K. S. Sodiqova Mirzo Ulug’bek nomidagi O’zbekiston

Noallel genlar o’zaro ta’sirining kombinirlangan tipi.
O’zbekiston Respublikasi Fanlar Akademiyasining akademigi D.A.Musayev o’z shogirdlari bilan qariyb 50 yil g’o’za genetikasi ustida olib borgan ilmiy tadqiqotlari natijasida dunyoda birinchi bo’lib G.hirsutum L. turiga mansub g’o’zaning genetik kolleksiyasi liniyalarida chigit ustidagi tola qoplami (tuk va tola) ning irsiylanishi haqidagi ilmiy nazariyani yaratdi. Bu nazariyaga ko’ra, chigit ustidagi tuklanishning irsiylanishi uch guruh genlarga bog’liq:
1. Gen ingibitor – I – i. Bu genning dominant allellari gomo va geterozigota holatlarda chigit tuklanishining asosiy va qo’shimcha genlari faoliyatini to’xtatib qo’yadi, natijada chigit tuksiz – yalang’och urug’li bo’ladi.
2. CHigit tuklanishining asosiy genlari. Ular asosan ikki juft – Ft₁-ft₁ va Ft₂ - ft₂ bo’lib, gen-ingibitor retsessiv gomozigota (ii) bo’lgan vaqtda chigitning mikropile qismidagi tuklanishni rivojlantiradi. Ular kumulyativ polimeriya tipida faoliyat ko’rsatadi.
3. CHigit tuklanishining qo’shimcha geni - Fc – fc. Bu gen genotipda gen-ingibitor retsessiv gomozigota holatida bo’lganda, asosiy genlarning birinchisi - Ft₁ – ft₁ bilan komplementar ta’sirda bo’lib, chigitning xalaza va yon tomonlaridagi tuklanishni rivojlantiradi.
Demak, chigit tuklanishining irsiylanishi kamida 4 juft genga bog’liqligini ko’ramiz.
Akademik D.A.Musayev olib borilgan tadqiqotlar natijasiga suyangan holda umumiy genetika faniga birinchi bo’lib noallel genlar o’zaro ta’sirining yangi tipi – kombinirlangan tipni kiritdi. Bu tipning asosiy xususiyati shundaki, bitta belgining – chigit ustidagi tuklanishning irsiylanishida bir vaqtning o’zida noallel genlar o’zaro ta’sirining barcha tiplari – komplementariya, epistaz, polimeriya, asosiy va qo’shimcha genlarning o’zaro ta’siri kabilar ishtirok etadi. Bu dunyo fanidagi buyuk kashfiyotlardan biri hisoblanadi.
Dastlab chigit tuklanishining irsiylanishida ishtirok etuvchi asosiy genlar – Ft₁ - ft₁ va Ft₂ – ft₂ ning faoliyatini ko’rib chiqaylik. Buning uchun genetik kolleksiyaning L-15 va L-110 liniyalarini olamiz. L-15 liniya o’simliklari normal mikropilyar tuklanishga ega, L-110 liniya o’simliklari esa retsessiv yalang’och urug’li. Bu liniyalarni o’zaro chatishtiramiz:
n – MS rGS
P ♀ L - 15 X ♂ L - 110
ii Ft₁Ft₁Ft₂Ft₂ fcfc ii ft₁ft₁ft₂ft₂ fcfc
g i Ft₁Ft₂ fc i ft₁ft₂ fc
F₁ ii Ft₁ft₁Ft₂ft₂ fcfc – m-MS (o’rtacha mikropilyar tuklanish)
m-MS m-MS

P ♀ ii Ft₁Ft₁Ft₂Ft₂ fcfc X ♂ ii Ft₁Ft₁Ft₂Ft₂ fcfc

g i Ft₁Ft₂ fc i Ft₁Ft₂ fc
i Ft₁ft₂ fc i Ft₁ft₂ fc
i ft₁Ft₂ fc i ft₁Ft₂ fc
i ft₁ft₂ fc i ft₁ft₂ fc
F ₂ 1. ii Ft₁Ft₁Ft₂Ft₂ fcfc – 1 1 n-MS normal mikropilyar tuklanish
2. ii Ft₁Ft₁Ft₂ft₂ fcfc – 2
3. ii Ft₁ft₁Ft₂Ft₂ fcfc – 2 4 p-MS oraliq mikropilyar tuklanish
4. ii Ft₁Ft₁ft₂ft₂ fcfc – 1
5. ii ft₁ft₁Ft₂Ft₂ fcfc – 1 6 m-MS o’rtacha mikropilyar tuklanish
6. ii Ft₁ft₁Ft₂ft₂ fcfc – 4
7. ii Ft₁ft₁ft₂ft₂ fcfc – 2 4 nz-MS juda kichik mikropilyar tuklanish
8. ii ft₁ft₁Ft₂ft₂ fcfc – 2
9. ii ft₁ft₁ft₂ft₂ fcfc – 1 1 rGS retsessiv yalang’och urug’li
SHunday qilib, F₂ da ikkita fenotipik sinf kuzatiladi:
- mikropilyar tuklanishli o’simliklar
- retsessiv yalang’och urug’li o’simliklar
Ularning nisbati 15:1.
CHigit tuklanishining asosiy genlari – Ft₁-ft₁ va Ft₂-ft₂ digenli polimeriya tipida faoliyat ko’rsatadi. Ularning ta’siri kumulyativ polimeriya tipida bo’ladi, ya’ni tuklanish dominant allellarning soniga bog’liq:
4 ta dominant allel (Ft₁Ft₁Ft₂Ft₂) normal mikropilyar (n-MS) tuklanishni beradi;
3 ta dominant allel (Ft₁Ft₁Ft₂ft₂, Ft₁ft₁Ft₂Ft₂) oraliq mikropilyar (p-MS) tuklanishni beradi;
2 ta dominant allel (Ft₁Ft₁ft₂ft₂, ft₁ft₁Ft₂Ft₂, Ft₁ft₁Ft₂ft₂) o’rtacha mikropilyar (m-MS) tuklanishni beradi;
1 ta dominant allel (Ft₁ft₁ft₂ft₂, ft₁ft₁Ft₂ft₂) juda kichik mikropilyar (nz-MS) tuklanishni beradi;
hamma retsessiv allellar (ft₁ft₁ft₂ft₂) retsessiv yalang’och urug’ni (rGS) beradi.
Asosiy genlar faoliyat ko’rsatishlari uchun gen ingibitor retsessiv gomozigota (ii) holatda bo’lishi lozim.
Endi gen ingibitor ishtirok etgan liniyalarni o’zaro chatishtirib ko’ramiz:
dominant absolyut normal mikropilyar
yalang’och urug’li tuklanish
DAGS n-MS
P ♀ L - 70 X ♂ L - 15
II ft₁ft₁ft₂ft₂ fcfc ii Ft₁Ft₁Ft₂Ft₂ fcfc
g I ft₁ft₂ fc i Ft₁Ft₂ fc
F₁ Ii Ft₁ft₁Ft₂ft₂ fcfc – yalang’och urug’li GS
GS GS
P ♀ Ii Ft₁ft₁Ft₂ft₂ fcfc X ♂ Ii Ft₁ft₁Ft₂ft₂ fcfc

F₁ duragaylari trigeterozigota bo’lganligi sababli, ota-onaning har biri

sakkiztadan gameta hosil qiladi:
g I Ft₁Ft₂ fc I Ft₁Ft₂ fc

I Ft₁ft₂ fc I Ft₁ft₂ fc

I ft₁Ft₂ fc I ft₁Ft₂ fc
I ft₁ft₂ fc I ft₁ft₂ fc
i Ft₁Ft₂ fc i Ft₁Ft₂ fc
i Ft₁ft₂ fc i Ft₁ft₂ fc
i ft₁Ft₂ fc i ft₁Ft₂ fc
i ft₁ft₂ fc i ft₁ft₂ fc
F ₂ 1. II Ft₁Ft₁Ft₂Ft₂ fcfc – 1 DAGS

2. II Ft₁Ft₁Ft₂ft₂ fcfc – 2 DAGS

3. II Ft₁ft₁Ft₂Ft₂ fcfc – 2 DAGS
4. II Ft₁Ft₁ft₂ft₂ fcfc – 1 DAGS
5. II ft₁ft₁Ft₂Ft₂ fcfc – 1 DAGS 16 DAGS
6. II Ft₁ft₁Ft₂ft₂ fcfc – 4 DAGS
7. II Ft₁ft₁ft₂ft₂ fcfc – 2 DAGS
8. II ft₁ft₁Ft₂ft₂ fcfc – 2 DAGS
9. II ft₁ft₁ft₂ft₂ fcfc – 1 DAGS
10. Ii Ft₁Ft₁Ft₂Ft₂ fcfc – 2 DAGS
11. Ii Ft₁Ft₁Ft₂ft₂ fcfc – 4 DAGS
12. Ii Ft₁ft₁Ft₂Ft₂ fcfc – 4 DAGS
13. Ii Ft₁Ft₁ft₂ft₂ fcfc – 2 DAGS
14. Ii ft₁ft₁Ft₂Ft₂ fcfc – 2 DAGS 32 DAGS
15. Ii Ft₁ft₁Ft₂ft₂ fcfc – 8 DAGS
16. Ii Ft₁ft₁ft₂ft₂ fcfc – 4 DAGS
17. Ii ft₁ft₁Ft₂ft₂ fcfc – 4 DAGS
18. Ii ft₁ft₁ft₂ft₂ fcfc – 2 DAGS
19. ii Ft₁Ft₁Ft₂Ft₂ fcfc – 1 n-MS 1 n-MS
20. ii Ft₁Ft₁Ft₂ft₂ fcfc – 2 p-MS 4 p-MS
21. ii Ft₁ft₁Ft₂Ft₂ fcfc – 2 p-MS
2 2. ii Ft₁Ft₁ft₂ft₂ fcfc – 1 m-MS
23. ii ft₁ft₁Ft₂Ft₂ fcfc – 1 m-MS 6 m-MS
24. ii Ft₁ft₁Ft₂ft₂ fcfc – 4 m-MS
25. ii Ft₁ft₁ft₂ft₂ fcfc – 2 nz-MS 4 nz-MS
26. ii ft₁ft₁Ft₂ft₂ fcfc – 2 nz-MS
27. Ii ft₁ft₁ft₂ft₂ fcfc – 1 rGS 1 rGS
48 DAGS + 1 rGS = 49 GS (yalang’och urug’li)

1 n-MS + 4 p-MS + 6 m-MS + 4 nz-MS = 15 MS (mikropilyar tuklanish)

F₂ da ikkita fenotipik sinf hosil bo’ladi:
- yalang’och urug’li (chigitli) o’simliklar
- mikropilyar tuklanishli o’simliklar
Ularning nisbati 49 : 15.
Gen ingibitor genotipda dominant gomo- va geterozigota ( II va Ii ) holatlarda ishtirok etgan taqdirda, fenotipda yalang’och urug’ (GS) hosil bo’ladi, ular dominant yalang’och urug’lilar hisoblanadi. Genotipda barcha genlar retsessiv gomozigota ( ii ft₁ft₁ft₂ft₂ fcfc ) holatda bo’lsa, u holda retsessiv yalang’och urug’ hosil bo’ladi.
Nihoyat, noallel genlar o’zaro ta’sirining komplementar tipida chigit tuklanishining irsiylanishini ko’rib chiqamiz:
to’liq tuk bilan normal mikropilyar
qoplangan tuklanish
OS n-MS
P ♀ L - 47 X ♂ L - 15
ii Ft₁Ft₁Ft₂Ft₂ FcFc ii Ft₁Ft₁Ft₂Ft₂ fcfc
g i Ft₁Ft₂ Fc i Ft₁Ft₂ fc
F₁ ii Ft₁Ft₁Ft₂Ft₂ Fcfc – chigitning mikropile qismida normal tuklanish,
chigitning xalaza qismida tuklanishning notekis taqsimlanishi PS
PS PS
P ♀ ii Ft₁Ft₁Ft₂Ft₂ Fcfc X ♂ ii Ft₁Ft₁Ft₂Ft₂ Fcfc

g i Ft₁Ft₂ Fc i Ft₁Ft₂ Fc

i Ft₁Ft₂ fc i Ft₁Ft₂ fc
F₂ ii Ft₁Ft₁Ft₂Ft₂ FcFc –1 OS
ii Ft₁Ft₁Ft₂Ft₂ Fcfc – 1 PS
ii Ft₁Ft₁Ft₂Ft₂ Fcfc – 1 PS
ii Ft₁Ft₁Ft₂Ft₂ fcfc – 1 n-MS
F₂ da uchta fenotipik sinflar hosil bo’lib, ularning nisbati 1 : 2 : 1.
Endi yuqorida ko’rib o’tilgan noallel genlar o’zaro ta’siri tiplarining bir kombinatsiyada chigit tuklanishining irsiylanishiga ko’rsatadigan ta’sirlari ustida to’xtalamiz:
DAGS OS
P ♀ L - 70 X ♂ L - 47

II ft₁ft₁ft₂ft₂ fcfc ii Ft₁Ft₁Ft₂Ft₂ FcFc

g I ft₁ft₂ fc i Ft₁Ft₂ Fc
F₁ Ii Ft₁ft₁Ft₂ft₂ Fcfc – GS
GS GS
P ♀ Ii Ft₁ft₁Ft₂ft₂ Fcfc X ♂ Ii Ft₁ft₁Ft₂ft₂ Fcfc
g I Ft₁Ft₂ Fc I Ft₁Ft₂ Fc
I Ft₁Ft₂ fc I Ft₁Ft₂ fc
I Ft₁ft₂ Fc I Ft₁ft₂ Fc
I Ft₁ft₂ fc I Ft₁ft₂ fc
I ft₁Ft₂ Fc I ft₁Ft₂ Fc
I ft₁Ft₂ fc I ft₁Ft₂ fc
I ft₁ft₂ Fc I ft₁ft₂ Fc
I ft₁ft₂ fc I ft₁ft₂ fc
i Ft₁Ft₂ Fc i Ft₁Ft₂ Fc
i Ft₁Ft₂ fc i Ft₁Ft₂ fc
i Ft₁ft₂ Fc i Ft₁ft₂ Fc
i Ft₁ft₂ fc i Ft₁ft₂ fc
i ft₁Ft₂ Fc i ft₁Ft₂ Fc
i ft₁Ft₂ fc i ft₁Ft₂ fc
i ft₁ft₂ Fc i ft₁ft₂ Fc
i ft₁ft₂ fc i ft₁ft₂ fc
F ₂ 1. II Ft₁Ft₁Ft₂Ft₂ FcFc – 1

2. II Ft₁Ft₁Ft₂ft₂ FcFc – 2

3. II Ft₁ft₁Ft₂Ft₂ FcFc – 2
4. II Ft₁Ft₁ft₂ft₂ FcFc – 1
5. II ft₁ft₁Ft₂Ft₂ FcFc – 1 16 Dominant GS
6. II Ft₁ft₁Ft₂ft₂ FcFc – 4
7. II Ft₁ft₁ft₂ft₂ FcFc – 2
8. II ft₁ft₁Ft₂ft₂ FcFc – 2
9. II ft₁ft₁ft₂ft₂ FcFc – 1
10. II Ft₁Ft₁Ft₂Ft₂ Fcfc – 2
11. II Ft₁Ft₁Ft₂ft₂ Fcfc – 4
12. II Ft₁ft₁Ft₂Ft₂ Fcfc – 4
13. II Ft₁Ft₁ft₂ft₂ Fcfc – 2
14. II ft₁ft₁Ft₂Ft₂ Fcfc – 2 32 Dominant GS
15. II Ft₁ft₁Ft₂ft₂ Fcfc – 8
16. II Ft₁ft₁ft₂ft₂ Fcfc – 4
17. II ft₁ft₁Ft₂ft₂ Fcfc – 4
18. II ft₁ft₁ft₂ft₂ Fcfc – 2
19. II Ft₁Ft₁Ft₂Ft₂ fcfc – 1
20. II Ft₁Ft₁Ft₂ft₂ fcfc – 2
21. II Ft₁ft₁Ft₂Ft₂ fcfc – 2
22. II Ft₁Ft₁ft₂ft₂ fcfc – 1
23. II ft₁ft₁Ft₂Ft₂ fcfc – 1 16 Dominant GS
24. II Ft₁ft₁Ft₂ft₂ fcfc – 4
25. II Ft₁ft₁ft₂ft₂ fcfc – 2
26. II ft₁ft₁Ft₂ft₂ fcfc – 2
27. II ft₁ft₁ft₂ft₂ fcfc – 1
28. Ii Ft₁Ft₁Ft₂Ft₂ FcFc– 2
29. Ii Ft₁Ft₁Ft₂ft₂ FcFc – 4
30. Ii Ft₁ft₁Ft₂Ft₂ FcFc – 4
31. Ii Ft₁Ft₁ft₂ft₂ FcFc – 2
32. Ii ft₁ft₁Ft₂Ft₂ FcFc – 2 32 Dominant GS
33. Ii Ft₁ft₁Ft₂ft₂ FcFc – 8
34. Ii Ft₁ft₁ft₂ft₂ FcFc – 4
35. Ii ft₁ft₁Ft₂ft₂ FcFc – 4
36. II ft₁ft₁ft₂ft₂ FcFc – 2

37. Ii Ft₁Ft₁Ft₂Ft₂ fcfc – 2

38. Ii Ft₁Ft₁Ft₂ft₂ fcfc – 4
39. Ii Ft₁ft₁Ft₂Ft₂ fcfc – 4
40. Ii Ft₁ft₁Ft₂ft₂ fcfc – 8
41. Ii Ft₁Ft₁ft₂ft₂ fcfc – 2 32 Dominant GS
42. Ii ft₁ft₁Ft₂Ft₂ fcfc – 2
43. Ii Ft₁ft₁ft₂ft₂ fcfc – 4
44. Ii ft₁ft₁Ft₂ft₂ fcfc – 4
45. Ii ft₁ft₁ft₂ft₂ fcfc – 2
46. Ii Ft₁Ft₁Ft₂Ft₂ Fcfc – 4
47. Ii Ft₁Ft₁Ft₂ft₂ Fcfc – 8
4 8. Ii Ft₁ft₁Ft₂Ft₂ Fcfc – 8
49. Ii Ft₁Ft₁ft₂ft₂ Fcfc – 4
50. Ii ft₁ft₁Ft₂Ft₂ Fcfc – 4 64 Dominant GS
51. Ii Ft₁ft₁Ft₂ft₂ Fcfc – 16
52. Ii Ft₁ft₁ft₂ft₂ Fcfc – 8
53. Ii ft₁ft₁Ft₂ft₂ Fcfc – 8
54. Ii ft₁ft₁ft₂ft₂ Fcfc – 4
55. ii Ft₁Ft₁Ft₂Ft₂ FcFc – 1 OS
56. ii Ft₁Ft₁Ft₂ft₂ FcFc – 2 OS
57. ii Ft₁ft₁Ft₂Ft₂ FcFc – 2 OS
58. ii Ft₁Ft₁ft₂ft₂ FcFc – 1 OS
59. ii ft₁ft₁Ft₂Ft₂ FcFc – 1 m-MS
60. ii Ft₁ft₁Ft₂ft₂ FcFc – 4 OS
61. ii Ft₁ft₁ft₂ft₂ FcFc – 2 nz-MS
62. ii ft₁ft₁Ft₂ft₂ FcFc – 2 nz-MS
63. ii ft₁ft₁ft₂ft₂ FcFc – 1 rGS
64. ii Ft₁Ft₁Ft₂Ft₂ Fcfc – 2 PS
65. ii Ft₁Ft₁Ft₂ft₂ Fcfc – 4 PS
66. ii Ft₁ft₁Ft₂Ft₂ Fcfc – 4 PS
67. ii Ft₁Ft₁ft₂ft₂ Fcfc – 2 PS
68. ii ft₁ft₁Ft₂Ft₂ Fcfc – 2 m-MS
69. ii Ft₁ft₁Ft₂ft₂ Fcfc – 8 m-MS
70. ii Ft₁ft₁ft₂ft₂ Fcfc – 4 nz-MS
71. ii ft₁ft₁Ft₂ft₂ Fcfc – 4 nz-MS
72. ii ft₁ft₁ft₂ft₂ Fcfc – 2 rGS
73. ii Ft₁Ft₁Ft₂Ft₂ fcfc – 1 n-MS
74. ii Ft₁Ft₁Ft₂ft₂ fcfc – 2 p-MS
75. ii Ft₁ft₁Ft₂Ft₂ fcfc – 2 p-MS
76. ii Ft₁Ft₁ft₂ft₂ fcfc – 1 m-MS
77. ii ft₁ft₁Ft₂Ft₂ fcfc – 1 m-MS
78. ii Ft₁ft₁Ft₂ft₂ fcfc – 4 m-MS
79. ii Ft₁ft₁ft₂ft₂ fcfc – 2 nz-MS
80. ii ft₁ft₁Ft₂ft₂ fcfc – 2 nz-MS
81. ii ft₁ft₁ft₂ft₂ fcfc – 1 rGS
192 ta dominant GS + 4 ta retsessiv GS = 196 ta yalang’och urug’li (chigitli) – GS o’simliklar;
16 ta nz-MS + 17 ta m-MS + 4 ta p-MS + 1 ta n-MS = 38 ta mikropilyar tuklanishli – MS o’simliklar;
12 ta PS va 10 ta OS tipli o’simliklar.
F₂ da to’rtta fenotipik sinflar hosil bo’lib, ularning nisbati 196:38:12:10.
SHunday qilib, chigit ustidagi tuklanishning irsiylanishida 4 juft noallel genlar ( I-i, Ft₁-ft₁, Ft₂-ft₂ va Fc-fc ) ishtirok etib, ularning faoliyatida bir vaqtning o’zida noallel genlar o’zaro ta’sirining komplementar, epistaz, polimeriya tiplari, shuningdek, asosiy genlar bilan qo’shimcha gen o’rtasidagi o’ziga xos ta’sir tipining mavjudligini ko’ramiz.
Asosiy Ft₁-ft₁ gen Fc bilan komplementar ta’sir tipida bo’lsa, Ft₂-ft₂ esa qo’shimcha gen bilan komplementar munosabatda emas.
ii Ft₁Ft₁ft₂ft₂ FcFc - OS tipli tuklanish
ii ft₁ft₁Ft₂Ft₂ FcFc - m-MS tipli tuklanish
ii Ft₁Ft₁ft₂ft₂ Fcfc - PS tipli tuklanish
ii ft₁ft₁Ft₂Ft₂Fcfc - m-MS tipli tuklanish.