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.
Koterov A.N. Low dose of radiation: the facts and myths. First book. The
basic concepts and genomic instability. oscow, Publ. by FMBC by A.I. Burnazjan FMBA of Russia, 2010 283 p.
The monograph; 283 pages.; 46 illustrations; 14 tables; 730 sources.
The first book of monograph of Full Doctor of Sciences, Chief of Scientific
Analytical Laboratory of Radiobiology of Technogenic-caused Diseases of FSE Federal Medical and Biophysical Center by A.I. Burnazjan FMBA of Russia, Moscow
(e-mail: govorilga@inbox.ru) Alexey N. Koterov is devoted to various aspects of low
dose radiation concept and the radiation-induced genomic instability. On the base of
the experimental sources analysis the conclusion is made that with reference to low
LET radiation (x-ray, -radiation, etc.) there are no facts of an induction of genomic
instability after exposure at low doses radiation for cells without obvious defects and
an organism non in utero. The critical consideration is present for works and corresponding ideas about low doses radiation and radiation-induced genomic instability in
the scientific world of Russia, Ukraine, Belarus and in the West.
The hypothetical molecular mechanism of radiogenic carcinogenesis and
hereditary genetic anomalies induction which connected with genomic instability in a
range of low doses radiation with low LET is absent.

.
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International Commission on Radiological Protection).

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AA1. Abramsson-Zetterberg L., Zetterberg G., Sundell-Bergman S. et al. Absence of genomic instability in mice following prenatal low dose-rate gamma-irradiation // Int. J. Radiat. Biol.
2000. V. 76. 7. P. 971977.
AB6. Baulch J.E., Raabe O.G. Gamma irradiation of Type B spermatogonia leads to heritable genomic instability in four generations of mice // Mutagenesis. 2005. V. 20. 5. P. 337343.

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Research Council. (http://www.nap.edu/catalog/11340.html, .
http://books.nap.edu/openbook.php?record_id=11340&page=11.)
2
ICRP Publication 99. Low-dose Extrapolation of Radiation-related Cancer Risk. Annals of the
ICRP. Ed. by J. Valentin. Elsevier 2006. 147 p.; ICRP Publication-99. Draft of December 10,
2004. 212 p. http://www.icrp.org/docs/Low-dose_TG_rept_for_web.pdf; Draft of May 10, 2005.
3
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Fifty-fourth session Vienna, 29 May to 2 June 2006. 101 p.(draft).
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United Nations. UNSCEAR 2006. Report to the General Assembly, with Scientific Annexes. Annex C. Non-targeted and delayed effects of exposure to ionizing radiation. United Nations. New
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,
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[AR13].

43

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39, AL34], , , ,
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.
[AU8,
AU13] (, , ),

,

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-1993 -2000 2040 [AU8, AU13],
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- : . 1.2.1 1.2.2,
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, .

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..
1988 1992 . [AL28, AL29].
.
28

44

, 2040 , , ,
.

45

1.3.1. [AL28].
() in vitro. :
Pohl-Ruling J. et al., 1983 [AP13], : Lloyd D.C. et al., 1992
[AL28]: 6- .
0 .

1.4. ()


200 (0,2 )
.
-, [AF24]:
64
(NCRP-1980) [AN9].
:
1986 . [AU5], 1993 . [AU8], 1994 . [AU10] ,
, 2000 . [AU13]. -
.
, .
, ,
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, ,
. .
in vivo ().
,
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19862000 [AU5, AU8, AU10, AU13]. , ,
. - ,
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46

47

48

49

1.4.1. .

() ( 19862000) [AU5, AU8, AU10, AU13].

50

( , ,
). ,
/? .
1) ,
, ,

, , ,
()
, in vivo.
2)
:
IBiol = D + D2 ( -1993 IBiol = 1 D + 2 D2,
, , ),

,
.
3) , (D)
.
:
D = D2, , = D, D = /.
, ,
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4) 1993 2000 . :
,

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,
. :
/ = 200 , , = /200 .
-1993 2000
20 . .
, , IBiol = D + D2, = /200 , :
IBiol = (20 ) + (/200 ) (20 )2,
IBiol = (20 ) + (2 ),
IBiol = (22 ).
, , 20
9% (2 22 ).
40 :
IBiol = (40 ) + (/200 ) (40 )2,
IBiol = (40 ) + (8 ),
IBiol = (48 ).

51

, 40
(8/48)100% = 16,7%.
, , , ,
, , , , -

917%.

2%, , ,
(.
. 1.2.1). , / = 200 ,
,
2% ( . ), 4,2 .
510
, .
5)
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.
, (..,
)
.

.. (D.A. Pierce) [AP9, AP10].
/ 1 .
, 100 (0,1 ) :
IBiol = (0,1 ) + (/1 ) (0,1 )2,
IBiol = (0,1 ) + (0,01 ),
, 100

10% (0,01/0,11).
200 (0,2 ), , 20%.
:
IBiol = (0,2 ) + (/1 ) (0,2 )2,
IBiol = (0,2 ) + (0,04 ),
17% (0,04/0,24 = 0,167).
6) ,
200 .
, , (
) , , 2040 .
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2040
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,

52

, , 2%
( ).

. 1017%, 20
40 100200 ,
2,53,5 (. . 1.2.1).
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3 .
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7) -1993 -2000
. (. . 1.4.1 , ),
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, 200 10%,
500 23%.
IBiol = (0,2 ) + (/1,7 ) (0,2 )2,
IBiol = (0,2 ) + (0,024 ),
0,0235/0,224 = 10,5%.
IBiol = (0,5 ) + (/1,7 ) (0,5 )2,
IBiol = (0,5 ) + (0,147 ),
0,147/0,647 = 22,7%.
-1993 -2000:
200 .
, , .
20%
200 (. ). , .
0,42 :
IBiol = (0,42 ) + (/1,7 ) (0,42 )2,
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( / = 1 . ),
20 (0,02 ):
IBiol = (0,02 ) + (/1 ) (0,02 )2,
IBiol = (0,02 ) + (0,0004 ),
35 .
. 1.4.1
().

53
1.4.1.
NCRP-1980 [AN9]
1986, 1993, 1994, 2000 ( -1993 [AU8] -2000 [AU13])

, %

2040

200

(
1991 .) [AP9,
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35

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: 0,22 ,
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29

54

, ; ,
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[7].
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AK26] . ,
, .

[7], .
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(BEIR-VII) [AB10],
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). ,
0 100 , ;
100 1 ( ..):
In this report the committee has defined low doses as those in the range of
near 0 up to about 100 milliGray (mGy) of low-LET radiation, with emphasis on the
lowest doses for which meaningful effects have been found. Additionally, effects that
may occur as a result of chronic exposures over months to a lifetime at dose-rates below 0.1 mGy per minute, irrespective of the total dose, are thought to be most relevant. Medium doses are defined as doses in excess of 100 mGy up to one Gy and high
doses encompass doses of one Gy or more, including the very high total doses used in
radiotherapy (of the order of 20 to 60 Gy).
, BEIR 20052007 .
- 100 . -

55

.
100 BEIR-VII
.
, Low dose
: , 100 .
733 , -
200 NCRP-1980 19862000.
, ,
100
42 100 70
, , , , , .
100 , BEIR,
[AB10].
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1.5.2. (DOE Department of Energy)
, , . DOE
.
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[AB34, AD8],
1998 . DOE .
10- DOE
.
? , 0,1 [AB34, AD8]30:
The DOE Low Dose Radiation Research Program is funding basic research
to determine the responses induced by radiation exposures at doses of 10 cGy and below These projects help identify research needs and integrate the basic data into
useful models of radiation cancer risks.
-
? ,
100 ? 31:
What is a low dose of radiation?
A low dose can be classified according to its health impact or in comparison to
natural background radiation. Background radiation, over which we have no control,
comes from such sources as cosmic rays, radon, radium, and other radioactive materials in the earth and has not been shown to cause adverse health effects. This dose of
radiation is considered to be low. For the Low Dose Radiation Research Program, a
. [AD8] : Frequently Asked Questions about the DOE Low Dose Radiation Research Program [AB34].
31
[AD8].
30

56

low dose of acute low-LET (Linear Energy Transfer) radiation such as X rays, gamma
rays, or beta particles is defined as being less than 10 rem or 0.1 Sv. This level of exposure (less than 10 rem) has been at or below the limit of detection for most biological changes observed in past research and is twice the yearly occupation exposure
limit.
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1.5.3.
2006 .

57

2003 . (,
)

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.
( 2006 .)32:
UNSCEAR Report 2006. Non-targeted and delayed effects of exposure to ionizing radiation. Fifty-second session Vienna, 29 May to 2 June 2006. 101 p.
,
(20032005 .), : In contrast to the risks associated
with exposures to low doses of ionizing radiation (<0,1100 mSv)....
, 100 . ,
( 200 ),
1986 . 2000 . (. . 1.4.1),
,
, . (R. Cox)
. (W. Morgan)
. ;

G -2000 [AU13].
, , ,

2006 . .
2009 ., ,
[AU18]. 100 (
),
2000 . 0,2 [AU18]:
exposures to low doses of ionizing radiation (less than about 200 mSv, UNSCEAR 2000 Report).
1.5.4. (Harvard Physics Department)

Web-

[AP29, AP30].
100 .
,
(00,1 ) .
2003 . Epigenetic effects of exposure to ionizing radiation; 2006 .
32

58

33 (00,1 ).

(
.
..)

,

, 0,2 (..,
1986 .). ,
.
,
34, :
At the same time, with the assumption (based on the above mentioned data for Hiroshima and Nagasaki) that radiation carcinogenesis is absent for low doses (00.1 Gy), the
number of radiation induced cancers among emergency workers will become much less and
the group of radiation risk will actually include only emergency workers who received external doses more than 0.2 Gy (the 1986 emergency workers only). This conclusion can be of
major medical as well as social and psychological significance35.

.. (
) [2] (. 1.5.1).
, 0,1 .
,
0,1 ,
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[AP29] ( Little, 1998)

, (
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35
, ( ) , :
Yet, the data of the National Registry gathered to date provide evidence to an increase in cancer incidence rate among emergency workers with external radiation doses more than 0.1 Gy.
0,1 , ,
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( ) [2].
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59

1.5.1.
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60

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61

62

1.5.6.
-99 (
20042006 . 150200
),
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, ,
:
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1 , 0,1 ,
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. (As a rough rule of thumb, effective doses on the order of 1 Sv,
100 mSv, 10 mSv, 1 mSv, and 0.1 mSv will be called moderately high, moderate,
low, very low, and extremely low, respectively, in this report.) [AI7].
-99 20042006 .,
, .
in vivo , (..,
10 ), , ( in utero ).

[AL27, AL28, AL29, AP13].

:

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: 10 (), , 100 ,
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:

63

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[2]. 2006 . - 0,350,7 [8],
-2006 11,5 () [4].
2005 . ( ,
0,251 [5] 3 [3]).
,
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0,4
[2, 7, 2], 0,5 : 1996 . [6], 1998 . [7],
2000 . [9], 2001 . [8], 2003 . [1], 2005 . [4] ( 0,5
2005 . , ) 2006 . () [5].
1 [7].

2006 ., , 0,5 ,
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64

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70 ) .
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(, )
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0,51 [AM50]. ( )
2000- . [AM40,
AM41, AM43]:
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[AM41, AM44].
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65

. : . ,
2000 . [AS55]:
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0,1 [AS55].
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8 2005 . ,

[AD17].
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(moderate) , 1 10
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[AK8].

1.6.1. Radiation Research,

(, 2001 .) [AM27].

, ,
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0,5 .
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66

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(, 2325 2007 .)
.
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67

, 2007 . :
(~0,2 )... [2],
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15-

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[1, 2]).
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0,20,5 (. 26 [AU10]).
39
40

68

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. 1.6.2 1.6.3
1994 . [AU10] 1996 .41; [AS34].

Biological effects of low-level radiation. Forty-fifth session of UNSCEAR Vienna, 17 to 21 June

1996.
41

69

1.6.2. -1994 [AU10],

. [P33] [AP25]. ,
(observed)
(expected) 0,20,5 .

1.6.3.
. -1996. [S5] [AS34].

70

, 0,5
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71

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[7].
43
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72

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1986 . [AU5] 1988 . [AU7] ,
0,05 () .
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.

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[AU8],

73

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74

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75
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1993 . [AU8] (. . 1.7.1, 331),

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(...with emphasis on the lowest doses for which
meaningful effects have been found. Additionally, effects that may occur as a result of
chronic exposures over months to a lifetime at dose-rates below 0.1 mGy per minute,
irrespective of the total dose, are thought to be most relevant [AB10]).
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76

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77

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78

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79

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81

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[AW29]
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120
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125
2.4.1. ,
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Kadhim M.A. et
al., 2004

,
,

Wright E.G.,

2004

Mothersill C.,

Seymour C.,

,
2000; 2001
;

,
Prise K.M., 2006

,
Schofield P.N.,

1998

Morgan W.F.,

2002

.
Little J.B., 1998
2007

;
Brooks A.L.,

2005

Schwartz J.L.,

2007

Matsumoto H. et
al., 2007
Clarke R., 2000

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126
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127

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128
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129

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132

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133

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134

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135

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136

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140

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141

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142

143

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144

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145

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146

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148

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149

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150

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151

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156

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157

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160

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161

2005 . [AK26].
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162

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AH18, AH19,
AH20*, AK33,
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[AB30]
[AK2, AK3, AK7]
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[AK7]
[AK5]

1; 3 8 (
)
3 6
1 ,

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5 ,

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0,5 5
3

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[AP22]

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14
1,5
2
3 4
1,5
4,5
3

[AC3]
[AC15]
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AE8, AG14, AM29,
AR17]
[AE6]
[AC15]
[AS15]
[AP21]

[AU2]

[AB29]
[AO4]

163

2.6.1 (-1). in vitro ( ).

(,
)
HPV-G (HPV-,
)
**

hTERT
HeLa**

HT1080


(SW 48)

SCL-II

SiHa



(U1-Mel) cells

CGL1 (HeLa
)

[AD30]

0,5; 1 3
(

0,5
)
16
115 (
)
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)**
0,55 (
)
2

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6
3
412 (
)
9
48 , 2
6
0,55

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18 (
)
0,550 (
)

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4,5 6,0

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AM21]
[AL7]

5 7 (
)

[AO6]
[AL39]
[AL35, AL36,
AL37, AL38,
AM39, AS21]
[AC14]
[AO3]

[AB36]
[AS59]
[AS62]
[AC14]

[AM2]
[AL6]
[AY2]

164

2.6.1 (-2). in vitro ( ).

GM10115

HT1080

C3H10T(1/2)
BALB/3T3

EF42

4T1


,
H2AX

( m5s)

V79

110
(

)

0,10,5 ;

110
15
5 10
7,513 (
)
10
10 25 (
70C)
4

[AN1, AN2, AN3]

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[AD6]
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AL15, AL16,
AP24]
[15]
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4 6
8
110 (
)
1

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[AC15]
[AG9]

46

[AL8]

1,5

[AB5]

4 6
6
0,5 6

[AM53]
[AT6]
[4, 1, 6,
7, 8, 9]
[AD2]
[AC15]
[AJ3, AM5, AT7]

4
8
112 (
)

[AS17]

165

166

2.6.1 (). in vitro ( ).

scid

CHO**

S. erevisiae

2 6

110 (
)
212 (
)
814
250400 [sic!]
(
)

[AU19]
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.

,
,
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.

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) [AB10, AU12, AN16]. 2004 . [12, 13]
, ,
.


. 2.2

,
,

. , . 2.6.1 ,
. ,
. 2.6.1 ,
. 2004 . ,
, ,

167

in vitro 0,10,2 -.
( 2.9),
, ,
TK6
( ),
HPV-G (
. ) HeLa [2, 7, 1, 5,
6, 8], ,
[1, AC10, AL5] (. 2.2),
[6].
.. .
, [12, 13, 18, AK26],
.. HeLa
,

[1, 4, 5, 6, 8, 10] (
, ).
,
(Non-targeted and delayed effects of exposure to ionizing radiation) [AU17, AU18].
,
( 2006 2009 .).
.. HeLa
[2, 4, 7, 1, 5, 6, 7, 8]. Pelevina
( et al.).

, 2004 . in vitro
,
?
J, 2007 . . 2007 .
( CHO)
[AS18]. , ,
, ,
: CHO 10
50 4 11
9 , .., 23 .
10 [AS18].
,
[1, 2, 6, 13, 3, AL4, AV5, AW20, AW21],
. ,
1050 in vitro [13, AW20, AW21] 100 in vivo [1, 2]

, ..,
- .

168

,
, :
.

169

2.6.2. , ,
.
,
[18], :
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, ,
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.
, - , .

, ,
[18, AK26], ,
, BEIR BEIR-VII
[AB10], [AI7].
, , ,
.
[18]. ,
, , , -
- -
. : ,
, , .
2004 . [12, 13], 2005 . [AK26] 2006 . [18].
, [16] [17]

, .
,
( . 2.4). ...
( ). V (2006 .) ..
, ,

HeLa ( ).
, 2004 .,
, ,
.
[AK26] [18] .
-
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,
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, , -,
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20- , -,
.

170

XX , , ,
. ,

.
. 2.6.1: -
, , in vitro, ..
. ,
:
1) 0,5

[AM13].

, ,
,
0,5 .
2) 2
[AS59].
3) GM10115.
[AL10]
(0,1; 0,25; 0,5 ),
.
(. 2.6.1.), , [AL10].
(. . 2.6.1), 0,10,5
.
4) ,
( HPV-G).
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. ,
(
) [AM39, AM46, AS23],
0,5 [AM38].
5) . ,

. ,
1 ( ) [AO4]
.
. 2.6.1 , HF-12
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) 1,5 ,
HF-19 [AK5]. [AK5] (Kadhim M.A. et al., 1998) ,
,
HF-12 HF-19, , .
- : ,
, (.
2.4). , . : ,
HF-19, -
, ?

171

2.6.1.
GM10115 [AL10].

, (Lawrence C. Dugan) (Joel
S. Bedford), 2003 . [AD28].
, AG1521A, 45
, .
,
. ,
, ,
.

.
() , ,
2025 .
. . [AG14]
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106. 2 :
,
,
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106

172

, ,
[AD28], 12%
[AG14].
.
. ,
.
:
, .
.
G0 (5 ;
3,9 /; 137Cs) (1,25 ; 56Fe; 1
) . 10%- .
;
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, 107. . 2.6.2
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(apparently simple symmetrical interchanges; FISH), .., .
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AG1521A
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[AD28]. , ,
56Fe : , ,
. [AD28] ,
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.
107

173

2.6.2.
- 5 [AD28].

, [AD28] - ,
, , , 0,2 , 5
. , ,
, ,
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- 40
, 150200 [13]. , ..
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[AD28]. .
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. [15] (
. ..): ,
: (
)....
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, [15]
? 2006 .

in vivo
[AK26]. 0,10,2 .
,
( , -
),
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(. . 2.6.1 [18, AK26]).

, ,
.
. [AD28]. ,

174

. [AD28]
[]. .
. ,
( ), -
, :
BALB/c, TK6,
()
. ,
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AB28, AW16].
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,
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:
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, :
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AG1521A.
( 5) (Kadhim M.A. et al., 1998) [AK5],
,
HF-12, [AK5]
.
. . [AD28], , ,
, ,
.
2003 . [AD28],
( ), ,
, .
. 2.6.1,
20042006 ., ,
in vitro

175

, ,
, , .
. . : ,

.
2.6.3. in vitro
,

. ,
- in vitro
, .
,
( ; ; ).
, ,
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-,
.
.
(
[42]) ,
, .
,
,
(, , HeLa,
[4, 5]),

( )
[1]. (
.)
, , .
(
) [1]. . 2.6.1,
, in vitro,
,
.
. 2.6.1
, . , (longterm) in vitro , , ,
: (,
), , .
, - ,
, .

176

,
.
, G0.
.
,
,
in
vivo, .

in vitro. , , -
(1996) [AA8].
, [AS31],
.
,
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). ,
.
. -
, , ( ..):
,
, -,
,
in vitro. ,
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,
.
, ( )
.. [AS31].
, , ,
[AB10].
, 2.3.3,

, ,
,
. ,
,
, ,
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in vitro (
) :
, , , (
),
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. 2.3.3).

177

, . , ,

.
, ,
, .
,
, , , ,
, ,
.
, ,
in vitro -
( , [42]).
, 2.6.2,
. ,
, ,
.
,
in vitro ,
, .

, in vivo, in vitro [26, 27]. ,

[24, 30,
AK28] [10, 11]. ,
,
,
.
, , 1993 .,
.
,

( ), .
, ,
, ,

.
, 1990- ., . ,
, ,
- in vitro .
,
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(
), . .
, 1993 ., ,
, ,

178
- in vitro
, . ,
, , ,
, [11].

, :

, ,
in vitro.

in vitro ,

.

,
in vitro ( )
in vivo,
.
, . 2.6.1,
in vitro ,
, .
, 0,5 .
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, in vitro [5]108; (. . 2.1.1)
// :
. ,
. 2.6.1, ,

.
. . ,
[AD28] (.
2.6.1). , ,
:

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, ,

, . (
. 2.6.1.)

-,
.
,
, ,
,

in vitro in vivo [12].
108

179
. ,
(
) ,
(. , [AB10, AU13, AU14, AU16]). , ,
,
.
,
, , .
;
.
, ,
,
, .

, , , . ; -
(
) , .
. , , ,
, ,
.
[8, 39, AC1, AF5, AH11, AL33, AL34, AP16, AS46].

2.7. in vivo
2.7.1. in vivo.
, ,

,

(. 2.2 2.3.4),
in vivo. ,
, , ,
, . ,
- ,
,
, .
? , : ,
,
.
,
[3]?
,
. ,
[AL26] 72-
, 40
.

180

. , 30%
, .
,

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(..,
).

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[AL26]
in vivo.
, [AH5]
( )
,
.
.
, ,
[AH5]109, . , [AH5]
:
-,
,
,
.
, ,
,

,
() .

,
[11, 20, 22] [11, 12, 13, 2, 3,
4]
.
,
.
- , ,
[2]110,
(Non-targeted and delayed
effects of exposure to ionizing radiation) 2006 . [AU17].
110
,
'

( , 0,5 ). [2]
.
109

181

? ,

? , ,
, ,
. ,
[AK31].
-2000 [AU15]:

- [AU15]. ,

.

.. [3].
2007 . ,
,

() .. .. [3].
,
- . [3], ,
( . ..):

,

( ),
de novo, ..
( ).
,
[10].
. [3] ..
..

- 111.
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11, 12, AV4] ( [AS47])
,
2007 . [3]
,
, (. 2.7.1).
(
80 ) ,
.. [AT2].
111
45- . ,
, -
, , - ( ) [3].

182

. 2.7.1 [3] ,
,
. ,
.

2007 . [3], , 145
112
, ,

.
2.7.1.
[3] ( ).

,
2007 . ,

(..,
; . 2.2 2.3.4),
,
.
,
[3, 6]
- , , ,
.
.. [3].
.. [3] -2000 [AK31, AU15].


, :

112

, .

183

[AL2].

[1].

(. [AS10,
AS44]).

[AH22].
(
) [AL3].
[AL3].

(fragile sites) [AB3].
[AR6].
in vivo ([AL3]113, [AQ1]114, [AI3]115, [AC7]116,
[AN21]117)
.

[5, PC23, AI3].

[AE2].

[AL3].

, ,
.
.
NEWSru.com ( ) 12 2005 .
(- . -)118.
.
, .
, ,
.
... ,
, , , , ,
, , , .
114
.
115
.
116
.
117
.
118
(http://www.e1.ru/news/daily/2005/08/12/news_id-236570-section_id-9.html;
http://www.intermed-spb.ru/news.php?id=1068&limit=77).
113

184
, 34%
,
, , , -.
,
(,
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, .
,
.

185

2.7.2.
in vivo
- , in vivo
?
. :

in vitro


( ).

in vivo , ,
,
in vitro , , in
vitro .
, ,
in vivo.
[AB28],
. .
2.6.3
.
.
in vivo ,
.
, :

,
,
( in utero).

(
).

.
,
,
(

[8, 5, 6, 7, 2, 19,
22]). , ,
, 2005 . International Journal of Low
Radiation [AK26],
2004 . (
).

(),
, (),
[AK26].

186

(
)
.

[AK26] (. 2.7.1; ).

2.7.1.
( )
. (Koterov A.N., 2005) [AK26].

, . 2.7.1,
0,10,2 . 0,25 (
: ,
) (.
, . 2.7.1).

187

, 1.6 2.4.
,
(
)
0,5 .
(. . 2.7.1).
,
.
[AK26],
[AU16, AC12].
.
,
, -, in vivo
, , -,
in utero.

() - (
in vivo)
[AK26] ( 50- ).
.
2.7.1, ,
, .., ,
,

.
, (
) -
; 0,10,2
[AK26, AK27].
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.
in vitro in vivo.
in vitro
. ,
in vivo ,
, ,
- . ,
G.E Watson, S.A. Lorimore E.G. Wright [AW5, AW6],
. [AW5, AW6], -,
in vitro - ,
, ,
,
. ( ,
.)

188

( ) ,
-
? ,
1996 . [AW6], : ,
. , ,
[2] [7], ,
, .
.
Watson G.E et al. [AW5, AW6] ,
- in vitro in vivo,
. 2001 .
[AW8].
.
.. (S.D. Bouffler)
[AB28],
226Ra in vitro, in vivo.
in vitro in vivo
100
[AB28]. ,
Watson G.E et al. [AW5, AW6, AW8],
. ..
, - in vivo
,
., , ?

. :
, . ,
, ,
, ( ,
in vivo ).

( ) 50- 100-
.
, , ,
, , ,
.
( 100 ,
) [AB28]. [AB28]:


.

,

.

189

, ,
. 2000- . [AW8]
, in vitro, ,
, in vivo.
, ,
.
in vivo . 2.7.2. ,
.
,
.
(
), .
in vivo
in vitro. . 2.7.2 ,
-, ,

[AS7].

,
. 2.7.2. -,
, -, , ,
( ),
() .
. 2.7.3 [AS7].
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,
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,
( 642 ). ,
, ,
(. 2.7.3). : %
, ,
( ),
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.
[AS7] ,
, ,
in vitro
.
.
,
,
[AS7].

190
2.7.2. in vivo ( ).

in vit in vitro ro
+

in vivo
in vivo
C3H 10T/2
,

in
vitro

C3H

CBA/H

in vitro

in
vivo

in
vivo
24

in
vivo, in vitro

[AW8]

;
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in vitro)

[AM1]

[AP4]

191

()

.
;

(
)

[AS7]

0; 0,1; 0,9
1,1

192
2.7.3.
[AS7].

1
(
)
2
(
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3
(
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4
(
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()

(%)

(%)
(%)
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1,9

0,9

88,7

1,1

85,2

8,2

3,0

1,6

2,0

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93,5

8,2

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2,6

3,5

86,7

4,8

1,9

2,9

86,2

10,3

1,7

3,5

75,9

9,3

7,4

7,4

[AS7]
,
,
, .
,
[33].
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. ,

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), ,
,
, , .

193

.
.. .. ,
T- ,
(,
)
[13].

, .
, [13] :
43 ,

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, ,
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. ,
[13] :

... .
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. 24
.
, ,
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, ,
, 1990 .
. -, ,
.
( ,
in situ). -, (
), in situ,
, [33]119. ,
, .
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in vitro (
), [AM23].
/
T-
,
[33]
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2,5 ),
.
119

194

. ,
,
, T- , ,
, [13]?

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(,
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.
,
/
.

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2.6.3
. ,
,
, , ,
. , ,
.

[AS30, AS3] (. 2.3.3).
,
, ,
in vivo.
,

. ,
,
, [10, 11, 12, 3,
AS47, AV4]. ,
(.
2.1). ,
()
,

195

-H2AX [AO4, AY3]

(PKcs) [AO4].

, , .
, 1960- 1970- .

[AS6, AB12].
[AD28]
,
AG1521A,
.
, 45- ,
, . 10-

12,6 120. 10-

,
0,03 , 45- , ,
0,11 .
, , ,
.
, .. ,
(10- ) . 3040%
[AD28].
[AD28],
(
- ). ,
, ,
[AD28].
, in vivo
,
, , ..
[AB28] (. ) .
,
,

?



,

?
:

,
( 56Fe; 1 )
45- 3
120

196

,
[AD28].

, ,
,
,
, [AS7]121.
, in vivo
in vitro ,
, .

[AS7],
,
. .
121

197

in vivo
in vitro. ,
, in
vitro in vivo.
, , .

in vitro,
in vivo
, ,
- - [4, 5, 7, 8, 19, 21, 22]122.
.. . ..
() , ,
, , ,
,
in vitro
10 100 , [22].
2006 . (. 2.7.2).

2.7.2. in vitro ,
. [22].

()
- in vitro [19, 21].

[10], ,
.
,
, .
,
, .
122

198

(. 2.7.2),
.. ,
[19, 20, 21, 22],
:

. -

, ,
, .
,
(
,
1986 . 1998 . . . 2.7.2).

, (
), (
2.8).

. ,
..
.

,
, , ,
(
, .
2.2 2.3.4).

, ,
.. [22],
,

..
,
, , ,
[AB10, AC12, AU16],

.
, , in vitro
[AB19] [AB2, AM15]
.

.. [22]
. ,
[22] , -

199

: ,
, , ,
.. .
, ,
.
, .
,
.. [22] (. . 2.7.2), ,

,
. in
utero, (
), ,
1,51,7 (
). .. . [22]
,
.
,
..

,
.. ,
[19, 20, 21, 22],
. . 2.7.2
.
, 2.4.4.
***
,
,
.. -
(-).
1992 . [5],
1995 . [7],
[8],
[4, 8] (..,

, ).
.. ., 1992 [5]
in vitro ,
, ,
. , - in
vitro
(
). , , ,

200

. 1992 . .. .
[5] ,
, 2002 .
..
[4]:
,
( 1992 . [5]).
2002 .
, .. ,
[4]:

.
,

( 1993 [AU9] 2001 [AU16], COMARE 2002 [AC12],
BEIR-VII [AB10]), .. 2002 .
[4] ,
, ,
[6].
[4, 8]
,
(),
in vitro [AB19] [AB2, AM15] ,
(),
, .. . [5, 7].
,

[AB2, AB19, AH15, AL9, AM15, AT3, AW18].
, ,
.. -
in vivo ,
.
,
[20, 22, 2, 19, 8, 5, 4, 6, 2]
[7]123 , ,
[AB10, AC12, AU9, AU16].

in utero.
2.8. in utero
, , ,
.
2003 .
[7].
123

201

,
. in utero ,
, , .
. 2.8.1.
(A. Stewart)
in utero (
) [AS48].
.
1,4 0,010,02 ([AB17, AB20,
AD18] (. [AC12, AU14]).

1,8 [AS49]. ,
in utero
0,2 [AS32]. -,
in utero .
, ,
.
,
.
, , , .
. 1992 . [AS49]
, 10

.
in utero [AS32].
, 2006 .,

[AM14],
. :

( ) ,
;

( );

( ),
;

202

;
-. [AM14] ,
, , ,

.

203

2.8.1. in utero
( ).

ICR CD-1

1-3-
*

,




,

;

Swiss
albino

912

CBA/Ca

36

0,06; 0,12;
0,241.88

1 2

[AW13,
AW14,
AW15]
[AP1, AP2]

[AD12]

0,251,5

[AD10,
AD13]

0,704,2 [AA1]
;

1
[AR11]
CBA/Ca
()

( )
0,5
[AK30]
CBA/H

, 0,095,22 * [AD19]

(6-) *

;

0,11,5 **
[AD11]
Swiss albino
F1

204
2.8.1 ().
in utero ( ).

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1
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,

205

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[AG15]. ,
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( 2008 .) ,
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206

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. BALB/c
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[20],
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AC15, AD28, AE5, AE6, AE8, AG14, AM29, AR17]. 2003 .
(
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207

NH32 WTK1, 0,1; 1,0 2,0 .
NH32 (
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. 2.9.1).

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TK6
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208

TK6
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to 2
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209

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210

(20042009) [AU17, AU18],
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[AW25]).
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125

211

1) HPV-G a priori
(.
2.6.3 in vitro).
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,
, . 0,010,05
[AS23] 3438%.
.
, ,

M.C. Joiner B. Marples [AJ6, AJ7, AM7, AM25].

, ,
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3) HPV-G
. , 2000 .
[AS23] 0,01; 0,03 0,05
3438%, 2002 . [AM46]
27% 0,01 0,05 .
, 0,010,1
[AS23].
. . 2002 . [AM46],

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212

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[AM49],
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2000 . [AS23] 2002 . [AM46]. .
(platting efficiency)
; . ,
2000 2002 .
, .

5)
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( 0,5 and 1,0 ).

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(110 ),
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7) ,
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) .

213

SW48 ,
[AM47, AS21]. GM10115
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[AM39].
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involves too few cells to compromise tissue function). [AM39, AS23]
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1.2 . 1.2.1). ,

214

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. 2.9.3),
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, , 5
[AM59].
2.9.3. HeLa
HeLa ;
[4, 5].

215

in vitro (. . 2.6.1),
, .
..
-.
2.6.1, [12, 13, 18, AK26]
.. . HeLa
,

[1, 4, 5, 6, 8, 10].
.
19851987 . .. HeLa
,
.
[9, 2, 3],
1986 .
:
, [3]126. -
.. 6 [9, 2,
3].

HeLa ,
. :

[9].
1991 . ..
HeLa - [7].
12- ,
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.
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, in vitro
.
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HeLa ,
, [1, 4,
5, 6, 8, 10].

2.3.1,
.. , , .
126

216

1992 . (4
; 1114 ) HeLa, 0,1; 0,2 0,4 ,
[2] (. 2.9.4).

217

2.9.4. HeLa.
(1)
( ) (; 2) ( .,
1992) [2]. 14- 1214- .

218

19951996 . HeLa, ,
[3, 6] (. 2.9.1).
2.9.1. HeLa 10-
( ( .. ., 1996) [6]).

. 2.9.1 ,
3- 10-
( , ).
2003 . [3, 6]
- [1] (. 2.9.2).
2.9.2. HeLa 10-
( ( .. ., 2003) [1]).
, , [3].

. 2.9.1
2.9.2
10- (1996 .) [6], , 3- 7- (1995 2003 .) [3, 1]. , ,

219

; , ,
.
. . , (. 2.9.3).
. 2.9.4 . 2.9.1 2.9.2, ,
, in vitro
. ..
-
2001 . [8], 2003 . [1] 2006 . [5].
19911992 . [2, 7], ,
, 19951996 . [3, 6], , .
, , .
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, .
[AS18] ,
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,
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,
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220

2.9.5. 416-
10 50 CHO [AS18].
. (
[AS18]
- .)

221
[AS18]:

. ,
, HPV-G , CHO
-
(0,112 ) - (Holmberg et al. 1998, Bortolleto et
al. 2001, Mothersill et al. 2000, Little et al. 1997, Ponnaiya et al. 2004). (A great variety
of papers have reported the induction of chromosomal instability by low dose radiation exposure...)
, ,
0,10,2 . (Holmberg et al.,
1998) [AH20] (Bortolleto et al., 2001) [AB25]
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20002002 .
( HPV-G), 510 [AM39,
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23 ,
[AS18]. , ,
1050

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-
[2].

222

. ( V79)
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-
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, CHO,
. . .
2.9.5. BALB/c
-
.
BALB/c
(Radiation-induced chromosomal instability in BALB/c and
C57BL/6 mice: The difference is as clear as black and white) [AP22].
in vivo
.
1999 . .. .. (R.L. Ullrich, C.M Davis)

in vitro,
BALB/c, in vivo 0,1; 0,25; 1,0 3,0 (
). 0,1 1,51,8 [AU1]
( . 2.4.1 ; 2.4.2).
, .
.. ()
F1 BALB/c,
- 0,1; 0,25 0,5 .
0,1 AP-PCR,
, .
AP-PCR .
( -) 0,1
13%, 0,25 0,5 37% 38% [3,

223

AV2]. , ..
(Yu.E. Dubrova),
F1 ,
0,5 [AD21, AD23, AD25, AD26].
,
.. .
..
F1
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0,1 30% [1].

BALB/c , , .
, ..
. 0,5 [4].
, BALB/c
. ,
BALB/c, HPV-G,
,

, ,
.
[18, AK26], ,
1989 . [AO1] in vivo127,
[AO1].
,
.
0,05; 0,29 1,73 ,
.
.
..
, ,
0,05 [AO1].


, .
, 1989 . [AO1] ICR CD-1
.


[AE1, AH8, AN13, AM4].

20042006 . [AU17]
2009 . [AU18].
127

224

2008 .128 (. . 2.6.1 [AK26, AK27]). , 12

(Low Dose Radiation Research of Department of Energy USA) [AD8]. Medline
.

,
.
128

225

2.10. (,
BEIR )

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226

, , .
,
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(.
. 2.9.1 2.9.2). . .
(J.B. Little) 1997 .,
412 [AL23].
HPRT [AL23].
[AG9],
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[AS18]), [AG9] .
, ,
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.
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(20042009 .) [AU17, AU18].

(27 ); , . ,
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[AU17, AU18].
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[37] (. 373).

; ,
TK6 [AS16] (. . 2.9.1)
[AM39, AM46, AS23] (.
. 2.9.2).

227

,
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. 1990- 2000 . ( ) [AM36, AM40, AM41, AM43] :
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228

(Non-targeted and delayed effects of exposure to ionizing radiation), ,
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[AU17] 2009 . [AU18].

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229

in vivo,
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230

BEIR-VII [AB10].
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inconsistencies) .

( -99).
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. (The data did not reveal consistent evidence for the involvement of induced genomic instability in radiation tumorigenesis, although telomere-associated
processes may account for some tumorigenic phenotypes.)
:
,
,

. (Knowledge on adaptive responses, genomic instability, and
bystander signaling among cells that may act to alter radiation cancer risk was judged
to be insufficient to be incorporated in a meaningful way into the modeling of epidemiologic data.)

231

, -
, BEIR-VII (2006 .) , ,
, ,
( 100 ).
:
,
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( 100 ) . (Until molecular mechanisms responsible
for genomic instability and its relationship to carcinogenesis are understood, extrapolation of dose-response data for genomic instability to radiation- induced cancers in
the low dose range <100 mGy is not warranted.)
, ,
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low total doses or low doses per fraction suggest that the dose-response relationship
over a range of 20-200 mGy is generally linear and not affected significantly by either
an adaptive or a bystander effect. No data are available in this dose range for radiation-induced genomic instability.)
' . 2006 .
BEIR ,
.
,
129

232

( 0,10,2 ). , ,
. , , BEIR-VII.
,
, in utero.
,
,
, BEIR-VII
[AB10]:

in vivo,

, ,
-
.
(Based upon the negative or inconsistent data on in vivo induced genomic instability in bone marrow cells, the non-sharing of genetic determinants, and the contention on data regarding A-bomb leukemias it is judged that induced genomic instability is unlikely to impact appreciably on the risk of lympho- hematopoietic tumors
after low dose radiation.)
BEIR-VII, ,
. , . ,
/
, (...it may be that genomic instability plays a more
important role in tumor progression than in tumor initiation).
, BEIR-VII, , -99,

. , BEIR-VII
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[5, 2], [7, 4] [7, 8, 9, 10, 11].
2010 . BEIR-VII [AD4a]
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233

.
, ,
, .
(There are studies that did not find evidence of radiation-induced genomic instability when blood chromosomes from normal, healthy workers were examined.
Certainly, one would not expect normal, healthy individuals to show detectable levels
of chromosomal instability regardless of whether they had been exposed to radiation.
Not unexpectedly, in those instances where exposed individuals showed convincing
evidence of instability, they also had diagnosed disease, for example, chromosomal or
microsatellite instability in acute myelocytic leukaemia and myelodysplastic syndrome patients among the Japanese A bomb survivors. This, however, raises the question as to whether the observed instability is a consequence of the disease, a non-targeted effect associated with the irradiation, or both.)

[AD4a].
(2004
2009 .) [AU17, AU18]. , ,
,
.
2004 .; 2009 .130
, 2004 . 2009 .
.

(
, ) in vitro in vivo.
, ,

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,
130

234

.
[AU17,
AU18] . , , , ,
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in vivo,
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dose and target size).
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in vitro, (2004)
(2006) ( 2005 .
2004 .;
2009 ., [AU18]).
. 2.10.1 2.10.2.

131

2009 . [AU18] .

235

2.10.1.
in vitro 2004 .
( 2005 .).
.

236

2.10.2.
in vitro 2006 .
.

237

2.10.3.
in vitro
2006 . ( 2009 . [AU18]). ,
2009 . .

238

. 2.10.1, 2.10.2 2.10.3 ,

2004 . 2009 .,
. 20042005 .
( ,
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. 2.10.1). 2006 2009 .
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239

, , ,
.

, ,
Medline,
.
.

. , in vivo
,
.
- ( ) ,
( ,
[AC12]), 2001 . .. (U.S. Weinberg) [AW12].
,
[AW10, AW11, AW12]. 2001 .

. . (A.J. Jeffreys) .. (Yu.E. Dubrova) [AJ4].

.. [AW12],
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: Questions of ensuring paternity, sample mixup...). in vivo ,
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240

, -99 BEIR-VII,
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BEIR-VII (2006 .), , ,
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241

,

(, , BEIR;
. ;
2.10)132. , ,
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;
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133
.. .
// . . . 1999. . 39. 1. . 2634;
. .: .. (.).
: .
, ., 1996. . 149182.
134
..
. .: . 1998. 100 .
132

242

,
,
1999 ., , 1997 .135. 2001
2005 .
.
2003 . I
WONUC ( ,
- )
136.
WONUC ( )

.
2005 .
International Journal of Low Radiation ()
137.
.
, ,

, .

, ,
,

.
,

,
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,
, , ,
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.
:
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.., .. in vivo // . .
. 1999. . 39. 6. . 648662;
// . . . 1999. . 71. 3. . 1325.
136
I
(WONUC). .. . .: , 2003. 458 .
137
Koterov A.N. Genomic instability at exposure of low dose radiation with low LET. Mythical
mechanism of unproved carcinogenic effects// Int. J. Low Radiation. 2005. V. 1. 4. P. 376451.
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243

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244

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245

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