tp tnh (gii tnh v phong tc): cc c th trong mt loi chn cc c th cng loi, ch khng phi cc c th khc loi lm bn giao phi. Cch ly khng gian (a l, sinh thi v mi trng): cc c th ca cc loi khc nhau giao phi cc a im v mi trng khc nhau,
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Loi v s hnh thnh loi

Cch ly trc giao phi Cch ly thi gian: cc loi khc nhau giao phi vo cc thi gian, thi im khc nhau trong ngy, trong nm. V d: mt s loi thc vt ch n hoa vo nhng thi im nht nh, Cch ly c hc: c quan sinh dc ngoi hoc b phn ca hoa cn tr s giao hp hoc ny mm ca ht phn, Cch ly do cc nhn t th phn: c nhng lai c quan h gn gi, nhng thu ht cc nhn t th phn (v d cc loi cn trng) khc nhau Cch ly giao t: V d: cc lai giao phi ngoi, cc giao t c hin tng hp dn chn lc. Tnh trng ca mt loi ch bi hng ti v kt hp vi trng ca chnh lai . thc vt, nhy hoa c th ngn cn s xm nhp ca ht phn mang tnh khng hp.
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Loi v s hnh thnh loi

Cch ly sau giao phi Con lai khng sng st: Mc d hai loi c kh nng giao phi, nhng hp t lai khng c kh nng sng st, hoc c sc sng gim. Hu qu l do s tng tc khng hp gia cc sn phm gen hai loi b, m khng ph hp vi nhau Con lai bt th: t nht mt trong hai gii tnh ca con lai F1 b bt th hoc gim kh nng hu th, Con lai suy thai: cc con lai t th h F2 khng c kh nng sng st, bt th, hoc gim sc sng hay kh nng hu th,

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Loi v s hnh thnh loi

S hnh thnh loi l qu trnh phn ly hnh thnh hai hay nhiu loi mi t mt loi ban u. Cy m t quan h h hng cng ngun gc c gi l cy chng loi pht sinh. S hnh thnh loi mi c th xy ra, do: (i) dng gen gia cc qun th ca mt loi b hn ch n mc lm cho chng phn ly, khng cn kh nng giao phi vi nhau na, (ii) mt thnh phn ca vt cht di truyn b phn ly trong qu trnh tin ha to thnh ro cn giao phi (cch ly giao phi). Trong qu trnh phn ly cc loi, cc ro cn c th l khng gian, thi gian, hoc thm ch l khc bit kh hu. Cc loi hnh thnh do cch ly a l gi l cc loi a l.
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Loi v s hnh thnh loi


(a)
(b) E H sinh thi suy gim C s gii hn vng
S cch ly

Cc hn o

Loi t lin C s di c A
F G

C s bin i Cc vng di truyn


lnh ln

(c) F thch nghi hn E Cch ly trc giao phi F

V d v s hnh thnh 5 loi t mt loi ban u do s cch ly v a l, kh hu, mi trng sng, tp tnh, v.v

F thay th E

C s m rng vng C s tip xc tr li B/C ha nhp Khng c c ch cch ly

S di c tr li F v G cng tn ti D

S phn ly sinh thi v cch ly trc giao phi

Vng lai Ch c s cch ly sau giao phi

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Tin ha hc phn t Nu nh trc y cc nghin cu v tin ha v s hnh thnh loi ch yu da trn cc bng chng ha thch. Th trong hn 20 nm tr li y, cc nghin cu di truyn hc phn t c nhng ng gp mang tnh cch mng cho cc nghin cu tin ha, m ra mt chuyn ngnh mi l tin ha hc phn t.
So snh h gen

Ly v d cc loi linh trng, xt trn ton h gen, ngi ging tinh tinh 99,5% cc gen m ha cho protein. Nh vy, ch 0,5% phn h gen m ho protein l khc nhau nhng to ra s khc bit r rt. Nh vy, s khc bit v cch ly gia cc loi i khi ch cn c quyt nh bi mt s lng gen hn ch. V d 1: 01 gen keratin tham gia to lng trn thn hot ng mnh i i v tinh tinh, nhng khng hot ng ngi (gen gi do mt t bin v ngha) to ra s khc bit v kiu hnh lng trn thn gia ngi v cc loi linh trng ny.
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Tin ha hc phn t s khc bit v cch ly gia cc loi i khi ch cn c quyt nh bi mt s lng gen hn ch. V d 2: 01 gen m ha mt hemoglobin c th biu hin ngi, nhng khng c hoc ch c lng t cc loi linh trng khc biu hin trong giai on pht trin t phi bo thai, gp phn ko di thi gian pht trin bo thai ngi tng ti a s pht trin ca no b khi mang thai pht trin nhiu kh nng mi (vd: ngn ng, kh nng t duy, v.v) V d 3: mt s gen n duy nht c tm thy iu khin ko di qu trnh trng thnh (trc dy th) ngi so vi cc loi linh trng khc.

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Tin ha hc phn t Trong qu trnh tin ha, t bit l c ch to ra s a dng ca cc gen, cn chn lc t nhin l c ch sng lc v duy tr cc vn gen. Hay ni cch khc, l cc gen mi khng ngng c hnh thnh chu s tc ng ca chn lc t nhin. V d: kh nng cm nhn mi ngi c lin quan n 906 gen va gen gi c trong h gen. Trong s 906 gen v gen gi lin quan n cm nhn mi c 60% l gen gi, tng cng chim khong 1% h gen. Ngi ta cho rng t tin loi ngi hng triu nm trc ph thuc nhiu hn vo th th cm nhn mi c th tn ti. Sau ny, chn lc loi b bt cc gen m ha cc th th khng cn quan trng cho s sng cn ca con ngi na, nn hnh thnh nhng t bin cm.

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Tin ha hc phn t
So snh h gen

S so snh kiu hnh nhum bng NST cng l c s cho thy mi quan h gia cc loi. Cc vng NST ging nhau nhiu loi khc nhau thng c gi l cc vng bo th.

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Tin ha hc phn t
So snh nhim sc th

Kiu hnh nhum bng NST ca ngi ging nht l vi tinh tinh, sau l kh gorilla v i i. Phn ln kiu nhn ca ngi khc vi cc loi linh trng khc l do o on xy ra trong cng NST. i vi cc loi vn khc xa hn, s khc bit ch yu l do chuyn on xy ra gia cc NST khc nhau. Nu c hai NST s 2 ca ngi b gy lm hai th th t bo s c 48 NST ging nh ba loi tinh tinh, kh gorilla v i i. ng ch NST X ca phn ln cc loi ng vt c v c kiu hnh nhum bng ging nhau.
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Tin ha hc phn t
So snh cc trnh t protein

Bi v tt c cc loi sinh vt trn tri t u s dng chung m di truyn, nn c th gi thit tt c mi sinh vt trn tri t u c t tin chung. Cc sinh vt d rt khc nhau, nhng c nhiu protein ging nhau v chc nng v cu trc. Trong c th ly v d v mt trong nhng protein c tnh bo th cao nht l cytochrome C.
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Tin ha hc phn t
So snh cc trnh t ADN

S so snh trnh t nucleotit cc gen hoc ton b h gen cho php nh gi chnh xc hn c mi quan h h hng gia cc loi. V nguyn tc, ngi ta c th phn tch cc ch th ADN, cc trnh t gen hoc dng phng php lai xc nh s khc bit ny.
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Tin ha hc phn t
So snh cc trnh t ADN

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Tin ha hc phn t
ng h phn t

Da trn s sai khc v trnh t nucleotit trn cc phn t ADN gy ra do t bin, ngi ta c th c tnh c thi im phn ly hnh thnh cc chng tc hoc cc loi mi. Tuy vy, cho n nay s pht sinh v tin ha ca cc chng tc ngi trn th gii cn nhiu ni dung cn cha c lm sng t.
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Tin ha hc phn t

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Tm tt v di truyn qun th v tin ha Di truyn qun th l chuyn ngnh theo di s bin i ca tn s cc kiu gen v alen ca cc qun th di s nh hng ca cc yu t khc nhau, nh t bin, chn lc t nhin, v.v i vi cc qun th ngu phi, tn s cc alen trong qun th c th c tnh t cc tn s kiu hnh trong qun th, da trn nh lut HardyWeinberg. Qun th t phi l qu trnh ng hp t ha. Chn lc xy ra khi cc kiu gen khc nhau c kh nng thch nghi (sng st v sinh sn) khc nhau. Chn lc lm thay i tn s alen trong qun th. C ba c ch chn lc c bn l bnh n, nh hng v tch ly. S bin d di truyn trong qnn th c th c ghi nhn c mc kiu hnh, nhim sc th v phn t (ADN, protein). S tch ly gia cc qun th ngy cng tng dn n s cch ly giao phi l mt c ch quan trng trong qu trnh hnh thnh cc loi. Qu trnh ny c th din ra gia cc qun th cch ly a l (allopatric) hoc trong cng vng a l (sympatric). Cc bng chng ha thch v tin ha phn t cho thy t tin ngi hin i xut hin khong 200.000 nm trc Chu Phi sau pht tn ra cc lc a khc. Cn tn ti nhng vn cha r v s tin ha ca loi ngi ni ring v nhiu loi sinh vt khc ni chung.

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