М???яр?е ??? э?лю?и, ???????и ???ия и ???ия
И???о, ?о ????е ????ы ДНК ??т ?????ся с ??? м?? ? ??? в ???? ??й ????ы
?и ?????ся в ??ую ????у.
С?????? ??аю?х ??в ???е ?? ???? Б. М?-К??? ?и ???? ???? ????. О? ?я??а,
?о э???ы ??лят??? ?? ????ю?я в ??? с ??го ??а ? ??? и ?ияют ? эк???ию
??в, ???я к ?я??ию ????ч??х ????. Э? ??? ? ????? в???я в ???е 30 ?т, ??
?????ия ? E. Coli ? ?????и ?????ь ??в к ?????ию и ????ю в ??? ??о в ???,
а Б. М?-К??? ?? ????? Н????ая ??ия.
С???? к ?????ю ????????? ДНК ???? ???? ?????ы (Tnэ???ы) ?и ??аю?е ??.
И???? ????я ???-? ???? ДНК ?????? ???ю?х ?о ?????????й ????ят ?в???? в
??? ????х, ??? ?????о ? ?х ??ю??ся ?????? ??о. М??е ???? ?????и ????ят в
????? ??л???й ????а?и ??у ????? ?с???????я? и ? ????ают ???? ????в ДНК.
В??е ? ??ря, ????а ??? ??? ????а. Э? и ? ??т ?? ??е, ????у н????о ??????
???????? ??? и ??в.
М??е ???? ?????и ? ?????и???, ?и ? ?я?? с ??м-? ??????м ?ия?? ? э????ю
??в и в ?х ?? э??? ?????и. С????и ??т ?? ???а ??х ???й, ?? ???? ДНК ?и ?
и ???. П???? ??х ???? ??? ???й ??? ?????ия ?????????й ДНК ? ??? ????о
??? в ??? ?и ????ия и ??е?ю?я ?????ия ????в ДНК. О??о ???е ?????? и
?????? ??т ?? ??я?? ??е с ??????, ??????о???и ????я?. Т?ие ?????????е
???я ??ют ?ю??ю ?? в ??ля?и э????и ????х ??в ? ??я ???????и и ???ия
?????? ??в ???.
Т?ы ????х э????
Р???ют ????о ??в ????х ДНК-э????, ? ?е ?и ???ют ??ую?? ??? ?????:
– ??т ?н (?и ????о ??в), ?????й ?я ?????? (?н ???о?? – ???? ???ую?? в
?????и);
– ?-???, ? ??? ???? ????е?? ???о ???????е ???яю??я ?????????, ??е ?????е
?я ??????
– ?? ?????ую??я э???ы ? ???ют ???х ?????? ?я ????а ???й, ??? ??о ????
?????к? ??, ???? ?н ?????? к ??????. Т?????я э?х э????, ?к ???о,
??о????я ???? ?????и э????.
– К ????и?ю??я э???? ?н?ят ?, ?????ия ???х ???а? ?з ???я ???? ??????
Т?????ю??я э???ы ????т
1. И?????е ????????? (IS – insertion sequences – ????????? ???и).
– Э? ????‚ ????е ?к ??е ?р????я ? ??? ???а ?????и в ??ой (?с 73 К???).
– Ип ???? ?? ? ??‚ ???е ????? ?я ? ?????? ?????я – ??????.
– ? ??х ??? ???о ? ?х ?е?а ?ею?я ???????е ???ы (?с. 10.4), ???е ??е
????? ?я ??????. Н?????е ????????? э?х ???? ???? ?я ??? IS и ???уют ?
??е ? 10 ? 40 п. н.
– Их ?????я в ??е ???? ??сы ??о ???ят к ???ям, ??ю?ю??я в ????? ??ля???
и ???ю?х ?а??? а ?????е э???ы ??? ?м? IS ??т ?ия? ? э????ю ?????х ??в.
– Ч??? ?????? у ?з?х э?м??в ?????а и ???ля? 10–5–10–7 ? ????е.
– П? ?????? IS в ??е ????е ???? IS – э??? о???я ? ???м ??е; т.е., ???ия
??????ся ??? ???? ??? ??и и ???т ? ??и???? ???й ?зя?а.
2. Т?????.
– Т. ??вают ???? ДНК‚ ???ю?е ?? ? ?????‚ ?о и ИП‚ ? ????? ??е ??‚ ?
?ею?е ????е ???????о к ??????
а) ???е ?????ы
– Ч?? IS – э???ы ?и?й ???? 1 т. п. н. ??ят в ??? ??е ???х ?????? (????а
1).
– Их ??а ??т ????ь ?сяч ?р ????й, а ?????е ???и ??т ????ь ?з???з?е ??,
(????‚ ?? ??й???и к ??????‚ ?? ???? ?и ?? ??????? ????в ????? ?????а)
– С ??х ??? э? ?????я ???ь ?????ся ?????и ISэ????и (? IS 1 ? IS 10) – IS
– L и IS – R с ??й и ??? ??? ??????но.
– Вся ?ф??ция, ?????я ?я ?????я ???? ?????а, ????ся в ?о IS – ч??; э? ?
??я ???а?я, ???я ?????ся ISэ???? ?к ???м.
б) П??? ?????ы.
– э? ?а???? ? ?я?? ? с ??? IS – э????и:
– ????ия, ?????я ?я ??????, ?????? в ?х ??х.
– В ???? ???а ??о ???? ????? ТnЗ (????а 3), ???й ???? ???????е ???? ???ы
??? 38 п. н., I R-L и I R-R, ?????е ?я ?????ии.
– М?? э?? ????и ??дя?я ?и ??: ?а ? ?х ???ют ????и???е ??и, ??? – ?-????у
(?н ?р, ? ?яз? с ??????й).
– К?? ??, ???я ????ю?й ???к ??? 170 п.н., ????? ????ы ??в ???????? ???,
а ??е ????е?? у???, ????? res и ?? ?????й ?я ??????.
С??-м??и
Э? ???? ДНК, в ???е ????ются Мэ
– О?и ?????ую??я э???ы ???? ????? и ???е ????ются в ???? ???? ????,
?????? ??? ?м?о ????? э???а.
– Д??е ??е ???? и ? ??ют я??о ?????ия ???м ???-??ням, ?тя ??ю??ся ????я
????я к ????нию ? в АТ-???е ???и.
– Н????о ? ?? Мэ. ?о ?????е в ??й ?????? ??с ??? ??ов???ся ?????й ????о
???а ДНК в в ??е-???.
– Э? ?п?????? ???? ??м ????ют ???в??я э???.
– П?? обя????я ?п???я ???-???й ?а??? ? ?, ?о ?и ???иях в ????е ?????о
???к ??с??т с ?????? в ??????х ???-??нях ????х ??????? ??ы?в, ?к э? ????
? ?с. 10.1. С?т-??? в ???, ????? ????й э???, ???о ???ют «??????, а ?
????? ???? – «?????.
Т?ы ??????:
Для ???? ????? т?????ю ? ??? ??? в ??? (????, ? ???? в ??????й ??м ?и
???о). В??????я?ая ?????ия ????т ??е ???.
1. К???????я ?и ??????я ?а????я (????у ????? ??ая ????ия э???а.)
– ???? ??м, ???й ??т ???о??ю??я э???, ????я с ?????? ????й ДНК (????а
4).
– О??????я ????? ???? ?ю ???ую и ?????ую ДНК, а ??е ? ??й ??и
?????ую??ся э???а в ??? ????ия э?х ДНК.
– К?????я ?лю?? ??? ???? ???иэ??? ?я?й и ?????е ??х, ????ию ??о э???а и
????ию ??а-???.
– К????т ??т ??м ?????я с ?????? ?? ???? ДНК, ??ая ? ???х н?? ??ю
??????а??о э???а. Для ????ия ????? ?о???????е ???? ?? tnp R‚ ??в???
????? (? ??. Resolution – р????е)‚ ???я ????т ????? ? ???? ?????ы.
– П? ?????? ??м ?????? ?????ся ? ??? ??р??я, ??????ая в ??м э???е, ? и
??????е ???и ???.
2. П??? ?????е ?и ????т??е (?и ???????м, ????у ???а?и ?к ???й ? ?????)
– Т?????ю??я э??? ?????ся в ??й ???? ??с, ?и э?м ???х ??? ?????, ??е
????? ??а-???, ? ????? (????а 4).
– Н???? ?????ую??я э???ы, ??и?р ДНК ?? Мu, ???уют ?к в ??????, ?к и в
???м ?????и.
3. С????т ?е ?? ??? ???????я ???, ???? ???й:
– ? ??? ??й ?о????? ????ы ????ля?ся ????и?я лю?? ? ?? ??в (?с. 10.9).
– В ?????? с э?й ???ю ??т ?????ь ?????ия ДНК ?? Мu и ??? ????х э????.
– На ???е, ?лю???ю?м ???, и ДНК ???, и ДНК ????? ???а??ы в ????й ??е,
????у в э?????? in vitro, ?????? ?я ???? ??????? ??? ???, ??????? ??ц?ая
????ая ДНК;
– ????м э???? в ???? ДНК ???а ???????ая ДНК ?? Мu. In vivo ??? и ?????,
???? ???? и ????ы Е.?li, ??е ??дя?я в ??ц?? ??е.
– С???о ??? ???, ?????ия ??н??я с ?????я ??????ых ???? в ????х ???? и
?????? ДНК.
– У ??? ??е ???ы ????ят с ??х З’-??? э???а, у ?ц???а – в ??е-??? с
??зо??? 5′-????.
– З?? ?????е ??ы ????? э??н? ???яю?я с ???и ?????, и ????ся ??й ??????й
???т.
– Е?и в ?пях, ???????х ? ??ше?ю к ?????м ??????о, ???х ??????? ???? ?
?????, ? ??т ??о ???е ?????е.
– В ????? ????ия ?е? и ?????я ????? ????ия ??а-???.
– Т?? ???? ??т ????ля?ся ???????я ?????ия Тn 10.
– Э? ???ю ?????? ?????а Тn 10, ???я ??ря? ????? ? ??? Тn 10, а ??е,
?-????, ??????е ?????х ???.
– П? ???а??? ?????? ??й ??????й ???т ?????? ??? ??? ?????я.
– ?????т ? ??й ????е ????? ????й ДНК с ??я ??????? ???и.
– Е?и ????ия ????ся в ??? ??е, ? ???? ???т ????ля? ??й ?????-???, ?????
и ???ую, и ?????ую ДНК, а ??е ??и ?????ую??ся э???а, ???????е ?????????
??м.
– П? ????? ?????а в ??е ??? ????ля?ся ?????ия ??й ??и ????? э???а в ???
??-???. Р????е ??т ?????ь ??м ?????? ?????? ??у ??я ??я? ????и?ю??ся
э???а с ???? ????????о ???? Е.?li. (????а 6).
Г?????я ?????? ???? ?и ??????
М???? э???ы ??вают ?????ую ??????? ????? ? ???а ??й ?????и, ???? в ??е?е
??????? ????а ??????.
В ?????и ? ??, ?к ???ы ???ы в ДНК-???, ????я ?? ???я (В???? ???а ????ы ?
? ????? ???и), ?? ???ия (??????) ??????о ????а ??у ??? ?????ия ?????а и
??нью ?о ?????я. В ?я? с э?м ?т??? ????, ?о ????ы ?????х ??в ???? ???ю?я
?? ? ??а ?о???е??и ??????и ??? э?? т?а. По-????, ????е э???ы ???и
?????ую ?? в ?????и и ???????и ????.
1) В????? ISэ???? ????? ? ????о ?? ??т ????ь к ?о ????и ? ?? ?????? с
????а IS – э???а, т.е. ??ня?ся ??ля?я ????л??о ?? (?. 8а).
2) О?? ??о, ?о ????е э???ы ??? ????? ????и ????и, ??????ая ?????ия ??у
???? ??т ????ь к ????иям ??в (?с. 76 К???). С???ся, ?о ????ия – ?? ?
???? ??й э?лю???? ??????я ??х ???й. Д??????, «??яя» к?ия ?? ???т ?-?д
???ия ???в??? ??? и ???? ?????ь ?????ь ????я. Ч?е ??о э? ???? к ??? ??й
? ? ? ?? ???и, ? ??? ??т ????ся ?н с ??? ???я?.
3) Н??я ???? и ?т ??, ?о ??? ??т ???? ?????е ?????? ? ?? ?????ия в ???е
?????а ??, ???й ?м ? ?? ???? ????я ???? ?я ???? в ?????? ???ях. Д??????,
? ?????? «?????ю? ?? ?????? к ????м ??????м я?м, в ?м ??е к тя??м ???? и
??????, ?? ??????? ??????? ??й, ???яю?е ?????? ????й и???к ????, ???ц,
?? ????х ????, ??ю?е ???? ?????и и ???яю?е ? ?м ??м ?????о ???? ??з ??и.
С???? в ??? ???и ????я и к ????м, ????у ?????о ???? ?я ???-?зя?а ?????
??в ????я в ???е ??????.
4) Е?и э??? ???н в ?? из ??в ???????о ???а, ? ????? э??? ??т ?ия? ?
э????ю ???ую?х ??в, ?? ???в??я ?????ию в ??е ?????, ??дя??я ??? ?о ???
(п???? 8,6), ?? ???яя ???я?ю ????? ???ю?х ?????????й ?НК (????а 8, в)
5) К?? ?? и ?т ? ??м ??? и ???м, и ?????м ?и о????? ?????а (??????ля?ая
?а????я), ?????ь ????х э??н?в ???? ?????и в ??д?х ???? ДНК ???? к ???
?????м э???м. В? ????я?е ?и э?м ???я, ?лю?я ???и и ????, ?я?? с ?????м
??? ?и ?? ??? ????? э???а с ??? ????????тя? ДНК (?с. 10.13); ??? э?х
????в ? ??а ? ?????ы.
П???? ?????? э???ы у э???т
С?????ю ??ь ??? э???т (10–30%) ???ляют ???яю??я ??е??????и, ?ею?е
?????ую ?????ю о????ию и ????е ?????ся в ??? ?к в ???? ??й ????ы, ?к и
??у ?????и. О? ???? ???? ????х ?????? э????. Э???ы ?лю?ют в ?? ??? ?
?ся? ? ?ся?? т?яч ?????? ?р.
Н????е ????? ????х э???? ????? в ??? ???? (? 50%). П???? ?????? э???ы
??? ??ея? ? ???, ? ??т ????????я в ????х ???? ????.
В?ы ????х э???? э???т
Р???ют ?а (? ???? ???ля?? ????? ?????я) ???? ??? ????х ?????? э????:
1. Т?????
Э? э???ы ????? и???????и ????и (????????тя?, ??а???? ????у ?? ??у), ?к и
????е ?????ы ????т. П???? ? ?г? ???ь Р-э??? ????ы и Ас-э??? ???? (?с. 92
К???). В ???х э?х ????? ??????я ? 30 – 50 ??й ??х э????. П??е ??и (??ь
??й д???а, ?к ?к ??т ????? ???и) ???? ???? ??и ????ия, ???ю?е ?????у
(?с. 92).
С?? ?р???ия ?????а ???? ? ?с. 93. И??????? ???ы ????? ?я ?????я э???а,
???е ????ля?ся ????я ? ????ю ?? с ??? и ????ю ??????и. И??????? ???ы
???ю?я и ??о ???ю?я ? ???? ???? ДНК ?зя?а. В????й ????он ??ря?ся в ??н
????о ?????? ???а в ????-??? и ????я с ДНК ?зя?а в ??м ??е. Р??в и ????е
????ляю?я ?????? и ???????? ???и. Т????? ??т ???в?ься ?к ??м ????м
э???ом, ???й ??т ?????ся, ?к и ??ой ??? э???а, ??????? в ?м ? ??? в
????и. Р-э???ы ??? ????ю?я в ?????? ??ы, ?????ая ????????? ???х ???:
5′-ГГЦЦАГАГ П? ?т???? Р-э???а э?т ??-??? ??????я.
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1fffffffff8067ffffffffffffffe000ffffffffffffffff3f9ffe7f867ffff3f9ffffff
f000fffff8fffffffffffffffffffe000fffffffff0067fffffffffffffff000ffffffff
ffffffff3f9ffcff013fffe7f9fffffff8007ffff8e7ffffffffffffffffff000fffffff
ff304ffffffffffffffff8007fffffffffffffff1f9ff1fe391fffc7f1fffffff8007fff
f007ffffffffffffffffff8007fffffffe721ffffffffffffffff8003fffffffffffffff
9f9ff9fe7c8fff8ff3fffffffc003ffff1cfffffffffffffffffffc003fffffffce71fff
fffffffffffffc003fffffffe1ffffff9f9fffff3c23fc1fe3fffffffe001fffffffffff
ffffffffffffffc003fffffff8c73ffffffffffffffffe001fffffff807fffff9f9fffff
3e10007fe7fffffffe000fffffff3fffffffffffffffffe001fffffff9ce7fffffffffff
fffffe000fffffff1c3fffff9f9fffff9e0403ffcfffffffff000fffffff1fffffffffff
fffffff000fffffff39c7fffffffffffffffff0007ffffff3f3fffff1f9fffffcf01ffff
9fffffffff8007fffffe0ffffffffffffffffff000ffffffe13cffffffffffffffffff80
07fffffe7f9fffff3f9fffffe7007fff1fffffffff8003fffffc47fffffffffffffffff8
007fffffe039ffffffffffffffffffc003fffffc7f9fffff3f9ffffff00ffffe3fffffff
ffc003fffffcf17ffffffffffffffffc003fffffce33ffffffffffffffffffc001fffffc
f79ffffe7f3ffffff8fffff87fffffffffe001fffff8787ffffffffffffffffc003fffff
8f03ffffffffffffffffffe001fffff9c3cffffe7f3ffffffc3ffff1ffffffffffe000ff
fff03c7ffffffffffffffffe001fffff87c7fffffffffffffffffff000fffff0c1c7fffc
ff3fffffff07ff83fffffffffff0007ffff38cffffffffffffffffff000fffff31cfffff
fffffffffffffff0007ffff03ce3fff8fe7fffffff80000ffffffffffff8007fffe3c1ff
ffffffffffffffff0007fffc388ffffffffffffffffffff8003fffe71c71fff1fe7fffff
fff0007ffffffffffffc003fffc1e1ffffffffffffffffff8007fffe3e1fffffffffffff
fffffffc003fffc3c6187fc3fcffffffffff87fffffffffffffc001fff88f3ffffffffff
ffffffffc003fffc0f3ffffffffffffffffffffe001fffc1e60e000ffcfffffffffffe7f
fffffffffffe001fff9e27ffffffffffffffffffc001fff9c63ffffffffffffffffffffe
000fff98e303801ff9fffffffffffe7fffffffffffff000fff0f0fffffffffffffffffff
e001fff1e07fffffffffffffffffffff000fff1e0100fbfff3fdfffffffffc7fffffffff
ffff0007fe078ffffffffffffffffffff000fff0f8ffffffffffffffffffffff8007fe0f
08003fffe7fcfffffffffcffffffffffffff8003fe739ffffffffffffffffffff8007fe6
39ffffffffffffffffffffff8003fe639e3fffffcffcfffffffff8ffffffffffffffc003
fc783ffffffffffffffffffff8003fc719ffffffffffffffffffffffc003fc713f3fffff
0ffcfffffffff0ffffffffffffffc001f83e3ffffffffffffffffffffc003f87c3ffffff
ffffffffffffffffe001f8783f8ffffe3ffc7fffffffe0ffffffffffffffe000f91e7fff
fffffffffffffffffe001f83e7fffffffffffffffffffffff000f81c7fc3fff87ffe7fff
ffffc07ffffffffffffff000f3c4fffffffffffffffffffffe000f38c7ffffffffffffff
fffffffff000738cfff05f81fffe7fffffffe07ffffffffffffff80061e0ffffffffffff
ffffffffff000e3c0ffffffffffffffffffffffff80061e0fffc0007fffe3ffffffff27f
fffffffffffff80020f1ffffffffffffffffffffff80061f1fffffffffffffffffffffff
fc0020f1ffffc03fffff3fffffffe3fffffffffffffffc000e33ffffffffffffffffffff
ff8000873ffffffffffffffffffffffffc000c73ffffffffffff9fffffffe7ffffffffff
fffffe000f03ffffffffffffffffffffffc001e33ffffffffffffffffffffffffe000f07
ffffffffffff9fffffffcfffffffffffffffff000787ffffffffffffffffffffffe000f8
7fffffffffffffffffffffffff000787ffffffffffffcfffffff8fffffffffffffffff00
63cfffffffffffffffffffffffe0007cffffffffffffffffffffffffff8031cfffffffff
ffffe7ffffff1fffffffffffffffff80f89ffffffffffffffffffffffff00f1cffffffff
ffffffffffffffffff80789ffffffffffffff3fffffe3fffffffffffffffffc3fc1fffff
fffffffffffffffffff81f81ffffffffffffffffffffffffffc1fe1ffffffffffffff9ff
fffc7fffffffffffffffffc7fe3ffffffffffffffffffffffffc7fe3ffffffffffffffff
ffffffffffe3fe3ffffffffffffffc7ffff8ffffffffffffffffffe7fe7fffffffffffff
fffffffffffc7fe7ffffffffffffffffffffffffffe3fe7ffffffffffffffe1fffe1ffff
fffffffffffffff3fe7ffffffffffffffffffffffffe7fe7ffffffffffffffffffffffff
fff3fc7fffffffffffffff03ff07fffffffffffffffffff1fcffffffffffffffffffffff
ffff3fcffffffffffffffffffffffffffff9fcffffffffffffffffe0001fffffffffffff
fffffff9f8ffffffffffffffffffffffffff1f9ffffffffffffffffffffffffffff9f8ff
fffffffffffffff800fffffffffffffffffffff9f8ffffffffffffffffffffffffff9f9f
fffffffffffffffffffffffffff9f8fffffffffffffffffffffffffffffffffffffffff9
f9ffffffffffffffffffffffffff9f9ffffffffffffffffffffffffffff9f8ffffffffff
fffffffffffffffffffffffffffffff9f8ffffffffffffffffffffffffff9f9fffffffff
fffffffffffffffffff9f8fffffffffffffffffffffffffffffffffffffffff9f9ffffff
ffffffffffffffffffff9f9ffffffffffffffffffffffffffff9f8ffffffffffffffffff
fffffffffffffffffffffff801ffffffffffffffffffffffffff9f9fffffffffffffffff
fffffffffff800fffffffffffffffffffffffffffffffffffffffff800ffffffffffffff
ffffffffffff801ffffffffffffffffffffffffffff800ffffffffffffffffffffffffff
fffffffffffffff9f9ffffffffffffffffffffffffef801fffffffffffffffffffffffff
fff9f8fffffffffffffffffffffffffffffffffffffffff9f9ffffffffffffffffffffff
ffcf9f9ffffffffffffffffffffffffffff9fcffffffffffffffffffffffffffffffffff
fffffff9f9ffffffffffffffffffffffffcf9f9ffffffffffffffffffffffffffff9fcff
fffffffffffffffffffffffffffffffffffffff801ffffffffffffffffffffffffcf9e1f
fffffffffffffffffffffffffff800fffffffffffffffffffffffffffffffffffffffff8
01ffffffffffffffffffffffffcf801ffffffffffffffffffffffffffff800ffffffffff
fffffffffffffffffffffffffffffff9fdffffffffffffffffffffffffcf859fffffffff
fffffffffffffffffff9f8fffffffffffffffffffffffffffffffffffffffff9fdffffff
ffffffffffffffffffcf9f9ffffffffffffffffffffffffffff9fcffffffffffffffffff
fffffffffffffffffffffff9fdffffffffffffffffffffffffcf9f9fffffffffffffffff
fffffffffff9f0fffffffffffffffffffffffffffffffffffffffff801ffffffffffffff
ffffffffffcf801ffffffffffffffffffffffffffff800ffffffffffffffffffffffffff
fffffffffffffff881ffffffffffffffffffffffffcf801fffffffffffffffffffffffff
fff8e8fffffffffffffffffffffffffffffffffffffffff9fdffffffffffffffffffffff
ffcf9f9ffffffffffffffffffffffffffff9fcffffffffffffffffffffffffffffffffff
fffffff9fdffffffffffffffffffffffffcf9f9ffffffffffffffffffffffffffff9fcff
fffffffffffffffffffffffffffffffffffffff801ffffffffffffffffffffffffcf9f9f
fffffffffffffffffffffffffff800fffffffffffffffffffffffffffffffffffffffff8
01ffffffffffffffffffffffffcf801ffffffffffffffffffffffffffff800ffffffffff
fffffffffffffffffffffffffffffff9fcffffffffffffffffffffffffcf801fffffffff
fffffffffffffffffff9fcfffffffffffffffffffffffffffffffffffffffff9fcffffff
ffffffffffffffffffcf9f9ffffffffffffffffffffffffffff9fcffffffffffffffffff
fffffffffffefffffffffff9fcffffffffffffffffffffffffcf9f9fffffffffffffffff
ffff9ffffff9fcfffffffffffffffffffffffffffffe1ffffffffff800ffffffffffffff
ffffffffffcf801fffffffffffffffffffff83fffff800fffffffffffffffffffffff371
1a0001fffffffff800ffffffffffffffffffffffffcf801ffff98180c00000000000003f
fff800fffffffffffe000000000000000000007ffffffff9fcffffffffffffffffffffff
ffcf9f9ffff00000000000000000001ffff9fcfffffffffffe00000000000000000403ff
fffffff9fcffffffffffffffffffffffffcf9f9ffff800eaf977ffffffff81fffff9fcff
fffffffffffffffffffffffffffc3ffffffffff800ffffffffffffffffffffffffcf9f9f
ffffffffffffffffffff8ffffff800fffffffffffffffffffffffffffffefffffffffff8
00ffffffffffffffffffffffffcf801ffffffffffffffffffffffffffff800ffffffffff
fffffffffffffffffffffffffffffff9fcffffffffffffffffffffffffcf801fffffffff
fffffffffffffffffff9fcfffffffffffffffffffffffffffffffffffffffff9fcffffff
ffffffffffffffffffcf9f9ffffffffffffffffffffffffffff9fcffffffffffffffffff
fffffffffffffffffffffff9fcffffffffffffffffffffffffcf9f9fffffffffffffffff
fffffffffff9fcfffffffffffffffffffffffffffffffffffffffff800ffffffffffffff
ffffffffffcf801ffffffffffffffffffffffffffff800ffffffffffffffffffffffffff
fffffffffffffff800ffffffffffffffffffffffffcf801fffffffffffffffffffffffff
fff800fffffffffffffffffffffffffffffffffffffffff8fcffffffffffffffffffffff
ffcf9f9ffffffffffffffffffffffffffff9fcffffffffffffffffffffffffffffffffff
fffffff9fcffffffffffffffffffffffffcf9f9ffffffffffffffffffffffffffff9fcff
fffffffffffffffffffffffffffffffffffffff800ffffffffffffffffffffffffcf9f9f
fffffffffffffffffffffffffff800fffffffffffffffffffffffffffffffffffffffff8
00ffffffffffffffffffffffffcf801ffffffffffffffffffffffffffff800ffffffffff
fffffffffffffffffffffffffffffff9fcffffffffffffffffffffffffcf800fffffffff
fffffffffffffffffff9fcfffffffffffffffffffffffffffffffffffffffff9fcffffff
ffffffffffffffffffcf9f9ffffffffffffffffffffffffffff9fcffffffffffffffffff
fffffffffffffffffffffff9fcffffffffffffffffffffffffcf9f9fffffffffffffffff
fffffffffff8fcfffffffffffffffffffffffffffffffffffffffff800ffffffffffffff
ffffffffffcf8f1ffffffffffffffffffffffffffff800ffffffffffffffffffffffffff
fffffffffffffff800ffffffffffffffffffffffffcf801fffffffffffffffffffffffff
fff800fffffffffffffffffffffffffffffffffffffffff9fcffffffffffffffffffffff
ffcf8f9ffffffffffffffffffffffffffff9fcffffffffffffffffffffffffffffffffff
fffffff9fcffffffffffffffffffffffffcf9f9ffffffffffffffffffffffffffff9fcff
fffffffffffffffffffffffffffffffffffffff800ffffffffffffffffffffffffcf9fdf
fffffffffffffffffffffffffff800fffffffffffffffffffffffffffffffffffffffff8
00ffffffffffffffffffffffffc7801ffffffffffffffffffffffffffff800ffffffffff
fffffffffffffffffffffffffffffff8f4ffffffffffffffffffffffffcf800fffffffff
fffffffffffffffffff8fcfffffffffffffffffffffffffffffffffffffffff8fcffffff
ffffffffffffffffffcf9fcffffffffffffffffffffffffffff9fcffffffffffffffffff
fffffffffffffffffffffff8fcffffffffffffffffffffffffc79fcfffffffffffffffff
fffffffffff8fcfffffffffffffffffffffffffffffffffffffffff800ffffffffffffff
ffffffffffc79f8ffffffffffffffffffffffffffff800ffffffffffffffffffffffffff
fffffffffffffff800ffffffffffffffffffffffffcf800fffffffffffffffffffffffff
fffc00fffffffffffffffffffffffffffffffffffffffff8fcffffffffffffffffffffff
ffcf800ffffffffffffffffffffffffffff8fcffffffffffffffffffffffffffffffffff
fffffff8fcffffffffffffffffffffffffc79fcffffffffffffffffffffffffffff8fcff
fffffffffffffffffffffffffffffffffffffff800ffffffffffffffffffffffffcf9fcf
fffffffffffffffffffffffffff800fffffffffffffffffffffffffffffffffffffffff8
00ffffffffffffffffffffffffc7800ffffffffffffffffffffffffffffc00ffffffffff
fffffffffffffffffffffffffffffff8fcffffffffffffffffffffffffc7800fffffffff
fffffffffffffffffffcfcfffffffffffffffffffffffffffffffffffffffff9fcffffff
ffffffffffffffffffcf9fcffffffffffffffffffffffffffffcfcffffffffffffffffff
fffffffffffffffffffffffdfcffffffffffffffffffffffffc79fcfffffffffffffffff
fffffffffffcfcfffffffffffffffffffffffffffffffffffffffff800ffffffffffffff
ffffffffffc79fcffffffffffffffffffffffffffffc00ffffffffffffffffffffffffff
fffffffffffffff800ffffffffffffffffffffffffc7800fffffffffffffffffffffffff
fffc00fffffffffffffffffffffffffffffffffffffffffcfcffffffffffffffffffffff
ffc7800ffffffffffffffffffffffffffffcfcffffffffffffffffffffffffffffffffff
fffffffcfcffffffffffffffffffffffffc79fcffffffffffffffffffffffffffffcfcff
fffffffffffffffffffffffffffffffffffffffcfcffffffffffffffffffffffffe79fcf
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00ffffffffffffffffffffffffe7800ffffffffffffffffffffffffffffc00ffffffffff
fffffffffffffffffffffffffffffffc0cffffffffffffffffffffffffc7800fffffffff
fffffffffffffffffffce4fffffffffffffffffffffffffffffffffffffffffcfcffffff
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ffffffffffc79fcffffffffffffffffffffffffffffc00ffffffffffffffffffffffffff
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fffc00fffffffffffffffffffffffffffffffffffffffffcfcffffffffffffffffffffff
ffc7800ffffffffffffffffffffffffffffcfcffffffffffffffffffffffffffffffffff
fffffffcfcffffffffffffffffffffffffe79fcffffffffffffffffffffffffffffcfcff
fffffffffffffffffffffffffffffffffffffffcfcffffffffffffffffffffffffe79fcf
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00ffffffffffffffffffffffffe7800ffffffffffffffffffffffffffffc00ffffffffff
fffffffffffffffffffffffffffffffc00ffffffffffffffffffffffff01800fffffffff
fffffffffffffffffffc00fffffffffffffffffffffffffffffffffffffffffcfcffffff
ffffffffffffffffff819fcffffffffffffffffffffffffffffcfcffffffffffffffffff
fffffffffffffffffffffffcfcffffffffffffffffffffffff839fcfffffffffffffffff
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ffc7800ffffffffffffffffffffffffffffcfcffffffffffffffffffffffffffffffffff
fffffffcfcffffffffffffffffffffffffc79fcffffffffffffffffffffffffffffcfcff
fffffffffffffffffffffffffffffffffffffffcfcffffffffffffffffffffffffe79fcf
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fffffffffffffffffffffffffffffffc00ffffffffffffffffffffffffff800fffffffff
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fffffffffffffffc00ffffffffffffffffffffffffff800fffffffffffffffffffffffff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В?????
Р?. 93 П????? ?????а
Б?? в ДНК ??е ?р??ия ?????а ??т ?????ся ?бо с ???ю ??????о ???а ДНК, ?бо
? ?? ???ия ?????х ??? (э? ???? ????? ?я ?????? ???ий).
Э??т ? ?????я ?????а э???т
Р-э??? ????ся ??? в ???р? ??ях D. melanogaster. С????? ?м? ?з Р-э???а с
???и, ???? Р-э???ы, ???? у ???? к ?????иям Р-э???а, ???е ??ю?ю?я ??? в
???х ?????? ??. С?о???? ??ю??? э???в ???ют ????м ?????. Д???? ?? ??о
???аю?я ?????и, а ?? ?и ????ы, то у ? ???? ??????ся ????о ???й и э? ????
???ю?я ??ую?м ????ям. Поэ?? ??и с Р-э???? и ?з ?? ??ядят ?к ??????о
??????е, ? ???й ?? ????. Б?????ая ??яция ??? ???ую ?? в ???? э?лю?и.
И?ля?я ??й ???а ????? ?????? Р-э???а, ?????ю?? в ??й ? ?х. П???, ???
?????? Р-э???а ????? ?????? ????, ?ъя?я?ся ?м, ?о ??? в ???х-???ст????
??т ????ля?ся ??й ?п ?????, ???й ???? к ?????ю ?????й ???? ??и ???я?и,
???ю?й ?????у (см. ?с. 92). О????? ?????? ???ш??и ???? ??т ?????? ??л,
????у ?????? ????е ???, ???х ???, в ???х ????и ???? ?????и ????? ??????
Р-э???а. П???й «???? ??, ?????ю??я ???й ???? ?????, ??т ????? ???ю ???ь
???? ??????, ???я ???? м????м ?я ??? в ?о??е э?лю?и.
П????ия Ас-э???а ????ят и в ?????? ???х ????. За ?? м?? ???ь ? ??????ю
????х ??т?в ??и, ???? ???? ???? ???т?е ?????и ??, ???ляю?? ????цию.
П???? ???, ????? ??? ???????? ??, ??е ??т ??? п???а. В???? ????? э???а
???? к ?????и ??. В ?я? с э?м ?л ??? ??д о ??ля??? ???и п???аю??я э????
и ?и ?? ???ы ?????ю?? (Б. Мак-К???).
В ??м ????е э???ы э???т п????яют ??й ?????о ?????ю ??ля?ю. С????т ??? о
?м, ?о ?и не ?ияют ? ???п ????а и ????ю?я в ??? ?? ????я ?????ям ??й
????ы, в ????? ?? ?с???о ??ляют ??м. П??????я, ?о ?и ???ляют ??ь ?к
????? э????? (?. с. 156) ДНК, ?????е ???й ??????ся ?????? ???м. О??
???яет ? ???, ??а ????е э????? ДНК ??????ся ???? ?????я?. На ??????
??нях э?лю?и э? э???ы ??т ??????ся ?я ??й, ???е в ????? ????т ?????е
?????? ????у.
2. Р???????ы.
А) с ???? ????и ????и
– С??ы ? ??й ????е с п???????и, ???е ??ряю?я в ??м, ????я ????ы ????
?????ии.
– Э? э???ы ???? «??» ???? 5–8 ?с. н.п., ?????? ?я?? ???? ????и ????и
(ДКП, ?и LTR – ? ??. 1?g terminal repeats), ??? ?????и ? 300–400 н. п.
(?с. 94).
– Ч?? к?? э?х э????, ??????х к ??? ????у, ????? ??оя?о ?я ??, ? ????т ?
????? ??й ? ??н ?сяч ??й в ?????и ? ?? ????????.
– В ??а? «??» э???а ?н???аю?я ???ые ??и ????ия ?я ???? ??????ы и ????
(????ы).
– Р??е ??и ??? ????а ??т ??ь ???и и ???и ?к в ДКП, ?к и в ?м? э???е.
– По ???м ДКП ??дя?я ???? ?я?е ???ы.
С??б ?????я ????????в с ???? ????и ????и
– ?????ет ?о ?????ию с ???ю РНК-????? П.
– В ??а? ДКП ?ею?я ??ы ????и ?????? и ???ы ????????ия. ДКП ??т с???
????и ?????и.
– Т?????я ????ся в ??м ДКП (???о ??м, 5′-ДКП), а ??????я в ??? (???,
3′-ДКП).
– С?????ая ???? РНК ??????я с ?????? ????в, ?????х ?я ???а ДНК и ????я ?
в ??м.
Э? ??а ????ю ???я? ??у о???а?я и ????? ????а (?с. 95). О???я ???? ???:
????и ? ?????ы ? ????????в ?и, ????, ???????? ???? ? ???в в ????? ???
?????и к ????ю? П??ляю?я ??ь ????????в (?и ????и ? с ??????и) ?? ??ря?
?н ???? ???, ?? ?е ? ????а ?о. Т?? ???м, ???????? ??с??яют ??й ???????,
??????? э?м??, ????е ?? к ?????????ю и «?????ию» ?? ? ???.
В???? ???????? ??? ?? ?и ??о ?? ???? ??? э???:
– ????? ?????я ?? (?с. 96 а).
– ?и ?т???и в ??н ????а ? 5′-??? ?? ???????н ??т ??о ?????? э????ю ??,
?????я ?????ию с ?????? ????а (?с. 96 б1). Е?и в э?м ??? ????й э???
???ся ??о ??????а, ? ?????м ??т б?ь ??????ия ??а и ???????е ?????? ???.
– э? ? ???? ?и ?????и э????в, ???с?? ????м э???? (?с. 96 б2).
– О??е ?????и ?я ?????я и ?????ия ??ля??? ???? ???ают ??а, ??а эл??т
(???????н) ??я?ся ? ?? ?????? ????? ??у ДКП с ?????и ????и (?. (?с. 96
в). В ?з???е ???я?ся ?? ?? ДКП ? ??е ????я ????????.
П??????е ??? ????я э???а ?????ю?я ???я? с ??? ???и ?н?????? ?оя??ия?
??????? ?????м ?????я ?и, ????, ???????й ??а.
П?е???е э???? ? ??? ??????
– ?????н?ию ??ля??? ???? (??? ????и ??????, ???? ????????ия, ?и э????в),
?о ??? ?????? ?? ???н? э???? в э?лю?и ??? ??ля?и.
Б) Р???????ы, ???е ? ?с? ???х ???? ????. М???м ????я ?
– ????о ??,
– ? ??е ????ля?ся с ???ю ???? ????п?и.
– К ?м ??ся?я ??????и ????а L1 ??ляющ? ??м ????.
– Р????я э???а ?з ДКП ??я?? ????с??? с ???м ???? ????я ????н???а
(?с. 97).
– РНК, ?р????яся ?и ?????? э???а, ?????ся к ????? ????? ??у ???а
ДНК-???,
– ??о с???ся с ??й ? ??й ДНК.
– Сю? ???ляю?я и ?????е ?я ????? ??и – ???ая ??????а и ????а.
– Д??я, ???????я ?? ДНК ??? ????й ?я ?????я РНК-??и э???а с ???ем ????
??????ы.
– Ф??? ???? ???а ????й ???к ДНК-?ш?и, а з?? ?ня? ???у и ???? РНК,
– ??е ч?о РНК ??я?ся и ????ся ??ая ?м?????ая ?? ДНК.
Р???????ы ?з ДКП ???уют в ????? ??? ???? в ря? ????й.
П???ю ???ю у ??? ????? ???я? ?????. У ря? ????х, в ?м ??е и у ????ы,
?????т ?????ая ?????. К?? ДНК у ?х ??няю?я ? ?? ?????й ??????з?? и в э?м
??? ???ают ?к ????? ???, ??аю?е ????у ? ??а???я. Р???????ы ????ю?я, ???я
???яю?юся ????у, в ???й эл??? ????ы ?? с ??? ? ?? «??? к ???». С??? ?
РНК-??с??? ?к ? ???е с ???ю ???? ?????? ???ся ???????я ?? ДНК, а ??м ??е
???ия РНК-???ы ?ст????я ??ая. Т?? ???м, ??м ?зя?а ?????л э? э???ы для
????ия ???? ???? ????, ?я ???ия ????ы ? ??? ??в.
3. Р?????
В ??? ????аю?х, ??, ???й и ????х ?????ю?я р????? – ?????ся в ??м
ДНК-??и, ???????е ? ??? ??х ????х РНК, ?к ? ????. С???? ?????? ???я? с
?????ью ???? ? ???и ???? ?????? в ???? ? ?????я. Т?? ???м, ?ря? с ????м
????? ? ДНК к РНК ????ля?ся и ???? ???с – ?????е ? в г?? в ?? ??????. У
????аю?х ?????ы ???ляют ??е 10% ДНК, ??????о, ???? ????? ??? ????? ? РНК
к ДНК ??т ?? ??????, ? ???й ?? ?и о??е ?о ? ????м э?лю???м ????.
Р???ют ??? ?? ??????, ??и ?х:
А) П????ы. Э? ??и ??в, ???? ??о?в и ???ю?е ?????????ю ??-?-??-? ?
3′-???. М???й ?и ? ?????и ????а «???????ая» п????????ая ?НК. Р?? ???я?и
??х «??? ??о «????? ??-???? и ???я?. О?, ?к и ???????, ????? ????и ?я??
????и, ????ляю?? ??й ????ию ???? ????????? в ???и ??а вн???я ДНК-??и,
?????? ? ?НК.
Б) Р????. Е?и ????? ????о ??????н, ДНК-??я ???? ???ую ??у ????ия и ????
?о э????я, ? ??е ?????ы ???ют ?????и.
Б??ую ??у ???яю??я э???? ?з?н??х ????ляют ДНК-??и ????х РНК, ???ю?х ??и
?????й ???ы, а ??е ????о ???????? ????х ?????? ?НК, 7S РНК и UPНК. О?
????ляют ??й ???е и ???? ???ы. С???? ???яю??я ?????????й ??о ???ют ??
???ия ? ??е?яю?м ? ?????? (????, ???? Aluп??? ?и ???е Крnпо??ы в ???
????).
К??й ?д ????аю?х ???? ???е?? ???м ????? ???х ????, ?л?аю?х ?о ? ??? ??в.
П?д??гают, ?о ??? ????а ???х ???? ???? ?????? ???о, ?е ??е эв?ю???й
?????и ??в ????аю?х.
К???е ???ы, ???? ???ы Alu – с???? у ????, ???ляют 3 • 105, ?о ??????т 5
– 6% ??ы ДНК ???. Их ?к????я ????????? ?????а ?????е???и ????х ???? 7S
РНК. Т?? ???ы ??ея? ? ???, ? ?????ют в ????, ? 5′-???х ??в, в ???е
З’-???????х ???? ??. М?? ??я? ??? ?? ??? ??ю?ю?я ???ия (?????м) в ????е
??в и ? б????о ????я, ??????е ????? ?р??х ?и ???х ????. В ??? ??с??т
????о ????? ????х ??й эт? э????, ?и ? ???? ????? т????? ?и ?р??ю
??????у. С ??? ???ы, ?и ???яют ?????ь к ?????ям, ?? в ??? ?????? ?ею?я
???????е ???ы, ????? ??????, а в ??? ????????в ????ы ???? и ?????ь
?????? э???а. М?же?? ??х ????х ??й ??? ?????ся, ?? ????, ??????е ?
?????е, ??т ?????ся ???и ?????? э????и. В ??? ???? ????? ???? ????
??????ы я?я?ся ???????н L1, ??о ??й ???? ????т 100 ?с. Из ?х ???о
????ю?я 30 – 60 ?с., ????е, ??????, ?????ся ? ??т.
М???? ?????? э???ы ??т ????ь ???е в ?????? ????. Н??? в ?о???х ?????
????? ? ?????? ???о?????и ??й ????? э??н? ???я? в ??? ??аях ?????ь
?????ию ? ???м ? ?????и. В ????? ????о ??????а ???ают д??? ????х ????
?и, ????, ??и??и (?с. 98).
Н???й ?????? ? ???м л????? ?????а mariner (?ряк) у ???? ?и??т к ???и
???а в ???? ??е 17й ????ы. Е?и э? ???е п????т в ?????й ??? ?и ????? ??т,
? ????а с ???? ??т ???? ????. Э? ???? к ??????? ?????ям ???й ???ы –
????иям и ????м. В???????ая ?????ия ??у ??я э????и ???? к ???? – ????
???а ????ы ? 180°. И???я ??т ?? ??? ?я ????а, а ??т ??????? э?лю?и ?н?а,
????у ???? ???? ????у ???? ?????я ????я?? с???? ??в, ??я??уя ??????у.
Р???????ы с ДКП ???уют в ????? ????? ???? ДНК. О??о ??й ??? ?????ся с
???ю ?????? ???лы ДНК, ???? ?????й, ??? ?о ???????й ??. У??? ????ую??я
ДНК ??о-?????? с ДКП в ???? ????ия ????? ???? ????? у ???й (?с. 99). Б??
в ????? ???и, ????ая ????????м, ???я? ?????ь ????ы, ? ??ня? ? ?????ую
?????????. Е?и ?й? ????ы, ?е ?? ??ь, ? ???? ??????о ??, ? ??? ???ся
???????.
П???? э???ы ? ??т ??????ь ??? ?к «э?????ую» ДНК, ??з??ую?ю ? ДНК
???-?зя?а. О? ??????ы и ?я ?? ??? ???: ?????ют ????ию ?????й ?????и
?????й ДНК; ?е?яя? в ??, ? ??? ?????ют, ?з?ая ???и, ? и ?няют ???? ?
э????и. В ?я? с э?м ?дв??е ?????? э???ы, ??? ???и ф???? ?????? ??в и
???уя в ?????? ????ы ????, ?еют ???? ???? в ????х э?лю?и ???в.
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