Chương 5 tương tác gen (bộ môn di truyền học)





 !"#$%!&
'() )*

!"#

1. Hiện tượng gây chết
+, /01-2314563236780-9:01-9.;01<=>764?@--A0<BC7D-9.;01
<=>1801EF3<G-HIEF7D-9.;01<=>7D?J4G0BK4-L7M-<8:BN0<7OP-
Q80R87B/014S00<T-H
UVWXYZ"[
F
\7]01^D01_-9@4`
6\B80<:a3b]3]76_7a0`
c<401.;4-67643<O@-^D01×^D01
d
e
\-<OB.=3<647:f43<O@-\g7]01^D01\e7]01h<23_b]3]76`iBj01-<;4
-9:013237k63<O@-Bl96-<mbn3:03o60CA-</0epqb:^r4323-K<=>764
h<23H
s230<P0tu-0DFB.=3B.6BG014S-<4G-7D3<O@-7]01^D013Ch4vO
180RN<=>-wx
F
6h<43<y01764^r40<6O7D?tOT-<4M03<O@-x
F
x
F
h<]013C

bk3bn01^D3<y01JN3<G-z1464B:f0br?3o6><]4H
{1.;4-67D?-<A01<4M?14S4><|O3<O@-3247]01^D01B601?601
-<64-9:01-K<=>76414567]01^D01×^D01B}Ot23BN0<<4M0 =01-9~0H
IC7D-9:01Rf3:03o63<O@-?•3C?@-bnJD:-<647]01^D01h<]01><2-
-94v0^m?@-bnJ@><P0-9:013€-<v?601Ba3B4v?RN<m0<H{<.-<G3<O@-
Bj01<=>-wx
F
x
F
h<]013Cbk3bn01R:6780x
F
7D67801EF3<G-h<]013<:
Bj01<=>-wbn01B.=3H•23B@013o66780x
F
^}?DO7]017D-9@4b:^r4
67806^m3/-<vRN<=>-wx
F
63C?DO7]01^D01H{<.01^}?a-bk3bn01-<m
‚ƒ
x
F
7f47a0b:^r46^m-K<=>x
F
6^|0bn01Jm0<-<.;01R:678067T02-b,
1EF3<G-3o6x
F
HIEF7D^AR„^}1803C-23B@010DF-9@40<.01-23B@01h46
7D7a0b:^r46780 /01k01H

 !"#$%&#'()#*+,#-" #/01#2130#454#4"+6)#71#"8-#)9#/:!2#;<!2%#'":!2#-",1#)=)

4,#454#4"+6)#>#)"(#"?#@0+#AB+#@C!2#@D)
…†
EFGH
IJKLIKLMNOPQR
6H s<O@-7.01B80iJ„01^D01_-9~0‡-924`iJ„01B80_-9~0‡><S4`^D3<O@-
760161:O-4_R.r4`H
JH c4vO180^Dh4vO<m0< /01k013o63236778783o618061:O-4H
3H ˆ641456323R‰01-<OŠ0-f:96?@-R‹F3C-<v3CŒ677878-<8:?@--<k-,
-9@4Hs<:32332-<vde146:><n4^r40<6O3<:96-<G<Mdg3C-•7Mh4vO
<m0<7DŒ\eHB4}O0DF3<:-<TFxi6-^D67D323677878h<230<6O3o63Ž01
?@-180
…e
•9.;01<=>0DF3‰01a>?@-bnBn4 =01h<230<.323<u>
c<4764323<u>hA0<^r40<6O
d
e
\epqxx \gpqx6\epq66
3<G-\g3<u>hA0<\e3<u>^SF
•01.;4JM0<-<4GO?2O<j013ŠO74}?R:B@-J4G0\
{1.;43Ch4vO180++\-<4GO?2O0a013<G--9.r3h<4-9.z01-<D0<H+x\bk3
h<€8Jm0<-<.;010<.01B]4h<43C-94MO3<k01-<4GO?2O0<•H
STUVWOHX
2. Sự tương tác giữa các alen của cùng một gen
•45632367803o63Ž01?@-1803C?n4‘O60<M-9@4^D7a0H•9:01
?n4 /01-231800DF01.;4-6><2-<4M096<4M0 =01?@-1803C0<4}O
</0g6780H
’AR„\
‡+,R4-9OF}00<C??2Oxi“i”R:Œ6780•
x
i•

“
i•
:
H{<C??2O3o6
Bf4146by33C</0e††6780H
‡+,R4-9OF}0?DO?–-9Oj414T?R:e1801j?eR‹Feg6780‘OF
BN0<i67803On43Ž01?–--9–01_—`^D-A0<-9@414S?RŠ0-<8:<.r01b6O\
…g
˜
™
š˜
b6-
š˜
3:
š˜
˜
š˜
6>Œ
š˜
3<
š˜
8
š˜
J7
š˜
6>
š˜
4
š˜
-

š˜
•./01k01\B€Rf4‡B€b6-bO?6‡b60<]_3:967`‡9.=O0<:_—408`‡
-924BD:_6>943:-`‡3<894‡b:0_8:b40`‡?2O_J7::R`‡-924BD:_6>943:-`‡01D
^:4_4^:9F`‡-9–01B„3_-4018R`‡-9–01_—<4-8`
+,J4vO<4M0-A0<-9f01?DO?–-R:b, /01-231456<646780^r40<6O\
˜
™
˜
J7
\<:601Rf4 ˜
3:
˜
J7
\B€b60<]
‡›‹F6780-9:01^4M3t23BN0<0<C??2Oz01.;4
s2367807D?tOT-<4M00<4}O0<C??2OBa3-9.01z01.;4i74~0
‘O60^r4Ba3B4v?h<20101OF~03o6-<v?2Oi‘OFBN0<b,tOT-<4M0h<201
-<vBa3<4MO-9:01<OFG--<60<?2OH+,><2-<4M0960<C??2O7DR:
ˆ60Rb-84089i]01-<TF-9:01?@-bn-9.;01<=>0<T-BN0<ih<4-9OF}0<j01
3ŠO01.;40DF^D:<OFG--<60<01.;4h<233C<4M0 =0101.01hG-323-<v
?2OHc<4-9OF}0?2Oi<4M0 =010DF3C-<v1EF3<G-H{1.;4-6B‹t23BN0<
-9:01<j013ŠO3Cgh<20101OF~0x^D“i3‰0-9:01<OFG--<60<3Cg
h<201-<v7D?01.01hG-3<y01HœOŠ0-<v01.;4B.=3><E096-<8:Ba3-A0<
3o6?2O-<D0<q0<C?\0<C?x3Ch<20101OF~0x^Dh<201-<vh<201“i
0<C?“3Ch<20101OF~0“^Dh<201-<vh<201xi0<C?x“3C3Sgh<201
01OF~0h<]013Ch<201-<vi0<C?”h<]013Ch<20101OF~0^D3C3Sg
h<201-<vH
•<S0k013o6q0<C??2O^r4<OFG--<60<3Ch<201-<vh<201“^D
h<201-<vh<201x0<.b6O\<j013ŠO0<C??2Ox“01.01hG-^r4<OFG-
-<60<3Ch<201-<vh<201“^Dh<201-<vh<201xHžj013ŠO0<C?x3<LJN

01.01hG-Jz4<OFG--<60<0<C?“H<j013ŠO0<C?“3<LJN01.01hG-Jz4
<OFG--<60<0<C??2OxHžj013ŠO0<C?”h<]01JN01.01hG--9:013Sg
-9.;01<=>H
•<E0-A3<‘O2-9m0<R4-9OF}03230<C??2Oz01.;4B‹3<k01?40<
9Ÿ01q0<C??2OB.=3‘OFBN0<R:b,R4-9OF}03o6Œ6780_•
x
i•
“
i•
:
`H{<C?
?2Ox“7D-<vRN<=>3Ch4vO180•
x
•
“
i0<C?x\•
x
•
x
i•
x
•
”
i0<C?“\•
“
•
“
i
•
“

•
”
i0<C?”\•
”
•
”
H
YZ
1. Tương tác át chế
‡•L7MeŒ\Œ
…Œ
1Dgh4vO180ss^D3344B}Ot23BN0<?DO7]01-901HQDO
-901zh4vO180ss7DR:180s-f:?DOJN1802-B4i30h4vO1803344
3<:h4vO<m0<-9017DR:180-f:?DOz-9f01-<24Bj01<=>7a0
\ 1D-901ì1D-901
ss 3344
d
e
s34_1D-901`
d
g
s\s44\33\e3344
e-901\?DO
L7Meg\\e
x780-9@4xhm?<?b,J4vO<4M03o6z7:3Obh<23H3<LJ4vO
<4M0z66Hx6JJ3Ch4vO<m0<h<23
V$YĂVÂÊ$ ÔƠ[[ƯƯVÂÊĐƠ
)Vă
e
[ƯƠ$ Đ*Ăă

e
^r40<6O3<:d
g
-7Meg
ĐYâ YêÔ
ƯVÂÊĐ ì ƯVÂÊĐ
d
e
\ [ƯôÂÊĐơ
d
g
\ x\xJJ\66\e66JJ
ê-ÂÊĐYâÂÊ YêÂÊÔ
[
\4#4"(#)]61#)^_#]0#)`#/?#a1b+#" !"#c&dedc#f0g#;<#)`#/?#cede#fhg
q
‡••7Mƒ\Œ\q
c4vO180663S0-9zb,J4vO<4M03o632367807:3Ob“i1®47D2-3<G
7a0Bn4^r47:3Ob“
•<A01<4M?\
•\ s<O@-B80×s<O@--9–01
 xxJJ 66““
d
e
\ x6“J_t2?0EO`
d
g
\ ƒx‡“‡\Œx‡JJ\Œ66“\e66JJ
 ƒt2?0EO\ŒB80\q-9–01
 !"#$%i##jkl!2#)54#5)#4"(#7_#Am!2#"8-#)9#2n!#/o!

2. Tương tác bổ sung
 ‡•L7Mƒ\Œ\Œ\e
’AR„b,R4-9OF}0<m0<Rf01?D:1D
•\1D?D:<:6<j01×<f-BPO
xxJJ 66““
…¯
d
e
\ x6“J_?D:‘OSC33<C`
d
g
\ ƒx‡“‡\Œx‡JJ\Œ66“‡\e66JJ
ƒ?D:‘OSC33<C\Œ<:6<j01\Œ<f-BPO\e?D:B/0
pKLHNIN

‡•L7Mƒ\°\e
•JAB€ig3a>6780t23BN0<<m0<Rf01‘OS\-9‰0iR•-^DRD4
•\-9‰0 × -9‰0
xxJJ 66““
d
e
x6“J_R•-`
d
g
ƒx‡“‡\Œx‡JJ\Œ66“‡\e66JJ
ƒR•-\°-9‰0\eRD4

…°
 !"#$%q##rs#" !"#)"<!"#454#7^!2#*+,#>#ht
‡•L7~ƒ\‚

uNHNIPvHLathyrus odoratus
•<A01<4M?zBPO-</?Lathyrus odoratus
x‡“‡\<:6B€
…‚
x‡JJi66“‡\<:6-9–01
• IPO-</?<:6-9–01×<:6-9–01
xxJJ 66““
d
e
\ x6“J_<:6B€`
d
g
\ ƒx‡“‡\Œx‡JJ\Œ66“‡\e66JJ
 ƒB€ \ ‚-9–01
3. Tương tác đa gen
6H•./01-233@011@>
s23-A0<-9f013<NOb,3<4><n43o60<4}O180B.=31®47Db,R4
-9OF}0B6180HQ±4180B}O3Ch4vO<m0<z?k3B@0<T-BN0<i0<4}O180B/0
0DF3C-23R„013@011@>-<8:?@-<.r01Hs2332-<v3CJ4vO<4M0h4vO<m0<
R6:B@01h<230<6OR:0<P00<4}O<6FA-180i3C-<vtG>3<y01-<8:?k3B@
J4vO<4M0-<D0<?@-R‹F74~0-„31®47Dbn7.=01H
•A0<-9f01R:0<4}O180t23BN0<i0~0J4vO<4M0h4vO<m0<3o63<y01
R²JNJ4G0BK4R:-23B@013o6?]4-9.;01H
JH••7M><E07Fzd
g
{³?eƒ†…0<DR4-9OF}0<®3•<„FI4v0{47b:0‡´<789h<401<4~03kOz7y6
?m\
•<G<Md
e
x

e
6
e
x
g
6
g
3C?DOB€<j01R:?601g6780-9@4x
e
^Dx
g
izd
g
\
x
e
x
e
x
g
x
g
‡‡‡‡‡‡‡‡‡‡‡š 6
e
6
e
6
g
6
g


I€BP? -9–01
•d
g
1456?DOB€BP?^D?DO-9–013C323?k3?DO-9O011460\B€
_Œx`iB€<j01_gx`^D<j01_x`
s<4}ORD4-924J–>^D?DO7]01t2?z3<O@-3µ017D0<501^AR„^}b,R4
-9OF}0B6180H
•01.;40<4}O-A0<-9f013Cb,R4-9OF}0bn7.=010<.-A0<-9f01
?DOR6i3<4}O36:HHH+,R4-9OF}0?DOR63C-<v74~0‘O60BG0Œ3a>180i
1456Rf01B80bP?x
e
x
e
x
g
x
g
x
Œ
x
Œ
^DRf01-9–016
e
6
e
6
g
6
g

6
Œ
6
Œ
3C323?DOR6
B80-9O011460H••7M><E07F1456R6?DO^DR6-9–017D°Œ\eH
……
4. Tính đa hiệu của gen:
{16F-¶-<;4Q80R87i]01B‹0<P0-<TFe1803C-<v-23B@01BG0
0<4}O-A0<-9f01H’AR„\zBPOžD760i180S0<<.z01BG0?DO<:6Bj01-<;4
S0<<.z013S?DO^€<f-i0<.<:6B€<f-t2?i3‰0<:6-9–01<f--9–01H
ž4M0 =01e180S0<<.z01BG00<4}O-A0<-9f011®47D-A0<B6<4MO3o6
180H
ž@43<k01Q69·60\JM0<R4-9OF}0180-9@4-9~0{+•-<.;01^r4J4vO
<4M0z?–-it./01^D<M-4??f3<H•801EF96<@43<k01Q69·60z-9~0
0<20<RD43o6{+•e¯i?‹<C63<:·4J94740_-<D0<><Š0?]74~0hG-`H“M0<
tOT-><2--¶-m0<-9f01-K3<k3?]74~0hG-JNhu:R‹0h<]01Jm0<-<.;01
1EF0<4}O<PO‘OSh<230<6OH{1.;4JM0<3C-6F3<E0RD4ih<O]0?a-<•>H
•01.;4ib64<€011803o6JM0<-<4GO?2O<j013ŠO<m0<74}?1EF96
<D017:f-3<k01JM0<h<23Hc<:S01g¯¸323JM0<R4-9OF}0b64<€013TO
-9y3-4?J¹?b40<3C-<vR|0BG0J4G0Rf013/t./01_ƒ¸`i<M-<Š0h40<
-9O01./01JT--<.;01_q¸`ib64<€01B.;01-4G-04MO<6F-<P0_¯¸`^D-4~O
<C6_q¸`H
•<,3-GJT-hº1800D:3µ013C-A0<B6<4MO^m?@-180h<]01A--<m
0<4}OB}O3CS0<<.z01BG0180h<23H{<5011803C<:f-B@01br?-9:01
‘O2-9m0<><2--94v032-<vb»3C-23B@010<4}O</0^D7EO</0H
wxyLxzIQ{|
1. Gen biến đổi (Modifier gene)
323180t23BN0<3C-A0<-9f01<6Fh<]013C-A0<-9f01B.=31®47D323180
3³0JS0<6F1800}0H•80J4G0BK47D0<501180h<]013CJ4vO<4M0h4vO

<m0<94~013o60C0<.017f43CS0<<.z01BG0b,J4vO<4M0h4vO<m0<3o6
0<5011803³0JS0h<23H
’AR„\•J‰i7]01B}OR:180-9@4+i3CBn?R:1807a0b‘OFBN0<H“‰3C
Bn?<6Fh<]013CBn?7DR:3231803³0JS0t23BN0<H{<.01zJ‰3CBn?i
Bn?A-<6F0<4}O7DR:-23B@013o6323180J4G0BK4H+nBn?z7]013<C
3µ013Cb,J4vO<4M0 /01-,\Bn?0<4}O<6FA-7DR:-23B@013o6180J4G0
BK4H
…ƒ
 !"#$%c}#~1b+#"1?!#4•0#2n!#h1(!#A€1#)]_!2#" !"#)"<!"#c}#/_^1#4"D#AC•
2. Các tính trạng bị giới hạn bởi giới tính
sC0<501-A0<-9f01?D1803o63<y013<LJ4vO<4M0z?@-14r4B.=3
1®47D180JN14r4<f0Jz414r4-A0<H•Bf4146by33Cb¶01bn7.=01bµ6^D
7.=01?¼-9:01b56B.=3J4vO<4M0z14n01324Hs233:0J‰B,3?601323
180t23BN0<7.=01b56^DB@?¼-9:01b56h<]013CJ4vO<4M00<.01323
1800DFB.=3-9OF}03<:J‰3243:0H›:BC-9:013<®014n01J‰b56><S4
-OFv0323J‰B,3?601323180-f:b56-n-BvB8?764H•1D-9n013µ01^PFi
3<y01?601323180Bl-9k010<4}O<6FA-i7r0<6F0<€0<.01h<]013C
J4vO<4M0H
3. Các tính trạng có sự biểu hiện phụ thuộc vào giới tính
sC0<501-A0<-9f01?Db,J4vO<4M0-9@4<6F7aX" 
-A0<H
Ġ
’AR„\zBf4146by33Cb¶01i1803Cb¶01\ž •80h<]01b¶01\<
c4vO180žž-f:b¶01z3S3:0B,37|03:0324ih4vO180<<h<]01
3Cb¶01z3Sg14r4Hc4vO180RN<=>-wž<0GOz3:0B,3b»3Cb¶01_ž7D
-9@4`i0GOz3:03247f4h<]013Cb¶01_ž7a0`H
•R~i-A0<-9f013C9EOtj?<6Fh<]013µ01-<O@37:f43CJ4vO<4M0
><„-<O@314r4-A0<i3:0B,3RN<=>-w3C9EOtj?iR~324RN<=>-wh<]01
9EOH
•01.;4<C4BŠOR:180“_J67708bb`t23BN0<Hc4vO180“J3CJ4vO

<4M0-9@4zBD0]01i0<.01J4vO<4M07a0z05HI4}O0DF14S4-<A3<^mb6:A-
3C><„05<C4BŠOH
 !"#$%cc##jt!"#"D1#A‚+#4D#@s#h1b+#"1?!#-"ƒ#)"+64#;<_#21„1#)t!"
{1C0-6F2>y-3o6JD0-6F01.;43C-<vRD4</0<6F01–0</001C0
-9€H{1.;4-63<:9Ÿ01b,14S?3<4}ORD401C0-9€R:?@-180-9@4z06?^D
7a0z05H
…†‡HJxzRMNPQ{Jˆxz‰‰IR
1. Độ thấm (độ thâm nhập)
I@-<T?7Dh<2404M?Bv3<L?k3B@-<6?1463o66780^D:h4vO
<m0<H
ƒe
AR\01.;43Cc4vO180
x

:
3C0<C??2Ox-<m
x
3CB@-<T?
eái30

-<má^mh<]01-<E?0<P>^D:h4vO<m0<H9.;01<=>01.;4
3C0<C??2Ox-<m3S
x
7|0

B}O3CB@-<T?eáH
{<4}O1803CB@-<T?h<230<6O-F7k6-OK4H{<50101.;4?601
1808>47:46_JM0<R4-9OF}0`3C01.;43<G-br?i3C01.;4bn01^D3C-<v
b40<3:0Hs23180h<]01-<T?^|03C-<v-9OF}03<:B;4b6O0<.3231803C
B@-<T?<:D0-:D0H

AR\
M0<x7ẵ<84?897D?9n47:f0-9A0<ri?T-BN0<<.r01h<]011460^D
-<;41460i3CJ4vO<4M0z01.;414Db6O-OK4H4M006FBt23BN0<B.=3
W$""ắ ê#%** %!ặ
ầ$"#%Ơ!"ầ$'ẩẫấ$ặ*
{<4}O0<E0^P-0K4-4G01?3JM0<x7ẵ<84?890<.3,O-K01-<n01QậèH
è86160H
M0<O0-401-:07DJM0<1EF96R:9n47:f0-<0h40<^r4J4vO<4M0
?T--9A_R8?80-46`^D146-01323^P0B@01h<]01h4v?b:2-B.=33o6323
3<4R|0BG0<4M0 =01?y6^;0_3<:986`0~0B]4h<4JM0<30B.=31đ47D
JM0<?y6^;0O0-401-:0H94MO3<k013o6JM0<-<.;01h<]01J4vO<4M0
-9.r3-OK4H
7M-<T?-<.;01B.=3B20<142R,6-9~0^4M3t8?tu-?@-bn
7.=017r0146Bm0<^Dt23BN0<-7M01.;4J-JO@3?601180^D0<501
%!ẻƠ*ẽé%
0DFA-</0eá-60C47D3C-A0<-<T?14S?<6F-A0<-<T?h<]01<:D0-:D0
_403:?>76-8>808-96038`H
2. hin hay biu hin
I@J4vO<4M0B.=3R01Bv3<L?k3B@0<4}OA-3o6-A0<-9f01h<4B
-<T?<:D0-:D0H
AR\b,3S?0<P0^NB013o63<T-><80F7-<4:369J6?4R8_s`<6Fh<]01
3S?0<P0z01.;4R:e180t23BN0<HOF0<4~00<50101.;43S?0<P0^N
B013CB@<4M0h<230<6O\3C01.;43S?0<P0^NB01z0j01B@eH
?1p7<:a336:</0i-9:01h<432J4M-3C0<50101.;43S?0<P0^NB01z
0j01B@9T--<T>7Die?1p7i0<4}O01.;4h<233S?0<P0^NB201z323
0j01B@-9O011460H
g
{<4}O-A0<-9f01z01.;43CB@<4M0K0BN0<bOn-R;40<.0<C??2Oi
?DO?–-HHH
‡“4vO<4M0B6Rf01_’6946J78´t>98bb4:0`

Q@--m0<-9f01><k3-f>h<237Db,J4vO<4M0B6Rf013o6JM0<H“M0<
3C-<v3C-A0<-<T?<:D0-:D00<.01?k3B@01<4M?-9®013o6JM0<3Cb,
-<6FBK49T-7r0H
’AR„\“M0<Ot/-<Š0h40<_08O9:·4J9:?6-:b4b`-F>8•_3‰01®47DJM0<’:0
Ì83h7401<6Ob80`H“n<:a3?•3C-<v3CJM0<9T-0<•i0<•BG00±4<®h<]01
01<Ñ7D?m0<?–3JM0<H{<.01<®3C-<v-9OF}0180JM0<3<:3:0^DB]4h<4
3<y013CJ4vO<4M0JM0<9T-0a01H“4vO<4M03o6JM0<7DJM0<0<E03C323
3<T?3D><~‡b56-9~0J„01^D323Ot/-<Š0h40<Jm<6F323Ot/-<Š0h40<
z><A67.01H
sSgh<2404M?B@-<T?^DB@J4vO<4M0B.=30<DR4-9OF}0<®3
01.;4{16•4?:·88^‡Ì8b:>bh40~O96^D:0³?eƒg¯BvB20<142?k3B@
-<v<4M096h4vO<m0<3o6323180H
{1:D496ib,J4vO<4M03o6h4vO18096h4vO<m0<3‰0><„-<O@3^D:
?k3><S0k01H
…[PFHZzX
c4vO1803‰03<NO0<4}O-23B@01h<230<6O3o6?]4-9.;01J~0
-9:01^D01:D43/-<v
1. Tác động của môi trường bên ngoài
6H{<4M-B@
•23B@013³0JS03o60<4}O1803<oFGO7Dh4v?b:2--n3B@><S0
k01b40<<C6-<]01‘O680½F?87D0<5013<T-B.=3323180t23BN0<^}?a-
R4-9OF}0H•4560<4M-B@^D-n3B@><S0k013Cb,74~0‘O603<a-3<»H{<4M-
B@S0<<.z01BG0b,J4vO<4M0h4vO<m0<-9:010<4}O-9.;01<=>H
’AR„\b,J4vO<4M0-A0<-9f017]01B80z3<C>?µ4i-64^D3<E03o614n01
-<€ž4?676F6z0<4M-B@-<T>h<4><2--94v0J@7]01H
{<4M-B@3C-<vS0<<.z01BG0B@-<T?^DB@<4M0H
JH›40<R.¼01
ƒŒ
•9:01?@-bn-9.;01<=>3<GB@R40<R.¼01S0<<.z01BG0J4vO
<4M0h4vO<m0<H’AR„b,J4vO<4M0?¼^D013o6-<€R:gFGO-n\b,<4M0R4M0

3o6180Bj01<=>-w7a0FF^D7.=01-<,3^P-t60<_t60-<:><F77`-9:01-<k3
³0H{GO-<4GO-<,3^P-t60<-9:01-<k3³0i?¼^D01h<]01tOT-<4M0H
“M0<><80F7h8-:0O946_•cÒ`B.=31a>z01.;4R6-9–01^r4-•7M
epe†H†††7Š0b40<Hs23B@-J4G0z7:3Ob?‹<C63<:80½F?83<OFv0<C6
><80F76760408<FR9:tF76b87D?3<:01.;4?601h4vO180Bj01<=>h<]01
-<v3<OFv0<C66?40:634R><80F76760408H•9:01h<40<501-9lb/b40<?–3
JM0<><80F7h8-:004MO3CJ4vO<4M0Jm0<-<.;01b6Ob40<-<mb,h<4G?
h<OFG-3<OFv0<C6b»7D?-A3<7µFRŠ0><80F76760408^D323bS0><¹?
3<OFv0<C6B@3h<23R|0BG0-K0-<./01<M-<Š0h40<-9O01./01^D1EF96
J4vO<4M03<P?><2--94v0-9A-OM-9Š?-9®01H{<.^PFh4vO180•cÒb»1EF
96h4vO<m0<JM0<7Ó01<4~?-9®01H•OF0<4~00GO-9l?601h4vO180JM0<
B.=3><2-<4M0br?‘O6bD017®3-9.r3b40<iJ4vO<4M03o6JM0<3C-<v-920<
JŸ013<GB@³001<Ô:><80F76760408H{<.^PFi?a3RŠO-9l?601h4vO180
•cÒ0<.010<;3<GB@³0><Ž<=>B‹-920<B.=3-m0<-9f013<P?-9AH
3HÕ0<<.z013o63/-<v?•
+6Oh<4-9k01B‹B.=3-<„-40<i3/-<v?•3C-<vS0<<.z01BG0b,><2-
-94v0H’AR„?2O01.;4?•3Ch4vO1809<
‡
_0<E0-n9<8bObE?`i0GOBk63:0
-<k0<T-3CÌ<
™
b40<96b»h<]01b6:i0<.01Bk63:0-<k<643C-<vJN3<G-H
2. Tác động của môi trường bên trong
œO2-9m0<><2--94v032-<v-9S4‘O60<4}OJ.r3-9O011460><k3-f>H
{16F-9:013/-<v3C0<4}O-23B@0114563233TO-9y3h<230<6O^D^r4h4vO
1800<.323 /01-23\180^r4180i180^r4{+•i{+•‡0<E0i0<E0‡-GJD:
3<T-i-GJD:‡?]HHHzBEF3<L0~O^D4-23B@013C-A0<3<T-3<O01-K01‘O2-\
6H•OK4
{<4}O-A0<-9f01^DJM0<R4-9OF}0z01.;43CJ4vO<4M0-9:01?@-B@
-OK40<T-BN0<H“M0<6736>-:0O946_0.r3-4vO3C634R<:?:180-4b43JNB80h<4

3C”
g
`J4vO<4M0016F7y3?r4b40<96H“M0<^SF32J4vO<4M0-9:01q-<201
BŠOH“M0<-4vOB.;01^D3<k013:14P-žO0-401-:0J4vO<4M0z0<4}OB@-OK4
h<230<6OH“M0<x7½<84?89J4vO<4M0b6O°†-OK4H
JH•4r4-A0<
ƒq
 •4r4-A0<3C0<4}OS0<<.z01BG0b,J4vO<4M03o61800<.14r4<f0
b,J4vO<4M0<6F-A0<-9@47a0><„-<O@3^D:14.C4-A0<H{1:D4963C0<501
18074~0hG-^r414r4-A0<HsC-<v<:9?:08b40<R„3-23B@01BG0J4vO<4M0
3o6323180H
{<.^PF?]4-9.;01J~0-9:01^D01:D43/-<v3C0<4}OS0<<.z01
><k3-f>h<230<6O7~0b,J4vO<4M0h4vO<m0<3o6323180H
{<4}OJM0<z01.;40<.-4??f3<i<OFG-2>HHH-9.r3BEFh<]013:4
7D323JM0<R4-9OF}0HœO60B4v??r4<4M006F3<:9Ÿ01BC7D0<501JM0<R4
-9OF}0?DJ4vO<4M0><„-<O@3?]4-9.;01H•9~0-<,3-G32318074~0‘O60
BG0JM0<0DFB.=3-9OF}0-<„-9:01323146Bm0<3CJM0<H
$%&'(
eH•<E0J4M-1806780^D180h<]016780
gH{~Ob,h<230<6O14562-3<G180-9@4^D2-3<G1807a0
ŒHÖ23BN0<JS03<T-R4-9OF}03o6b,t23BN0<323Rf01?D:1DH
qHHc<401<4~03kO<4M0 =01R4-9OF}0i3230<DR4-9OF}0<®3B‹><2-
<4M0-•7Mƒ\°\e-9:01-<G<M3:0Hž‹F14S4-<A3<hG-‘OSR4-9OF}00DFHsC
-<vh4v?-9614S-<OFG-0DF0<.-<G0D:H
¯H•<E0J4M-B6180^DB6<4MO3o6180H
)*+,-./0
•<f?•<D0<žK_g†††`H›4-9OF}0<®3H{Ö“•42:›„3H
ˆ~Im0<ˆ./01i•<60s,{<E0_eƒƒ…`Hs/bzR4-9OF}0<®3H{Ö“•42:
›„3H
ž:D01•9®01•<20_eƒƒ¯`H›4-9OF}0<®3><E0-wH•9O01-E?ID:-f:

•¶t6iIf4<®3žOGH
x0-<:0F  ×H  dH  •94··4-<bi  +Ob60  ÌH  ˜8bb789i  Ì43<69R  sH  ˆ8—:0-40i
˜47746?  QH  •87J69-i  ›6^4R  •H  +O½Oh4i  ×8··98F  žH  Q47789H  g††qH  x0
40-9:RO3-4:0-:1808-43b6067Fb4bH˜HžHd988?60•OJ74b<89bH
ž697-›HˆHi×:08b´H˜H_eƒƒ…`H•808-43b‡•94034>7860R6067Fb4bH×:08
60R“69-78 •OJ7b<89bH•:9:0-:is606R6H
ƒ¯
+-60b·487R˜H›HeƒƒeH+3<6O?Øb:O-7408:·-<8:9F60R>9:J78?b:·
1808-43bHQ3•96—‡ž477i•03Hi{8—Ù:9hH
ƒ°