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5. Cotman C. W., McGaugh. Behavioral neuroscience. N. Y. Academic Press. 1980.
6. S. Ramon y Cajal. The structure and connexions of neurons // Nobel
Lecture, December 12, 1906.

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// Annual Review of Psychology, 1996. V. 47. P. 1-32.
7. Singer S. J. & Nicolson The fluid mosaic model of the structure of cell
membranes // Science, 1872. V. 175. P. 720-731.
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1. . . //
. . 1974. . 5. 2. . 5-93.
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// / . . . . .:
- . 2006.
3. . . // XXI / . . . . ., 2004. . 39-85.

1. ., ., ., ., ., .
. .: , 1986. .. 1-5.
2. . . . .: , 1989.
3. . . // . 1995. . 16. 1. . 26-30.
4. . . //
2004. T. 21. 1. C. 5-14.
5. . ., . . : // / . . . . .: - , 2005. C. 119-157.
6. . . - // . 1996. .: -. . 23-47.
7. . . . .: , 1975.
8. . . . . // . . 1980. 4.
.185-187.
9. . . . .: . ., 1991.
10. . . // :
/ . . . . .: . 1986. . 229-240.
11. . .
. .: - - , 2004.
12. . . . .: , 1990.
13. .. //
. 1998. . V. 1. . 60-67.
14. . . // . .: . 1960. . 1. . 93-141.
15. . . : // . . . . 2000. . 50.
5. . 743-774.
54

16. . , ( ) . ., 1859.
17. . . //
. 1997. 6. . 28-32.
18. . . .
.: - . 1993.
19. . . .
. .-.: . - . -. 1934.
20. . ., . . : // , , : / . . . ,
. . . .: , 2008. . 133-160.
21. . . , , //
. .: , 1982. . 47-85.
22. . . .: , 1975.
23. . . .: . 1980.
24. . -
. .: . 1995.
25. . . . .
//
: . . .: , 1999. .11-37.
26. . . .
: . . . .1998.
27. . . .: . 1983.
28. . . :
// . . . . 1990. . 26. 5. . 733-740.
29. . . . .: . . . 1922.
30. . . . : . 1997.
31. . ., . . // . . . 1992. . 112. 2. . 265-272.
32. . . . .:
. 1966.
33. . ., . ., . .
// . .: . 1987. . 319-352.
34. Alexandrov Yu. I., Grinchenko Yu. V., Laukka S., Jarvilehto T., Maz V. N.,
Svetlaev I. A. Acute effect of ethanol on the pattern of behavioral specialization of
neurons in the limbic cortex of the freely moving rabbit // Acta Physiol. Scand.
1990. V.140. P.257-268.
35. Alexandrov Yu. I., Grinchenko Yu. V., Laukka S., Jarvilehto T., Maz V. N.,
Korpusova A. V. Effect of ethanol on hippocampal neurons depends on their behavioral
specialization // Acta Physiol. Scand. 1993. V.149. P.429-435.
55

36. Alexandrov Yu. I., Jrvilehto T. Activity versus reactivity in psychology

and neurophysiology // Ecological Psychology. 1993. V.5. P.85-103.
37. Allsopp T. E., Fazakerley J. K. Altruistic cell suicide and the specialized
case of the virus-infected nervous system // Trends in Neurosciences. 2000. V.
23. P. 284-290.
38. Baily C. H., Kandel E. R. Structural changes accompanying memory storage //
Annual Review of Physiology. 1993. V. 53. P. 397-426.
39. Bar M. The proactive brain: using analogies and associations to generate
predictions // Trends in Cognitive Sciences. 2007. V. 11. P. 280-289.
40. Bonner J. T. The evolution of complexity by means of natural selection.
Princeton: Princeton University Press. 1988.
41. M. Bota, Swanson L. W. The neuron classification problem // Brain
Research Reviews. 2007. V. 56. P. 79-88.
42. Buzsaki G., Kandel A. Somadendritic backpropagation of action potentials
in cortical pyramidal cells of the awake rat // J. Neurophysiol. 1998. V. 79.
P. 1587-1591.
43. Chang J.-Yu., Sawyer S. F., Lee R.-S., Woodward D. J. Electrophysiological
and pharmacological evidence for the role of the nucleus accumbence in cocaine
self-administration in freely moving rats // The J. of Neurosci. 1994. V. 14. P.
1224-1244.
44. Edelman G. M. Neural Darwinism: The theory of neuronal group selection.
N. Y.: Basic, 1987.
45. Edelman G. M., Finkel L. H. Neuronal group selection in the cerebral
cortex. Ch.22 // G. M. Edelman, W. E. Gall, W. M. Cowan Eds. Dynamics
aspects of neocortical function. N. Y.: Willey. 1984. P. 653-695.
46. Ellis R. D. Dynamical systems as an approach to consciousness: emotion,
self- organization and the mind-body problem // New Ideas in Psychology. 1999.
V. 17. P. 237-250.
47. Engel K. A., Fries P., Singer W. Dynamic predictions: oscillations and
synchrony in top-down processing // Nature Rev. Neurosci. 2001. V. 2. P. 704-716.
48. Fiorillo C. D., Williams J.T. Glutamate mediates an inhibitory postsynaptic
potential in dopamine neurons // Nature. 1998. V. 394. P. 78-82.
49. Fisher K. W., Bidell T. R. Dynamic development of action, thought and
emotion // W. Damon and R. M. Lerner (Eds.) Theoretical models of human
development. Handbook of child psychology. 6th ed., Vol. 1. 2006. New York:
Wiley. pp. 313-399.
50. Fodor J. A., Pylyshyn Z. W. Connectionism and cognitive architecture: A
critical analysis // Cognition. 1988. V. 28. P. 3-71.
51. Forti L., Cesana E., Mapelli J., DAngelo E. Ionic mechanisms of autorhythmic
firing in rat cerebellar Golgi cells // J. Physiol. 2006. V. 574. P. 711-729.
52. Freeman W. J. Three centuries of category errors in studies of the neural basis
of consciousness and intentionality // Neural Networks. 1997. V. 10. P. 1175-1183.
53. Fregnac Y. A tale of two spikes // Nature Neurosci. 1999. V. 2. P. 299-301.

56

54. Guillery R. W. Relating the neuron doctrine to the cell theory. Should
contemporary knowledge change our view of the neuron doctrine? // Brain Res.
Reviews. 2007. V. 55. P. 411-421.
55. Jordan J. S. Recasting Deweys critique of the reflex-arc concept via a
theory of anticipatory consciousness: implications for theories of perception //
New Ideas in Psychol. 1998. V. 16. P. 165-187.
56. Klopf A. H. The hedonistic neuron. A theory of memory, learning, and
intelligence. Washington: Hemisphere publ. Corporation. 1982.
57. Koch Ch., Laurent G. Complexity and the nervous system // Science,
1999, 284, 96-98.
58. Koshland D. E. The bacterium as a model neuron // Trends in Neurosciences.
1983.V.6. P.133-137.
59. Koyama T., Kato K., Tanaka Z., Mikami T. Anterior cingulate activity
during pain-avoidance and reward tasks in monkeys // Neurosci. Res. 2001. V. 39.
P. 421-430.
60. Kuo Z. Y. The fundamental error of the concept of purpose and the trial
and error fallacy // Psychological Review. 1928. V. XXXV. P. 414-433.
61. Matsuzaka Y., Picard N., Stric P. L. Skill representation in the primary
motor cortex after long-term practice // J. Neurphysiol. 2007. V. 97. P. 1819-1832.
62. Milburn N. Falling stereotypes and new cell models in neurobiology //
Amer. Zool. 1990. V.30. P. 507-512.
63. Leist M., Jaattela M. Four deaths and a funeral: from caspases to alternative
mechanisms // Nature Rev. 2001. V. 2. P. 1-10.
64. Lewontin R., Levins R. Dialectics and reductionism in ecology // Synthese.
1980. V.43. P. 47-78.
65. Nelson S. B., Sugino K., Hempel Ch. M. The problem of neuronal cell
types: a physiological genomics approach // Trends in Neurosciences. 2008. V.
29. P. 339-345.
66. Nicola S. M., Deadwyler S. A. Firing rate of nucleus accumbens neurons is
dopamine dependent and reflects the timing of cocaine-seeking behavior in rats
on a progressive ratio schedule of reinforcement // The J. of Neurosci. 2000. V.
20. P. 5526-5537.
67. NSF Task Force. Newsletter of the Animal Behavior Society, vol. 36 (4).
. de Waal, 1996.
68. Pape H.-C., Pape D., Driesand R. B. Two types of intrinsic oscillations in
neurons of the lateral and basplateral nuclei of the amygdale // J. Neurophysiol.
1998. V. 79. P. 205-216.
69. Quian Quiroga R., Kreiman G., Koch C., Fried I. Sparse but not
Grandmother-cell coding in the medial temporal lobe // Trends in Cognitive
Sciences. 2008. V. 12. P. 87-91.
70. Ramus F. Genes, brain, and cognition: a roadmap for the cognitive scientist //
Cognition. 2006. V. 101. P. 247-269.

57

71. Ranck J. B. Studies on single neurones in dorsal hippocampal formation

and septumin unrestrained rats: I. Behavioral correlates and firing repertoires //
Experimental Neurology. 1973. V. 41. P. 461-555.
72. Rose C. R., Intracellular Na+ regulation in neurons and glia: functional
implications // The Neuroscientist. 1997. N 3. P. 85-88.
73. Raoul C., Pettmann B. , Henderson C. E. Active killing of neurons during
development and following stress: a role for p75NTR and Fas? // Current Opin.
Neurobiol. 2000. V. 10. P. 111-117.
74. Schall J. D. Neural basis of deciding, choosing and acting // Nature Rev.
Neurosci. 2001. V. 2. P. 33-42.
75. Sherrington Ch. Man on his nature. The Gifford lectures 1937-8. Edinburgh:
Penguin Books, 1955.
76. Smolensky P. On the proper treatment of connectionism // Bahavioral
and Brain Sciences. 1988. V. 11. P. 1-23.
77. Strange K. The end of nave reductionism: rise of systems biology
or renaissance of physiology? // Am. J. Physiol. Cell. Physiol. 2005. V. 288.
P. 968-974.
78. Strassmann J. E., Zhu Y., Queller D. C. Altruism and social cheating in the
social amoeba Dictyostelium discoideum // Nature. 2000. V. 408. P. 965-967.
79. Subak-Sharpe H., Brk R. R., Pitts J. D. Metabolic co-operation between
biochemically marked mammalian cells in tissue culture // Journal of Cell Science.
1969. V. 4. P. 353-367.
80. Thompson E., Varela F. J. Radical embodiment: neural dynamics and
consciousness // Trends in Cognit. Sci. 2001. V. 5. P. 418-425.
81. de Waal F. Good natured. The origins of right and wrong in humans and
other animals. Cambridge, London: Harvard Univ. Press, 1996.
82. Webb B. Neural mechanisms for prediction: do insects have forward models?
// Trends in Neuroscience. 2004. V. 5. P. 278-282.
83. Wilson E. O., Consilience. The unity of knowledge, A. A. Knoff, New York,
1998.
84. Woese C.R. A new biology for a new century // Microbiology and Molecular
Biology Reviews. 2004. V. 68. P. 173-186.
85. Yuan J., Yankner B. A. Apoptosis in the nervous system // Nature. 2000.
V. 407. P. 802-809.

58

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89

 ,
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 ,
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91

1. ., ., ., . / . .
.: , 1992.
2. . . .
. 1. .: , 1964.
3. ., . / . . .:
, 2008.
4. . . . . 3. .-.: - , 1950.
5. - . .
. .: - , 1974.
6. - . . . . .: , 1978.
7. - . . . : ,
1995.
8. . . . .: . , 1955.
9. . . . .: -17, 1998.
10. . . . .: -, 2001.
11. . . . .: , 2005.

1. Barbanera F., Erra F., Banchetti R. Non-Adaptive Behaviour of Isotropically

Heated, Inert Populations of Oxytricha bifaria (Ciliophora, Stichotrichia) //
J. Eukaryot. Microbiol. 2002. Vol. 49, . 1. P. 54-62.
2. Bream J., Davis R.M. Rain, wind- and touch-induced expression of calmodulin
and calmodulin-related genes in Arabidopsis // Cell. 1990. Vol. 60. P. 352-364.
3. Hertwig O. Das Nervensystem und die Sinnesorgane der Medusen. Leipzig:
Verlag von F.C.W. Vogel, 1878.
4. Iftode F., Prajer M., Frankel J. Nuclear and Cortical Regulation in Doublets of
Paramecium: II. When and How do Two Cortical Domains Reorganize to One? //
J. Eukaryot. Microbiol. 2001. Vol. 48, 6. P. 690-712.
5. Kuhlenbeck H. Invertebrates and Origin of Vertebrates. Geneva: Karger
Press, 1967.
6. Lipps J. H., Signor W. Origin and early evolution of the Metazoa. N. Y.:
Plenum Press, 1992.
7. McConnel J. V. Comparative physiology: Learning in invertebrates //
Annu. Rev. Physiol. 1966. Vol. 28. P. 107-136.
8. Meglitsch P. A., Schram F. R. Invertebrate zoology. Lond.: Oxford Univ.
Press, 1991.

92

 III.
,

93

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.
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94

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(Na+, K+, Ca2+ Cl-).
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120

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. AMPA -NMDA , Na+/K+-.
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121

. (-75 ) NMDA- Mg2+. , -NMDA

, , Na+/K+/Ca2+-, . NMDA-
. NMDA- ,
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5) (, Zn2+).

. 5.16. NMDA-. NMDA , : 1) ; 2) , , ; 3) , ; 4) - Mg2+ ; 5)

(, Zn2+).
122

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123



. (6 , 3 , 3 , , , ).
(2 -,
2 - 1 -, 22),
(. 5.17). .
, , ,
. : , .

. 3.17. .
(2 -, 2 - 1 -, 22), . -.
-, -.

Cl-. ,
, Cl- , .. Cl-.
124

 , .
Cl-,
.

Cl-. Cl- . ,

.
- , Cl-. ,
,
.

(
) ,
,
, . .
-, (15 , 25 , 35)
(1m2n , 2m3n , m+n=5). . ,
, . .
Cl-,
, Cl-
.
B
B ( , ,
) .
(GABABR1 GABABR2), -, .
B ,
.

125

B - 2+-, , .
2+- - G-.

. 5.18. B GABABR1
GABA BR2. GABA BR1
G-, - +- ().
GABABR2 G-, - () ().

B G, - +-, - (. 5.18). +- , B
.


.
, , .
- , . , - , -
- ( ), ,

126

 Cl HCO3-. .

, .
5.4.4.

() . ,
, . .

( ), .
- .
P2X 7 , .
(Na+ Ca2+), -.
, , .
, P2X1,
(, ), 2+ 3.
P2Y 6 , , , Gq- (
), Gi- Gs-, ,
.
,
.
P2X 2+-
, ,
.
127

 2 (P2T) (P2Z).
.

, : 1, 2 3.
1 ,
Gi- ,
, , .
2-, Gs- .
2+- +-,
.
3 .
5.4.5.
, . ,
-,
,
. , , . ,
.
,
.

, ,
, . 11 . (NK1, NK2, NK3),
G-. , -

128

 , (. 5.19).

. 5.19. .
, ,
,
.
, , ,
.

, . (Met- Ley-).
, -

129

 . ,
( ).
,
, (. 5.19). . ,
-, 2+- +- .

.
-.
, Go- - 2+-. - - Gi-
, , +-.
. - .
.

: -, -,
(), , - ,
.


(). ,
( ),
( ), .
, ,
, , , ,
- , ( ), .
130

 , . , ,
.

1. . . . .: - .- -,
1999.
2. . ., . ., . ., . .
. .: , 2003.
3. . . . .: ,
1997.
4. ., ., . .
. .: , 1991.

1. . .
.: , 1973.
2. . . . .: , 1990.
3. Zigmond M. J., Bloom F. E., Landis S. C., Roberts J. LSquire., L. R.
Fundamental Neuroscience. Academic Press, San Diego, London, 1999.

131

 6.

6.1.
, : ,
,
. ,
.
, , - . . ,
.
, (),

( , ) -60
-100 .
,
.
+, .
, +. , , . . 1902 .
,
+, .
, -70 :
[K+]in

RT
EK+ = -

ln
nF

,
[K+]out

[K+]in [K+]out +, ,
.
132

 , - (. 6.1).
6.1

, ,

. .

6.1. , + . .

+ ,
. ( )
+ (. 6.1). + (dC) (dx).
, , + .
+
(C), (z) (u),

133

 (X).
+:
dC
D

= - uCzX
dx
, ,
+ (. 6.2).
, ,
+ . , , , Na+ Cl-,
.
, Na+
110% +, Cl 15% +. . .
, :

[]in []out , ,
, P . Cl
,

.
,

( )
.

. .
, .

, . 134

, ,
.

. 6.2. +. , , -
Na+ Cl-.

,

.
,
.
.
6.2.

( ,
, ).
.
, .
135

 , .
(),
,
. , , (-) .
, , ,
.
. , ,
, (. . 5 ).
6.2.1.
- . (), ( ), () .

. 6.3. , . .

136

 ,
, (). , 5 ;
(. 6.3). ,
.

, (. 6.4). . , ,
. ,
, .

. 6.4. ,
.

, . Na+ -

137

, , , Na+.
Na+ K+.

. 6.5. ,
(), . ()
. ()
-120 ; , -3
, . ()
.

,
, . ,
(. 6.5). -3 , .
138

 . ,
(IK+ = INa+ , .. ), .
(-)

, .

. 6.6. , , , .
. ( ) ... .

, , .
(), (. 6.6).
.
-60 -40 , , - Na+-,
139

 . , , ,
. . , , , .

(. 6.6), (...) .

. 6.7. , -,
()
().
. , .

140

 , ,
, ,
Na+/+- (. 5.4.1) - ,
. ,
, ,
(. 6.7).
,
. , , , . , (),
(). ( ),
.
6.2.2.
, , ,
. , ,
, . , ( ),
, , .
, ,
().
, , , ( ) . ,
(), ().
, Cl. Cl , .

141

 - ,
.
.


(. 6.8 ). (-74 ) .
(-64 ) . -82
( ). -82
( ) . , -80 .

. 6.8. , , , . .
( ) ... .

142

. 6.9. () ,
, . ()
.

, , (. 6.9). -70
143

, Cl . -70 ,
. , -70 Cl-, , , .

,
, .
,
, ...
(. 6.8 ). , , Cl-. , Cl , , .

. 6.10. ,
, Cl
SO42. , Cl.

144

 ,
Cl
, SO42-.
(H-) Cl-, ()
(. 6.10). Cl SO42 - - , Cl
.
, .
6.3.
, -
. , .

. 6.11. - .
, . .
145

 - ,
.
, , (. 6.11, , ). 1,5 , (. 6.11, ),
( . 6.11, ). (. 6.11, ),
( . 6.11, ) , , .
, Cl--. , Cl-- .
.
,
- 2+- .
. , .
( )
,
.
- , -
.
, . , ,
, , .
6.4. ?
.
, ,
, .
146

 , , .
, , .

. . ,
. , , Na+ +,
Na+ ( +55 ) + ( -90
) . , , ..
, .

. 6.12. . V
, V , .
.

(),
(. 6.12, ). (), (. 6.12, ).
. , +-. + , +
(. 6.12, ).
. +-.
147

 - , + , ,
,
, . .

, .
6.5.
, , ,
, ( 5.3).
,
+- .
(.
6.13, ). , (. 6.13, ).
,
, , . - ,
, -
.
- . -
, +-
, -. .
6.6.
, ,
,
. ( ) ( ).
.
148

. 6.13. . () () () . ()
,
(). - Cl--, . - +- ( +-), . -
( ).
(
).

149

6.6.1.
() -, Na+,
K+ Ca2+. 5 .
, ( ),
( ), .
, : (1) AMPA,
(2) (3) NMDA.
AMPA-
Na+, K+ Ca2+. 0 .
(-75 ) NMDA- Mg2+;
, Na+/K+/Ca2+-,
. NMDA ,
.
6.6.2.
- A-, Cl-. Cl--
75 . Cl-- , ,
, .
A- - Cl-- ,
.
, , ()-, Cl-.
6.6.3.
+- , , , , ,
, , , , . , (
-) +-. +-
,
.
150

 Ca2+-. , ,
Ca2+- .

,
. Ca2+ , Ca2+- .
6.6.4.
( +-)
- +-
( 6.8.10), .
IAHP (+- )
, , , , .

, .. . IAHP (),

.
-
.
I ( +-) -. , IAHP ,
,
65 , IAHP , . I
, ,
, .
6.7.
(
, ) .
( ),
( ). ,
151

 .

; ; , .

, ..
, , - , .
6.7.1.
,
. ,
. , - . -
, , .
, i, r, v = ir.
6.7.1.1.
,
, .

(. 6.14). 6.15
. cmrm ri. rm , ri
.
( ri) , ,
(I = U/R). rm
, -

152

, .
.

. 6.14. () . , . , .
. () .
153

. 6.15. .
: ri , rm , cm .

, (Vx) :
Vx = V0e-x/
V0 . ,
. , , .. 37%
.
6.7.1.2.

rm , ri :
= (rm/ri)
,
, , , , , .
:
rinput = 1/2(rm/ri)1/2
1/2 , . rinput
, rm ri.

154

6.7.1.3.
rm ri, 1 .
,
.
,
.
,
Rm, 1 2 2.
1

2 2. rm Rm/2
,
rm.
Rm=2
Rm
+- Cl- . Rm,
2000 2. Rm, , 1000 2
50000 2 .
Ri 1 1 2.
2, Ri = 2ri.
, ri = Ri/
Ri , 30 . Ri
. Ri 125 ( )
250 ( ).
6.7.1.4.

, , :
rinput = 1/2( RmRi /2
23)1/2
, 3/2
.

155


(Rm Ri)
:
Rm/2Ri)1/2
= (
, . , . ,
( 1 ), ( 50 ) ( 1 )
, , 13, 1,4 0,3 .
, rinput
, , . , ( rinput) ,
. , -
, .
6.7.1.5.
,
. ,
, . 5 .
1 /2.
, ,
, :
Vt = V0(1-e-t/),
t , Vt
t, V0 ,
,
(RmCm).
, 63% (11/)
. .

. ( ) ,

156

 .
, , (RmCm) . ,
, , .
1 20 .

. ,
. , ,
.
, , , , .
.
(Vx = V0e-x/).
(, ) , .
, .
6.7.2.
. 10-710-10 .
,
. (. 6.16), ,
,

. , ,
, .

, (. 6.17).

157

 ,
, ,
. ,
,
.
, ,
.

. 6.16. . .

6.7.3.

, RC- (. 6.18). , .
, . RC-
= RC.
158

. 6.17. .
()
50 .

. 6.18. ()
. , R ,
R R ,
. R R << R.
159

 . : , , , ,
- ,
- .
( ) .
. ,
(. 6.19). , di3/2=const.
, , , .
. ,
, , .
,
(. 6.20). , , ,
.
6.8.
6.8.1.
. , , + ,
. ,
.
. , 1930- .. . . . -

160

 , ,
( 1000 25 2), , , . ,
, .

. 6.19. () ( ), . ()
. ( 1).
()
1, 5 9 . ,
( ).
161

. 6.20. ()
.
. () () .

. 6.21. . . .

, . . , , ,
(. 6.21). ,
Na+ 162

 , (. 6.22).
, ,
Na+. Na+ 10
, (. 6.1), ENa+ 50-60 .

. . . ,
Na+
Na+-:
[Na+]out

RT
ENa+ =

ln
nF

,
[Na+]in

[Na+]in [Na+]out Na+, , .

. 6.22. Na+ [Na+]o . (100%) .

+- . ,
Na+ +, . ,
, . -163

, , , , .
, . .
. -, Na+ + , .
Na+
(), . .
6.8.2.

.

(. 6.23).
. ,
,
. ,
,
.

. 6.23. . .

-
.
164

 , . , , ,
.

( - ) , .
,
-. ,
.
, .

. 6.24. ,
( . . ).
, B , C ( ).
165

6.8.3.
1952 . . , .
. ,
(. 6.24). , , ,
(. 6.24, ). Na+
,
(. 6.24, ).
(. 6.24, ). .

. 6.25.
. ()
( , ),

. () ( Na+-) . () ( +-)
.
166


(. 6.25). , (TTX, Na+-) ,
(. 6.25, ). , (TEA, +-),
, (. 6.25, ). .

. 6.26. () - () ( ).

,
, (. 6.26). 20 .
, . ( +-), .
- . ( 167

 )
- +-.
6.8.4.
, Na+ 0,5 3 , , . Na+- Na+-.
, . .
Na+-.
-
Na+ (. 6.27). ( 30 )
( )
Na+- (.
6.27, , ) , (.
6.27, , ). Na+- (. 6.27, , , ). 6.27 ().
-30 Na+-
( ).
-95 ( 70%) Na+-.
40%
. Na+- .
6.8.5.
- Na+-
. . , Na+-
,
().
(. 6.28, ) . , ,
, ..
. , Na+-, ,
. (. 6.28, , )
(. 6.28, , ).
(. 6.28, , ),
(). ,
+-, +- .
168

. 6.27. () Na+-.
() . () .
(, ) . () Na+- (INa,/INa,), , ) .
Na+- (h, Na+-), 0 , 1 .
169

. 6.28. . ()
Na+
, ().
, (). ( , )
, , (). () Na+- . () Na+-.

Na+-
, . ( )
Na+-.

. , , .
170

6.8.6.
( . )
Na+- .
.
Na+- . (
) Na+, m- (. 6.29, , I). m, Na+ (. 6.29, , II)
h-, ,
Na+ (. 6.29, , III).
h- m, .
m- h- (. 6.29, ).
-50 m-
, 50% -30 (
), h-
0.
Na+-.

. 6.29. () m- h- -
Na+- (). () . . .

6.8.7.
, -
: (1)
Na+-,
; (2) - +-, 171

 . , ,

(. 6.30). ,
. Na+ - +-.
, .
. .
. , , .

. 6.30. .
(),
. , . ,
.

172

. 6.31. () . () , Na+-. (, Vp) Na+- (, Ip).

Na+- (),
.
173

6.8.8.

, , . 1 ,
(. 6.31, ).
, ,
.
(. 6.31, ). , ,
. , Na+-, , , . ,
, m- h-.
Na+-.
Na+- , .
6.8.9. Na+
. Na+-
.
Na+- 0,30,5 , Na+- ,
.

, . . : (H3N+-OH),
(H3N+-NH2) (H3N+-CH3). , - 0.370.38 , H3N+-OH 100
, H3N+-CH3. ,
. 0.08 ,
0.30 . . ,
,
.
174

. 6.32. Na+ . Na+- Na+ . ,

.

(Na+- ) Na+ -
(. 6.32). () ,

. , .
175

 , Na+ - , 104 -1,

Na+- 107108 -1.
6.8.10.
, . 1960- . . .
, , .
( ) - Na+- +-, . , , ,
2+, +-, , , (. 6.33). Aplisia.

. 6.33. , . ( Na+-
Ca++-) ( I+,fast I+,slow).
176

. 6.34. , . ( INa+,fast INa+,slow, ICa++) ( I+,fast I+,slow). Na+-, , .

, ,
Na+-,
1015% (. 6.34). Ca2+-, , 2+- 2+ Cd2+.
- , . ,
- 2+-, (. 6.35).
, Na+/K+ .
.
- , (. 6.36).
177

. 6.35. . (-), 2+- - 2+-. ,

Na+/K+ ().
().

- ( ) . ,
, , ,
. . (., transient),
,
(sustained), , (persistent),
(long lasting).
Na+-
Na+-
. ,
.

178

. 6.36. -
.
-100 -10 () -10 -100 ().

INa,t (transient). . INa,t, INa,p .

.
INa,p (persistent, . 6.34 INa+,slow). , . , . .
2+-
2+-, ,
179

 (). Ca2+- 20 10 .
IL, IN IP.
Ca2+- , ,
Ca2+, , Ca2+- K+-,
.
IL (Long-lasting, ) ( 10 ).
IN (Neither, , ; IT, IL;
IN, IT IL) 20 ,
. IN
.
IP (Purkinje) (. 6.35),
.
Ca2+-,
, Na+/K+ .
2+- IT (transient, ) () . 65 (Na+/K+ 55 )
. IT Ca2+- 50-100 .
Na+/K+ , .
+-
, , . K+-
,
, .
IK (, +-) -40 ,
.
, INa,t-, . IK .
2+- +- (IK,Ca) 2+. 2+- .
180

IC .
, . IC , . IC 10 .
IAHP (AfterHyperPolarization, ) IC . ,
, .
IA ( +-) 60 .
. IA ,
.
IM (- +-) 65 ,
. - ,

. IS , .
Ih, IQ If, , . Ih
+ Na+. Ih
, (, ),
Na+/+- / 2+-.
IK,leak ( ) .
.
6.8.11.
-
Na+- 2+-
- Na+-
, .

181

Na+- , 1832
, (I-IV), (. 6.37). , 300-400 , (S1-S6). Na+- .
, , 5- 6-
. S4, ,
. ,
,
- .

. 6.37. Na+-.
(I-IV), . (S1-S6). , , 5- 6- .

- :
, . 2+- , Na+-.
+-
K+-,
:
Kv- (voltage-gated, -) (), (S1-S6);
Kir (inward-rectifier, )
(), -

182

 M1 M2, S5 S6 - .
Kv-
Kv- (. 6.38). +-
Na+-. , Na+-, ,
, 5- 6- , S4 .

. 6.38. () V-. V- Na+-. ()

. ().

Kir-
, , .

G-, Ca2+. ,
.
+ ,

+ (, Ih). , M1 M2 (. 6.39).
183

. 6.39. IR-. 1 2 IR- S5 S6 . IR- , V-, ().

6.8.12.
: , ( , Na+).
6.8.12.1.
,
,
, . ,
25
Na+ 6
( 15 15,006 ). Na+,
+ ,
. , ,
.

184

 (). 1950- .
. , Na+/+. ,
24Na+,
(. 6.40).

( ) 24Na+ . , 24Na+
.

. 6.40. 24Na +
( ) (). 0,2 24Na +
. , 24Na +
.

Na+/+- (. 6.41). (Na+/K+-) Na+ +,
,
. , Na+
.
, +
.
185

. 6.41. Na+/K+-. .

, 2+ . 2+- 2+ . 2+-
( ) ( ) , . 2+-
2+.
2+ , . 2+ 2+-
2+.

+, PH.
186

6.8.12.2.
. 2+ Na+/Ca2+-, 2+ Na+, .
( ) .
, 2+, . Na+ + 2+
+.
Cl - , Cl () Na+
() HCO3 (). Cl
+/Cl (Cl +).
Na+/+, + () Na+ ().

.
Na+ +,
- ( 3-). ,
(, )
Na+ Cl-.
, , .
. , .
, ,
Cl-.
6.9.
. ,
187

: .
.
. .

. 6.42. . .

6.9.1.
. , ,
188

. , - Na+-, (. 6.42). - + -,
.
, Na+-
.
, , .
.
( 6.7), , , . , (RmCm), .
6.9.2.

. , . , (. 6.43),
. , .
( ). 100
. ,
, , ,
(. 6.44). ( . saltore ) .
.
.
189


,
, .

. 6.43. ( ), .

6.9.3.
, , 200 /.
.
.
,
. , , 190

. 6.44. .
(0 ).
(), . ,
, .

, . , 0,7 , 191

 0,6 0,8. , ,
100 .
6.9.4.
Na+-
, +- . , - Na+-, +-,
Na+- .
, , +-, Na+- .
6.9.5.
, , - Na+-.
5. ,
( ),
.
,
. -,
. , .
6.9.6.
, - Na+- 2+, . , 2+-. (. 6.45).
192

. 6.45. . ()
. () . () 2+- .

, (. 6.45, ).
. 2+- , (. 6.45, ).
, , . -,

193

 . , Na+/+- 2+- ,
- . , ,
.
.

, , + ,
+. (+ ) (+
)
+.

(Na+, 2+, Cl H+), , .

( ),
(Na+/K+- Ca2+),
().
(), .
,
,
.
, Cl--, . (), ,
, ().

. -

194

 , 2+- .
( ) , , , +-. , .
, , +-, ,
.
- 2+- ( 5).
( , ) .
,
. ,
.
, ,
, .
- Na+.
- Na+-, - +- .
Na+- .
, .
.

- 2+-.
- Na+- 2+- , . - +-

195

 , .

( ) .
, .
Na+, +, 2+ + ( )
.
,
Na+ H+.

.
.
.

1. . . . .: - .- -,
1999.
2. . ., . ., . ., . .
. .: , 2003.
3. . . .: . ,
2005
4. . . . .: ,
1997.
5. ., ., . .
. .: , 1991.

1. . . .: , 1973.
2. . , , . .: , 1968.
3. . ., . .
. .: , 1981.
4. . .
.: , 1973.
5. . . .: , 1965.
6. . . . .: , 1990.
196

 7.
, .
,
.
.
, .

, ,
.
7.1.
XIX
. 1836
,
. (1865) , .
(1886) , (1887) ,
. , () . ,
,
.
() (1973). (1875), 1885
(),
(. 7.1).
, 197

 (Golgi impregnation, Golgi staining).

.

. 7.1. (A) ()
(Golgi C., 1885. A monograph on the fine anatomy of the central nervous organs).

, ,
.
.
. ( , neuron
) , I II.
, (. 7.2).
, XIX .

,
1906 . 1906 --. ,
.
--, , , (1988), (. 7.3).
198

. 7.2. (1885. A monograph on the fine anatomy of the

central nervous organs). :
I (1, 2) II (3, 4).
, .

. 7.3. ()
(, 1985) () ,
- (2003).

199

 , ,
, . . : .
( , ,
).
,
.
. :
( . 1887 .). 1906
: . . 1932 ( )
.
. : .
, .
. , , , , , .

, () (,
)
.
XIX
. ( -

200

 ) . , .
XX .

. . . . .
(. . ., 1974, 1990).
1936 , .
XX
.
7.1.
7.1

1843
1850
1904
1906
1914
1921
1924
1926
1933
1934
1935

- .
.
. .
.
. .
.
.
. . .
. , .
. . .
.
. . () ()
.
201

1936
1936
1959
1963

1970

.
. . .
.

, (
).


, .

(S. Palay).
. , . ,
, .
. . . , : .
7.2.
7.2.1.
()
. ,
, . .
. .
, (, 1974, 1990).


202

 , ,
.
,
. :
1) : , ;
2) ( : - );
3) (, );
4) ( );
5) , , ;
6) .
,
, ,

( ). . .
. .
7.2.2.


. (25 ) , ,
, , (gap junctions). 1
. () , . ( ,
connection ). -, . , , 1.
. ,

203

: 1) ; 2)
, , ; 3) - , .

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, 2+ + ( ). ( . ., 2000).

204

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205

 , (. 7.4). , (postsynaptic
density, PST). ,
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206

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211

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, , .
(, ).
, .

. 7.11. :
. , . (1966). , . : Sir John
Carew Eccles Nobel Lecture, December 11, 1963: The Ionic Mechanism of
Postsynaptic Inhibition (: , 1966).

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( mediator , ) ().

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215

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. (, 1974, . 164).
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. : - (-), - (-), - (-), (-), , - (-) ,
, (--, --) (, 1987).

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(, 1987). -, -
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-- -- (. 7.12). () () ,
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, 1990).
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(, H3 , HA)
().
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(, TRK-R).
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(-, IP3, NO Ca2+).
217

. 7.13. .

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.

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218

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(, 2006).
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(, 2003). , , , ,
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- .

219


, (. 7.14).

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. 7.14.
(Rollenhagen and
Lbke. 2006).
a. (MNTB), (,
, ).
b.
CA3 .
c-1. .
d. - :
.

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-
(. 7.15). (110 , 36
, ),
, ,
(
)
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 , ( )
.

. 7.15. .
, , Helix pomatia/Helix lucorum).
( : LPag,
RPag) (Vg) . (, ) -
(LPa2), (Aortal. n.). ( ).
. 100 .
221

 , , (Aplysia),

(. 7.16). , , , 2000
. . .

. 7.16. , (Aplysia) (gill).

(. . ).

7.9.2.
- - ( ). 222

. 7.17. .

, (, 1972; , 1980; Kandel, 2000).

-
. 20-

, , , - (, Katz B.,
1972), (, 1995),
, ( ). . ,

(patch clamp), .
,
,
: (the enddulbs of Held)
(the calyxes of Held). , (. 7.17). ,
19- (Held, 1893).
,
, :
.
223

 -- (, 1985).
(. 7.18).

. 7.18. .
(MNTB). MNTB

, MNTB
.

. .

.
, , , .
.
. ( ), . , .
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224

 , . ,
.
, ,
.

- .

1. . ., . ., . ., . .
.: . , 2003, 672 .
2. . . : . .:
, 2003, 287 .

1. ., ., ., ., ., .
p . .: p. 1994, . 2, . 12, . 338-383; . 3,
. 16: . 67, 78, 81, 94-100, . 19: . 287-375.
2. . .
// . . . ., 1997. . 47, 2, . 261-286 (
, , 2000, . 217-230).
3. . . (.) . .: , 1974. 158 .
4. . . / . .
. . . .: - , 1975, 160 .
5. . ., . . , , (
). , 1983, . 2. 3. . 327.
6. ., ., . , . .:
, 1988. 64 .
7. . ., . . :
. .: , 1992. 152 .
7. . . (2000) :
// . . . . 2000. . 50. 50. . 743-774.

225

8. . . .
.: - , 2006, 332 .
9. . . . 1997;
. 2002; .
2003.
10. . . - . .: ,
1979. 165 .
11. . ., . . . .: , 1978. 64 .
12. ., . . .: . 1979.
13. . . .: , 1980. 598 . Kandel E. R.
(2000) The Nobel Prize in Physsiology or Medicine 2000. (Kandel_Nobel
lecture_2000.ppt).
14. . . .: , 1968. . 1-8.
15. .
. .: , 1983. . .:
, 1987.
16. . . .: , 1975. . 17-64.
17. . : .: , 1995. 380 .
18. . . . .: , 1974. 183 .; . .
: . ., 1973. 46 .
19. . . (1990) : . . . . ., 1990, . 26, 5, . 733-749.
20. . . . // . . . . 1995, 2, 3, 4, 5, 1996 5, 1997 4.
21. ., . . .:
, 1976. 198 .
22. . ., . . . .: , 1983.
23. . . . .: ,
1981. 181 .
24. . . . : . ,
1986. 264 .
25. . ., . .
. .: , . ., 1987. 173 .
26. . : 2- . .: , 1987. 454 .
27. . . (.) . .: , 1996. 4- .
28. ., . . .: , 1981. 135 .
29. Basic Neurochemistry. 6th Edition. Eds.: G. J.Siegel, B. W. Agranoff,
R. W. Alberts, S. K. Fisher, M. D. Uhler. Part two: Intercellular signaling.
30. Cell and Tissue Research/ September. Biomedical and Life Sciences and
Medicine. Springler Berlin / Heidelberg 23, 2006, v.326, pp. 201-669.
31. Klaus Unsicker (from the Editor:) The synapse recent advances. P. 201;
32. Frotscher M., Gundelfinger E., Jonas P., Neher E., Seeburg P. (Editorial)
The most important advances in synapse research from my point of view and
what remains to be done. P. 203-204.
226

33. Jan L. Gibbins and Judy L. Morris Structure of peripheral synapses:

autonomic ganglia. Review, pp. 205-220.
34. Astrid Rollenhagen and Joachim H.R. Lbke The morphology of excitatory
central synapses: from structure to function. (Review), pp. 221-237.
35. Scheggenburger, Ralf and Forsythe, Jan D. The Calyx of Held. (Review),
pp. 311-337.
36. . . .
.: , 1980, 196 . ( (1898-1974). .: , 1987. 128 .
37. . . . .: , 1975. 333 .
38. . . ( ) / .
.. .. . . . .: , 1985. 272 .
39. . (Katz B.) The release of the neural transmitter substances. Liverpool
University Press, Liverpool, 1969.
40. . . / . . . . .: -
, 1951. 616 .
41. . . (1926, 1927, 1935, 1952) . : , 1952. 287 .
42. . . : .
. . (. . , . 5-24). .: (
), 2007, 127 .
43. . . . .:
. . . , 1958.
44. . . ( ). - , 1969. 176 .
45. . (George Somjen, 1972.
Sensory Coding in the mammalian nervous system). .: , 1975. 415 .
46. . . .: , 1957. 187 .
47. . . .: , 1971. 222 .
48. ( ). .
. . .: , ., 1981. 144 .
49. . . .: , 1965, 125 . (Hodgkin A.L. The
Conduction of the Nervous Impulse/Liverpool University Press, 1964).
50. . . .: ,
1959. 298 .
51. . . .: , 1966. 395 . (
. .. . . . . ) / John
Carew Eccles (1964) The physiology of synapses. Spriner-Verlad, Berlin, Gettingen,
Heldelberg.
52. Hebb D. O. (McGill Univ.) The Organization of Behavior/ A Neuropsychological
Theory. USA, N.Y., John Wiley & Sons, Ink.; London, Charman & Hall, Limited, 1949,
P. 335.
53. Sir John Eccles Nobel Lecture, December 11, 1963: The Ionic Mechanism
of Postsynaptic Inhibition.
227

 8.
. , . ( )
.
. , .
-, . .
8.1.

. ( , , , ), ( ) () .

( , , ) ( ). , . ()
,
.
, (), , (), .
228

 . , ,
.
(
) .

,
.
8.2.
(Aplysia)
(Helix lucorum). ,
( 1 ), .
, .
, . .
, , . , , .
8.2.1.

, , , . ,
(. 8.1).
, (. 8.1, ).
229


(. 8.2, ), , , , .

. 8.1. ,
. ,
.

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, (. 8.1, ).

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.
230

 ,
(. 8.2, ) .

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()
(). ICa++ 2+-,
. .

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,
.
8.2.2.


, 54- , Gs- (. 8.3). - , - s+-. +- (. 8.3,
1), 2+- (. 8.2), N-.
, - 2+- L-, ,
2+ (. 8.3,
3).
2+
2+/- II ( II) ( ). , 231

 , .
, (. 8.3,
2). 2+
( ). 2+
.

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. .

52 Gq-,
. Gq 2
3 . (), 2+- L-, (. 8.3,
2 3).
232

8.2.3.


,
. , .

. 8.4. . 1 2 ,
(. . 8.3).
.
233

 8.4. , , .
- , (. 8.4, 3)
ApCREB1 (Aplysia CRE-binding protein),
( . 8.4 CREB). , ApCREB2 ( )
, -
.
, . , - (
), ApCREB2 .
, , . . (, . 8.4)
, . ,
(. 8.4, 1 2). ,
CREB1, . .
8.2.4.


. (Kandel, 2000). (Dullock, 2000).

. -

234

 (Helix lucorum).
, , , .
, ,
. ,
, , () (
, , uEPSP) ,
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facilitate , , , ), ( ), . ( )
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236

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237

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238

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. 1
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2+ ,
1 . 2+
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( AMPA),
, .
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) , Na+/K+-
AMPA .
, NMDA- -
Mg2+-, (. 8.8, ,
, ). (
, . 8.6)

239

-30 , 2+- Na+/K+/Ca2+-,

(. 8.8, , Mg2+, ). NMDA ,
. NMDA- 2+- 2+
. , ,
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,
2+ (. 8.8, ). NMDA- APV (. 8.8, ).
NMDA- , NMDA- APV .
2+- , , 2+. ,
2+
. 2+ . II - . , II .
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, ,
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.
, - 240

 .
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. 8.8. NMDA-. () . () . () . ()
. .
241

. 8.9. (, 2000).
1. - , .
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3.
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4. , ,
.
242

 .
, ,
1, NMDA ,
-. -.
,
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) . , ,
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, 2+ . II : 1)
- ,
, ; 2) ()
- , .

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AMPA- . ( 3 .) . 2+
2+- ,
(CREB),
, ,
.

243

8.4.
, ,
, 2+ . ,
2+ ,
. 2+ 2+- . ,
.
-
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.

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244

. , 2+-,
.
2+ ,
2+/- II. (1) - ,
, (2) .
, 2+
,
CREB, ,
, .


, . , .

1. . ., . ., . ., . .
. .: . , 2003. 672 .
2. . . : . .:
, 2003. 287 .

1. . .
.. . . ., 1997. . 47, 2, . 261-286 (
, , ., 2000. . 217-230).
2. . ., . . :
. .: , 1992. 152 .
3. .. . .: ,
1979. 165 .
4. . . .: , 1980. 598 .
5. Kandel E. R. The Nobel Prize in Physiology or Medicine 2000. (Kandel_Nobel
lecture_2000.ppt).
6. . : . .: , 1995. 380 .
7. . . . .: ,
1981. 181 .
245

8. . . .: , 1968.
9. Kandel E. R., Schwartz J. H., Jessell Th. M. Essentials of neural science and
behavior Appleton & Lange, Stamford, 1995.
10. The synaptic organization of the brain. Ed. by M. Shepperd, 4th ed., Oxford
University Press, NY, Oxford, 1998.
11. Zigmond M. J., Bloom F. E., Landis S. C., Roberts J. LSquire., L. R.
Fundamental Neuroscience. Academic Press, San Diego, London, 1999.

246

 9.

. ,
. , ,
. , ,
.
9.1.

,
, . .

, . , ,
. , ,
, ,
30 , ,
.
. , , , , 50%
50- . . . . , ., , 5070
1516 , .

. 247

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90% ,
35 .

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9.2.

.
, , : -
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.
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:
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,
, - ,
.
1893 ., ,
. (E.Tanzi, 1893) :
248

, - ; , ; , ,

, , ,

, .
C. -- (Ramon y Cajal, 1893) ,
,
, 20 . , . .
(R. Semon), 1904 . Die Mneme,
c .
, ,
.
(K. Lashley, 1950). , 30 ,
:
, , . , . 20 ,
,
.
.

249

9.3.

, , , . (Ebbinghaus, 1885). , .
, , . , , ?
, , , WUX, CAZ, JEK, ZUP, RIF ..
2300 , , , 7 36 ,
, 50
.
, , ,
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, .
. , .
,
. . , 2060
, .
, 2030%
, .
, .

. (James, 1890).
250

 , ,
.
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9.4.

. . .
1890- ,
.
.
. 24
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(Mueller and
Pilzecker, 1900).
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1949) , , 15 ,
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253

 ,
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. 9.1.
. (, ,
) , c-fos c-jun.
. Fos, Jun
, - . -
- ( )
( Curren & Morgan, 1991).

c-fos.
, . ,
c-fos
. ,
, c-fos,
-, . , , AP-1. c-fos,
fos-B, c-jun, jun-B, jun-D, fra-1, fra-2
. ,
254

 , , ets-1, ets-2, Myc, Myb, rox-20, zif/286,

NGFI-B, mKr2, Arg 3.1. , ,
, .
,
, ,
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(. 9.1).
1980- , ,
, ,
c-fos. .
9.1.
c-fos
. , cfos ,
. c-fos .
, . () .

, . , , zif268,
Egr-1, NGFI-A Krox-24. .
.
, , , .

255

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, , , .
, . , , : 1) , 256

 (BDNF, Narp); 2) (RheB, RGS-2, Homer 1a); 3) (Arc, TPA) 4) (COX-2).

, , 500,
.
9.8.

, , . ,
, , :
, -. 1969
.
(Glassman, 1969):
, : -1 -2... -1
, . ,
,
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257

 3-5 . , , c-fos c-jun,

.
?
fos/jun, -1
. -1
,
. , -1 , , , , S-100,
, N-CAM.
N-CAM (neural cell adhesion
molecules), ,
.
, , , .

. N-CAM ,
. N-CAM
, .
,
.
. 6 8 4 6
, - .
, N-CAM -1 ,
fos/jun,
, , . ,

.

258

9.9.

, - .
, . , ,
, ,
.
, . -

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.

. 9.2.
(, 1996).

259

 .
9.10.

, ,
,
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,
. c-fos : , . 6 ,
, , c-fos . ,

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, ,
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, ,
, -
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260

 , .
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-
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 . ,

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262

 . ( ) ( )
.
,

. , , .

.
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.

.
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, . .
,
, .
, , . ,
. . ,
,
.
,
263


.
,
, , ,
c-fos, c-jun, zif/286, NGFI-B . ,
, ,
.
, ,
,
, .
,
, , . ,
. , , ,
(BDNF, Narp), (RheB, RGS-2, Homer 1a), (Arc, TPA)
(COX-2).
,
, ,
. , , c-fos c-jun, . ,
,
(
). , S-100,
, , N-CAM .
,

.
, , 500 .
, , ,
. , , -

264

 .
.
,
, .
,
.

.

( )

.
-
.
, . ,

.


: -
,
.
,
, -


, . -
, . ,
( ) ( ) .

265


, .
,
, ,
,
.

1. . .
// . . . . 1997, 47:
262-286 .
2. . . - // , , 1996, 23-65 .
3. . . . . . -, 1911.

1. . . .: , 1950.
2. . -. . . . . -. 1935.
3. . . . .: -, 1996.
4. . ., . ., . ., . .
c-fos .
. 1989, 25: 1119-1121.
5. Pfenning A. R., Schwartz R. and Barth A. L. A comparative genomics approach
to identifying the plasticity transcriptome. BMC Neuroscience. 2007. 8: 1-18.
6. Davis H. P., Squire L. R. Protein synthesis and memory: a review. Psychol.
Bull.1984, 96: 518-559.
7. Dingman,W.,Sporn,M.B. The incorporation of 8-azaguanine into rat brain
RNA and its effect on maze- learning by the rat: an inquiry into the biochemical
bases of memory. J. Psychiat. Res. 1961. 1, 1-14.
8. Dityatev A, Schachner M. Extracellular matrix molecules and synaptic
plasticity. Nat Rev Neurosci. 2003,4: 456-68.
9. Duncan, C.P. The retroactive effect of electroshock on learning. J. Comp.
Physiol. Psychol., 1949, 42: 32-44.
10. Hyden H. Quantitative assay of compounds in isolated fresh nerve and
glial cells from control and stimulated animals. Nature, 1959, 184: 433-435.
11. Hyden H. and Lange P. Brain cell protein synthesis specifically related to
learning. Proc. Natl. Acad. Sci. USA. 1970. 65: 898-904.

266

12. Flavell, S. W. and Greenberg M. E. Signaling Mechanisms Linking Neuronal

Activity to Gene Expression and Plasticity of the Nervous System. Annu. Rev.
Neurosci. 2008. 31: 56390.
13. Flexner, J. B., Flexner, L. B.,Stellar, E. Memory in mice as affected by
intracerebral puromycin. Science, 1963, 141: 57-59.
14. Frankland P. W. and Bontempi B. The organization of recent and remote
memories. Nature Rev. Neurosci. 2005, 6: 119-130.
15. Glassman E. The biochemistry of learning: An evaluation of the role of
RNA and protein. Annual Rev. of Biochem. 1969, 38: 605-646.
16. James, W. The Principles of Psychology. 1890, 2 vols., Holt, New York.
17. Lashley K. S. In search of the engram. In: Physiological Mechanisms in
Animal Behaviour Cambridge University Press., Cambridge. 1950. 454-482.
18. McClung C. A. and Nestler E. J., Neuroplasticity Mediated by Altered
Gene Expression. Neuropsychopharmacol., 2008, 33: 317.
19. McGaugh J. L. Memorya Century of Consolidation. Science. 2000, 287:
248-251.
20. Muller G. and Pilzecker A. Experimentelle beitrage zur lehre vom gedachtni.
Z. Psychol. Physiol. Sinnesorg. Erganzungsband. 1900, 1: 1-18.
21. Nedivi E., Hevroni D., Naot D., et al. Numerous candidate plasticity-related
genes revealed by differential cDNA cloning. Nature. 1993, 363: 718-721.
22. Ramon y Cajal S. Archives of Anatomy and Physiology. 1893, 19: 428-465.
23. Rose S. P. R. Cell-adhesion molecules, glucocorticoids and long-termmemory formation. Trends in Neurosci. 1995, 18: 502-506.
24. Semon R. Die Mneme. 1904/1921. London, George Allen & Unwin.
25. Tanzi E. I fatti e le induzioni nell odierna istologia del sistema nervosa.
Riv. Sper. Freiat. Med. Leg. Alien. Ment. 1893, 19: 419-507.
26. Tischmeyer W., Kaczmarek L., Strauss R., et al. Accumulation of c-fos
mRNA in rat hippocampus after acquisition of a brightness discrimination. Behav.
Neural Biol. 1990, 54: 165-171.
27. Tiunova A. A, Anokhin K. V. and Rose S. P. R. Two critical periods of
protein and glycoprotein synthesis in memory consolidation for visual categorization
learning in chicks. Learning and Memory. 1998, 4: 401-410.
28. Welzl H and Stork O. Cell Adhesion Molecules: Key Players in Memory
Consolidation? News Physiol Sci , 2003, 18: 147-150.
29. Wang H., Hu Y. and Tsien J. Z. Molecular and systems mechanisms of
memory consolidation and storage. Progr. in Neurobiol. 2006, 79: 123135.

267

 10.
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., 2003). , , , , , , .
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- (Arakelov et al., 1991, Marakujeva et al, 1992) (. 10.1).

269

. 10.1. .
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.
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, 1990). , , (. 10.2).
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270

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271


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.
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274


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275

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,

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.

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.
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,

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,
.

.
, .
276

 .
, ,
. .
,
, .

,
, .

1. . ., . ., . ., . .
. .: . 2003. 672 .
2. . ., . .
:
// . . . . 1990. . 40. 6. . 1186-1189.
3. . ., . ., . . - // . . . .
1992. . 42. 6. .1170-1179.
4. ., ., , ., . ,
. : . 2004.
5. ., . , , :
. .: . 2000.
6. . ., . . //
. .
. . . . (.). .: . 1985. . 64-84.

1. Arakelov G. G., Marakujeva I. V., Palikhova T. A. Structural and functional

analysis of monosynaptic connections between identified neurones of Helix lucorum
// In: Simpler nervous systems. D.A. Sakharov, W. Winlow (Eds.). Manchester
University Press. Studies in Neuroscience. 1991. 13. Ch.16. PP. 258-269.
277

2. Atwood H. L. and Wojtowicz, J. M. Silent synapses in neural plasticity:

current evidence // Learning and Memory. 1999. Vol. 6. 6. PP. 542-571.
3. Bailey C. H. and Chen M. The anatomy of ling-term sensitization in Aplysia:
morphological insights into learning and memory // In: Memory: organisation
and locus of change. L. R. Squire, N.M. Weinberger, G. Lynch, J. McGanyh (Eds.),
New York Oxford, Oxford University Press, 1991. PP. 273-300.
4. Eccles J. C. Evolution of complexity of the brain with emergence of
consciouness // In: Rethinking neural networks: quantum fields and biological
data. K.H. Pribram (Ed.). Hillsdale, Lawrence Erlbaum Associates, Publishers. 1993.
PP. 3-28.
5. Gall Ch. M., Lauterborn J. C. Activity-dependent neuronal gene expression:
a potential memory mechanism? // In: Memory: organisation and locus of change.
L. R. Squire, N. M. Weinberger, G. Lynch, J. McGanyh (Eds.), New York
Oxford, Oxford University Press. 1991. PP. 301-329.
6. Hagan S., Jibu M.,Yasue K. Consciousness and anasthesia: an hypothesis
involving biophoton emission in the microtubular cytosceleton of the brain // In:
Origins: brain and self-organization. K. H. Pribram (Ed.). Hillsdale, Lawrence Erlbaum
Associates, Publishers. 1994. PP.153-171.
7. Hameroff S., Dayhoff J.E., Lahoz-Bettra R., Rasmussen S., Insina E. M..,
and Coruga D. Nanoneurology and cytoskeleton: quantum signaling and protein
conformational dynamics as cognitive substrate // In: Rethinking neural networks:
quantum fields and biological data. K. H. Pribram (Ed.), Hillsdale, Lawrence Erlbaum
Associates, Publishers. 1993. PP. 319-365.
8. Marakujeva I. V., Palikhova T. A., Trepakov V. V., Arakelov G. G. Identified
synapses in the snails brain // Neuroinformatics and Neurocomputers. RNNS/
IEEE, Rostov-on-Don. 1992. V.1. PP. 497-508.
9. Prigogine I. Mind and matter: Beyond the Cartesian dualism // In: Origins:
brain and self-organization. K. H. Pribram (Ed.), Hillsdale, Lawrence Erlbaum
Associates, Publishers. 1994. PP. 3-15.
10. Sokolov E. N. Local plasticity in neuronal learning // In: Memory:
organization and locus of change. L. R. Squire, N. M. Weinberger, G. Lynch,
J. McGanyh (Eds.), New York Oxford, Oxford University Press. 1991.
PP. 364-391.

278

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280

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281

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286

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287


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.

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288

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289

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290

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1999.
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2005
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1997.

1. . . .: , 1973.
2. . , , . .: , 1968.
3. Zigmond M. J., Bloom F. E., Landis S. C., Roberts J. LSquire., L. R.
Fundamental Neuroscience. Academic Press, San Diego, London, 1999.

291

 12. :

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292

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293

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294

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, 297

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1990).
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1975; Nelson, 1977; Ahnelt e.a., 1990).
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299

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1998).

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1998).
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301

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1965).

302

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303

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1979).
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, (=500 ) ( , , I Meta I) II (Meta II). II
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304

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GRK). (GRK1) (GRK7) , . , (,
) (GRK1) ,
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, (GRK7 GRK1) (Weisse.a.,
2001).
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2006) -, , , . , , ,
.

305

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306

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GDP) - ( GTP) , , G-.
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. 12.7. . , ( ) ( ) ( Baehr, 1996).

307

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, , -, . -
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- , , - (GTPase activaiting protein GAP).
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RGS) (He e.a., 1998),
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308

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Calvert e.a., 2006).

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Lobanova e.a., 2007).
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, .

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1982). - , . - -
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(Catty .., 1992). (PDE*)
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) 2000 / (Leskov .., 2000).
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100:10:1.

309

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311

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312

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1984; Morgans, 2001).

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313


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314

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315

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2000).
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316

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, 317

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Studholme, 1997).
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-) . , - .
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.

318

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2+
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319

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, .
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, , .
,
.
.
, . .
320


(), .
, :
, .
, , -
, ,
.
, G-. G- .
-
,
.
, , Na/Ca,K ,
- .
, , - L-.
- ,
( ),
, , .

1. . ., . ., . ., . .
. : . 2003. . 19. . 414-442.
2. . . .: . ,
2005.
321


1. Eliasof S., Werblin F. (1993) Characterization of the glutamate transporter
in retinal cones of the tiger salamander. J. Neurosci. 13, p. 402-411.
2. Fain G. L., Matthews H. R., Cornwall M. C., Koutalos Y. (2001) Adaptation
in Vertebrate Photoreceptors. Physiological Reviews. 81, . 117-151.
3. Fu Y., Yau K. W. Phototransduction in mouse rods and cones (2007) Pflugers
Arch. Eur. J. Physiol. 454, . 805819.
4. Gaal L., Roska B., Picaud S. A., Wu S. M., Marc R., Werblin F. S. (1998)
Postsynaptic response kinetics are controlled by a glutamate transporter at cone
photoreceptors. J. Neurophysiol. 79, p. 190196.
5. Hosoi N., Arai I., Tachibana M. (2005) Group III metabotropic glutamate
receptors and exocytosed protons inhibit L-type calcium currents in cones but not
in rods. J. Neurosci. 25, p. 4062-4072.
6. Hurley J.B. (1995). Recoverin and Ca2+ in vertebrate phototransduction.
Behavioral and Brain Sciences 18, . 425-428.
7. Kaneko A., Tachibana M. (1986) Effects of gamma-aminobutyric acid on
isolated cone photoreceptors of the turtle retina. J. Physiol. 373, p. 443461.
8. Kawai F., Horiguchi M., Ichinose H., Ohkuma M., Isobe R., Miyachi1 E.
(2005) Suppression by an h current of spontaneous Na+action potentials in human
cone and rod photoreceptors. Invest. Ophthalmol.&Visual Sci. 46, p. 390-397.
9. Koulen P., BrandsttterJ.H. (2002) Pre- and postsynaptic sites of action of
mGluR8a in the mammalian retina. Invest. Ophthalmol. Visual Sci. 43, p. 1933-1940.
10. Leskov I.B., Klenchin V.A., Handy J.W., Whitlock G.G., Govardovskii V.I.,
Bownds M.D., Lamb T.D., Pugh E.N. Arshavsky V.Y. (2000) The gain of rod
phototransduction: reconciliation of biochemical and electrophysiological
measurements. Neuron 27, . 525537.
11. Mendez A., Burns M.E., Sokal I., Dizhoor A.M., Baehr W., Palczewski K., Baylor D.A.,
Chen J. (2001) Role of guanylate cyclase-activating proteins (GCAPs) in setting the
flash sensitivity of rod photoreceptors. Proc Natl Acad Sci USA 98, . 99489953.
12. Menon S., Han M., Sakmar T.P. (2001) Rhodopsin: structural basis of
molecular physiology. Physiol. Rev. 81, p. 1659-1688.
13. Nelson R. (1977) Cat cones have rod input: a comparison of the response
properties of cones and horizontal cell bodies in the retina of the cat. J. Comp.
Neurol. 172, 109-136.
14. Picaud S., Pattnaik B., Hicks D., Forster V., Fontaine V., Sahel J., Dreyfus
H. (1998) GABAA and GABAC receptors in adult porcine cones: evidence from a
photoreceptor-glia co-culture model. J. Physiol. (Lond) 513, p. 33-42.
15. Schwartz E. A. Transport-Mediated Synapses in the Retina. (2002) Physiol.
Rev. 82, . 875891.
16. Sjostrand F.S. (1976) The outer plexiform layer of the rabbit retina, an
important data processing center. Vis. Res., 16, p. 1-14.
17. Trifonov Y.A. (1968) Study of synaptic transmission between the
photoreceptor and the horizontal cell using electrical stimulation of the retina.
Biofizika. 13, . 809-817.

322

18. Wassle H., Boycott B.B. (1991) Functional architecture of the mammalian
retina. Physiol. Rev. 71, p. 447-480.
19. Xu J.W., Slaughter M.M. (2005) Large-conductance calcium-activated
potassium channels facilitate tansmitter release in salamander rod synapse J.
Neurosci. 25, p. 7660-7668.
20. Yazulla S., Studholme K.M.(1997) Light adaptation affects synaptic vesicle
density but not the distribution of GABAA receptors in goldfish photoreceptor
terminals. Microsc. Res. Tech. 36, p. 43-56.

323

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2006, Armus et al., 2006, Geisler et al., 2005, Moortgat et al., 2000,
, 2005).
326

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327

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328

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329

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331

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332

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333

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334

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et al., 1999, Christensen et al., 1998). GTP
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335

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336

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1967; Frazier, 1968, Kandel et al., 1967).
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). 30
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1968) , , .


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13.5).
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337


. ,
, (,, 1979, Baumgarten et
al., 1971). Na+ ,
, NH4+.
Na+ Ca++.

. 13.5. . ,
. ,
. ( Alving B. O. Spontaneous activity in isolated somata of
Aplysia pacemaker neurons J Gen Physiol. 1968 Jan; 51(1):29-45).

, ,
(Alving, 1968).

. Waziri et al. (1965) , ,
, Na+
- K+ .

338

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. , Chaplain (1979) ,

.
fructose-6-phosphate
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. , .
, .
(2000)
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Rpa1
-6567 .
. V7, , V7 . , , ,

339

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,
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R15 . Lewis (1988) .

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1 mM, .
, - , , .
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, . (Prosser et al., 1989).
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340

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525 12 .

. 13.6. Helix lucorum: ,

0,75 5 ;
()
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341

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. - , . : 10 , 1 .
342

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343

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1985). ,
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344

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Gartside, 1968).
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200 ). : 20 , 1 .

345

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? (1979),
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,
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1975, Gmez-Beldarrain et al., 1998, Niktarash, 2003, Parnas I., Sturmwasser,
1975).
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.
. .
: 10 , 1 . (: . . // , 1975, . . 87-108).

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347

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. () . (, 1975, , 2007). ,
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348

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(): . 1 1 .;

1 30 . : 10 , . . 14.
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349

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.
1;
2; 3.
: 10 , 1 .

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, . : 10 , 1 .

.
,
1, 1 Helix lucorum
(1987) .
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,
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350

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351

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., 2004.).

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352

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353

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354

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355

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357

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359

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Leznik and Llins, 2005, Meijer J. H., Rietveld, 1989, Mieda, 2006, Rabbah
P, Nadim, 2007). , ,
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361



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, ). ,
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II
II (MII PMmouse). , (F), , . ( ).
(ER) () () (MI Pasc MII Pasc) (MII Pmouse).
, Ins(1,4,5)P3 ( ). (
R. Dumollard, J. Carroll, G. Dupont and Christian Sardet. Calcium wave pacemakers
in eggs Journal of Cell Science 115, 3557-3564 2002).
363

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. , , ,
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(McClung, 2006). -

364


. (-,
, ,
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, ,
. (LakinThomas and Brody, 2004), , (Welsh et al, 1995),
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365

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.

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24 . ,
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, (Iwasaki
and Kondo, 2004). . ,
( 1
24 )
. Johnson CH, Golden SS, Ishiura M, Kondo T. (1996)

, .
,
.
: KaiA, KaiB KaiC.
. -

366

 , . , . KaiA, KaiB, KaiC

Synechococcus elongatus PCC 7942. KaiC
(kaiBC) , KaiA kaiBC. KaiC KaiA ,
, , ,
kaiBC. , , , , Arabidopsis (Young and Kay,
2001), Kai
. , ,
, . ,
(Young, Kay, 2001).
: 24-
. ,
, , 24-
, (Roenneberg et al.,
2008).
KaiA, KaiB, KaiC
. , KaiC, in vitro
, .
, (. 13.21).
KaiA KaiC, KaiC. , , (Kageyama et
al., 2006).

367

. 13.21. KaiC
Kai. .
KaiC, ,
in vitro. - KaiC
KaiA (2/3 1/4 . . D KaiC KaiB
(2/3 1/3 . ( Molecular Cell 23, 161171,
July 21, 2006 yanobacterial Circadian Pacemaker: Kai Protein Complex Dynamics
in the KaiC Phosphorylation Cycle In Vitro Hakuto Kageyama, 1 Taeko Nishiwaki,
1 Masato Nakajima,1 Hideo Iwasaki, 1, 2 Tokitaka Oyama, 1 and Takao Kondo1).

368

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.
fisZ , ,
, .
. ,
,
(Mori et
al., 2001).
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. . , .
, .
, .
,
. Mixed-strain
, , (Johnson,
2004).

369

 Synechococcus elongatus PCC 7942

(kaiA, kaiB, and kaiC) . KaiC

, . KaiC in
vitro: KaiA, KaiB KaiC; , , ,
, Kai.
(Kondo, 2007).
in vitro KaiA, KaiB
KaiC, with ATP. iC
10 . , ,
, , KaiC ( ) KaiC. , KaiC
KaiC hexamers
. , , resilience
,
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,
. , , . ,
, , , . ,
,
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370

 . : , .
1932 . ,
eclosion .
- .
. 1948 , ,
.

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,
Synechococcus
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Simons MJ, Merrow M., 2008). , , , , . ,
,
. , , : , , common themes.
, -, (Golden, 2003).
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, (1975) . , R15 -

371

 . in vivo
. R15
,
.
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R15. ,
. R15 - -
. R15 , , . ,
,
R15 in vivo, R15
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1932 (Bnning, 1932). ,
() :
. Drosophila
melanogaster (Konopka and Benzer, 1971).
, ,
, Xenopus, Arabidopsis Synechococcus (Merrow et al., 2005,
McClung, 2006).
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, .
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,
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372

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. , , ,
, . (Barnard, Nolan,
2008).

373

. 13.22. .
,
- , , (). ,
. -,

24 .
, . ( The circadian cycle:
daily rhythms from behaviour to genes First in the Cycles Review Series Martha
Merrow1+, 2, Kamiel Spoelstra1& Till Roenneberg2 EMBO reports VOL 6. NO 10.
2005. P.p. 930-934).

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24 .

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374

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).
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, , 46 (Balsalobre et al, 2000). ,

, .

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. Ba Bb -52 -59 mV
Ih 0 45 pA ( Zhi-Gen Jiang, YuQin Yang,
Zhao-Ping Liut and Charles N. Allen Membrane properties and synaptic inputs of
suprachiasmatic nucleus neurons in rat brain slices Journal of Physiology 1997,
499.1, pp. 141-159).
375

 . .
.

, ,
3-4 ,
, . - . (0,87+- 0,12 ) ,
(0,24+-0,12 ).
, .
, (Kononenko et al, 2008).
Klisch et al. (2006) 12- , - .
- . .
. ,
. , - , (10pM 100 nM) 30 .
. - ,
. , .
,
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, (Klisch et al., 2006).

.

376

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2,5 .
,
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.
, , . , ,
, . , - ,

. Nahm et al. (2005)

- .
mRNA - .
L-, R N-.
,
. , , rPer2 rBmall. viability. ,
-
(Nahm et al., 2005).

, Period1 (Per1). , +
.
Pitts et al. (2006)

377

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. - Per1 GFP
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neurotransmitter. J Neurophysiol. 1997 Jun; 77 (6): 3075-84.).
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Naohiro Koshiya, 1, 2, Robert J. Butera, JR., 3 AND Jeffrey C. Smith1 Persistent
Sodium Current, Membrane Properties and Bursting Behavior of Pre-Botzinger
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jn.00081.2002).

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and Edward G. Lakatta, MD Normal heart rhythm is initiated and regulated by
an intracellular Calcium clock within pacemaker cells Heart Lung Circ. 2007
October; 16(5): 335348).
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418

1. / . . . , . . . , , 1975.
2. . . : // .:
. 2003.
3. . . // . ., 1987.

1. . ., . ., . . // . 2004. . 49., 24.

2. . ., . ., . ., . .

Helix pomatia // , 1986, . 31, 5, 921.
3. . . //
. . ., .14, , 2001, 1, 64-80.
4. . . // , 1984, . 36, 5, 982-985.
5. . ., . . .
.., 1981, . 2, 3, 95-99.
6. . ., . . . // , 1979, . 29, 5, 1093-1095,
7. . . // . 2006. . 3, 2. . 62-72, 24, 31.
8. . . - // . , 2008. , 1 , .248-249.
9. . ., . . Spirostomum ambiguum // . . ., 1976, 12
(6), 539-543.
10. . ., . ., . .
// . . . . 1990. . 40. 2. . 230-237.
11. . ., . ., . ., . .
// .
. . . 2007. . 93. 7. . 799-805.
12. . ., . ., . ., . .,
. X., . . // . 2000.
. 45(3). . 514-519.

419

13. . . . . Opalina
ranarum // . ., 1962, 23, 74-76.
14. . . . , 2001. http: // masters.donntu.edu.ua /2001 /fvti /kuznetsov
/diss/lib/neuroosc/
15. . . ( 100 ,
- //
. . , 2006, -., 37 (3), 87-94.
16. . .,. . . . 100-
. . //
. . 77, 9. 2007. . 831-836.
17. . . . . , , 1975,
130-176.
18. - ., . . // ., , , 1975,
. 109-129.
19. . . , . . , - . .
,
http: // nisms.krinc.ru/papers/icnc02_2.pdf, 23
20. . ., . ., . , . .
: //
. . ., 2007, 43(5), 379.
21. . ., . . ,
// . . . 2003. 53 (4). 451-463.,
22. . ., . . : // . . . 2007, Jan.-Feb.; 57 (1): 5-22.
23. . ., . .
// . .: , 1973, . 163.
24. . ., . . // , 1972, . 21, 3, . 536-547,
25. . . //
, 1975, , , . 87-108.
26. Allen GC, Farnell Y, Bell-Pedersen D, Cassone VM, Earnest DJ. Effects of
altered Clock gene expression on the pacemaker properties of SCN2.2 cells and
oscillatory properties of NIH/3T3 cells. Neuroscience. 2004; 127 (4): 989-99.
27. Alving B. O. Spontaneous activity in isolated somata of Aplysia pacemaker
neurons J Gen Physiol. 1968, Jan.; 51 (1): 29-45.
28. Antonov I, Antonova I, Kandel ER, Hawkins RD.The contribution of activitydependent synaptic plasticity to classical conditioning in Aplysia. J Neurosci. 2001
Aug. 15; 21 (16): 6413-22, 32, 36.

420

29. Arinaminpathy Y., Philip C. Biggin, Indira H. Shrivastava1, Mark S.P.

Sansom_ A prokaryotic glutamate receptor: homology modelling and Molecular
dynamics simulations of GluR0 FEBS Letters 553 (2003) 321-327, 1.
30. Armus HL, Montgomery AR Aversive and attractive properties of electrical
stimulation for Paramecium caudatum. Psychol Rep. 2001. Oct.; 89 (2): 342-4,
32, 36.
31. Armus HL, Montgomery AR, Gurney RL Discrimination learning and
extinction in paramecia (P. caudatum). Learn Behav. 2006, Nov.; 34 (4): 361-5.
32, 36.
32. Arvanitaki A, Chalazonitis N Inhibitory processes of intrinsically generated
current of low frequency from autorythmic neurons . C R Seances Soc Biol Fil.
1964;158:1674-7, 30.
33. Aton S, Colwell C, Harmar A, Waschek J, Herzog E Vasoactive intestinal
polypeptide mediates circadian rhythmicity and synchrony in mammalian clock
neurons. Nat Neurosci. 2005, 8. 476483., 12.
34. Aton SJ, Herzog ED Come together, right...now: Synchronization of rhythms
in a mammalian circadian clock. Neuron. 2005, 48. 531534., 11.
35. Audesirk G.and Strumwasser F. Circadian Rhythm of Neuron R15 of
Aplysia californica: In Vivo Photoentrainment Proc. Nat. Acad. Sci. USA. 1975.
Vol. 72, No. 6, pp. 2408-2412, 18.
36. Bal T, McCormick DAMechanisms of oscillatory activity in guinea-pig
nucleus reticularis thalami in vitro: a mammalian pacemaker. J. Physiol. 1993,
Aug.; 468: 669-91., 1.
37. Balsalobre A, Damiola F, Schibler U (1998) A serum shock induces gene
expression in mammalian tissue culture cells. Cell 93: 929937, 2.
38. Barnard Alun R., Nolan Patrick M. When Clocks Go Bad: Neurobehavioural
Consequences of Disrupted Circadian Timing PLoS Genet. 2008. May 30; 4 (5).
39. Barnes S, Jacklet JW Ionic currents of isolated retinal pacemaker neurons:
projected daily phase differences and selective enhancement by a phase-shifting
neurotransmitter. J Neurophysiol. 1997, Jun.; 77 (6): 3075-84.
40. Baumgarten R.I.von, Takeda R., Chen C.F. Isolation of pacemaker neurons
in Aplisia californica // Feder.Proc., 1971, vol. 30, pp. 490, Abs.
41. Bernard S., Gonze D., Cajavec B., Herzel H., Kramer A. SynchronizationInduced Rhythmicity of Circadian Oscillators in the Suprachiasmatic Nucleus PLoS
Computational Biology. April 2007. Volume 3. Issue 4. e68 668-679.
42. Brazhnik ES, Fox SE Intracellular recordings from medial septal neurons
during hippocampal theta rhythm. Exp Brain Res. 1997, May; 114 (3): 442-53, 16.
43. Brehm P.and Eckert R. An electrophysiological study of the regulation of
Ciliary beating frequency in paramecium // J. Physiol. (1978), 283, pp. 557-568.
44. Brian D. Burrell,1 Christie L. Sahley, 3 and Kenneth J. Muller1, 2 NonAssociative Learning and Serotonin Induce Similar Bi-Directional Changes in
Excitability of a Neuron Critical for Learning in the Medicinal Leech The Journal
of Neuroscience, February 15, 2001, 21 (4): 14011412.

421

45. Brucet V. G. and Pittendrigh C. S. Temperature independence in a unicellular

clock*PROC. N. A. S. 1956. VOL. 42, 676-682, 1.
46. Brugger P., Macas E., Ihlemann J. Do sperm cell remember? Behav. Brain
Res., 2002, Oct 17; 136 (1): 325-328.
47. Brunner M, Simons MJ, Merrow M. Lego clocks: building a clock from
parts. Genes Dev. 2008, Jun 1; 22 (11): 1422-6., 2.
48. Bnning E ber die Erblichkeit der Tagesperiodizitt bei den Phaseolus
Blttern. Jb wiss Bot 1932 81: 411418, 2.
49. Bnning, E. (1973). The Physiological Clock. 3rd ed. (New York: SpringerVerlag, 30.
50. Cacciatore TW, Rozenshteyn R, Kristan WB Jr Kinematics and modeling
of leech crawling: evidence for an oscillatory behavior produced by propagating
waves of excitation. J Neurosci. 2000. Feb 15; 20 (4): 1643-55., 14.
51. Camborov P, Hubka P, Sulkov I, Huln I. The pacemaker activity of
interstitial cells of Cajal and gastric electrical activity. Physiol Res2003; 52 (3)
275-84.
52. Caroline Geisler,1,2 Nicolas Brunel,3 and Xiao-Jing Wang Contributions
of intrinsic membrane dynamics with irregular neuronal discharges //
J Neurophysiol 94: 43444361, 2005., 25.
53. Casadio A., Martin K.C., Giustetto M., Zhu H., Chen M., Bartsch D.,
Bailey C.H., Kandel E.R. A transient, neuron-wide form of CREB-mediated longterm facilitation can be stabilized at specific synapses by local protein synthesis //
Cell. 1999, Oct. 15; 99 (2) pp. 221-37., 25.
54. Cerminara NL, Rawson JA Evidence that climbing fibers control an intrinsic
spike generator in cerebellar Purkinje cells. J Neurosci. 2004, May 12; 24 (19):
4510-7.
55. Chaplain R. A. Metabolic Control Of Neuronal Pacemaker Activity And
The Rhythmic Organization Of Central Nervous Functions J. exp. Biol. (1979), 8i,
113-130.
56. Chaurasia SS, Pozdeyev N, Haque R, Visser A, Ivanova TN, Iuvone PM.
Circadian clockwork machinery in neural retina: evidence for the presence of
functional clock components in photoreceptor-enriched chick retinal cell cultures.
Mol Vis. 2006, Mar 30; 12: 215-23.
57. Chen C.F.,Baumgarten R.I., von, and Harth O. Metabolic aspects of the
rhythmogenesis in Aplisia pacemaker neurons // Pfluegers Arch. der Physiol.,
1973, vol. 345, pp. 179-193.
58. Chen GQ, Cui C, Mayer ML, Gouaux E. . Functional characterization of a
potassium-selective prokaryotic glutamate receptor. Nature 1999, Dec 16; 402
(6763): 817-21.
59. Chen Z, Zheng M, Friesen WO, Iwasaki T.. Multivariable harmonic balance
analysis of the neuronal oscillator for leech swimming. J Comput Neurosci. 2008, Jul 29.
60. Chevalier M, Mironneau C, Macrez N, Quignard JF. Intracellular Ca(2+)
oscillations induced by over-expressed Ca(V)3.1 T-type Ca(2+) channels in NG10815 cells Cell Calcium. 2008, Jun 19. [Epub ahead of print], 2.

422

61. Christensen ST, Leick V, Rasmussen L, Wheatley DN. Signaling in unicellular

eukaryotes. Int Rev Cytol. 1998; 177: 181-253., 33, 38.
62. Clark K., Hennesseyt T., and Nelson D.External GTP alters the motility
and elicits an oscillating membrane depolarization in Paramecium tetraurelia //
Proc. Natl. Acad. Sci. USA May, 1993. Vol. 90, pp. 3782-3786, 25, 37.
63. Cuninkova L, Brown SA. Peripheral circadian oscillators: interesting
mechanisms and powerful tools. Ann N Y Acad Sci. 2008; 1129: 358-70., 3.
64. Curras MC, Kelso SR, Boulant JA . Intracellular analysis of inherent and
synaptic activity in hypothalamic thermosensitive neurones in the rat. Physiol.
1991; 440: 257-71, 24.
65. Del Negro Christopher a.,1,* Koshiya Naohiro,1,2,* Butera Robert J., Jr.,3
and c. Smith Jeffrey 1 Persistent sodium current, membrane properties and bursting
behavior of pre-Botzinger complex inspiratory neurons in vitro J Neurophysiol 88:
22422250, 2002;, 28.
66. Dumollard Rmi 1,2,*, Carroll John 2, Dupont3 Genevive and Sardet
Christian 1 Calcium wave pacemakers in eggs Journal of Cell Science 115, 35573564 2002.
67. Eckert R. and Naitoh Y. Passive electrical properties of paramecium and
problems of ciliary coordination // General Physiology. 1970. V. 55. P. 467-482.,
25, 28.
68. Eisenhart FJ, Cacciatore TW, Kristan WB Jr A central pattern generator
underlies crawling in the medicinal leech. J Comp Physiol [A]. 2000, Jul-Aug; 186
(7-8): 631-43., 14.
69. Fliessbach K, Trautner P, Quesada CM, Elger CE, Weber B.Neuroimage.
2007 Apr 15; 35 (3): 1330-7. Epub 2007, Feb 13. Cerebellar contributions to
episodic memory encoding as revealed by fMRI, 4.
70. Forti L, Cesana E, Mapelli J, DAngelo EJ Physiol. 2006 Aug 1; 574 (Pt 3):
711-29. Epub 2006, May 11. Ionic mechanisms of autorhythmic firing in rat cerebellar
Golgi cells, 3.
71. Garca-Muoz A, Barrio LC, Buo W Membrane potential oscillations in
CA1 hippocampal pyramidal neurons in vitro: intrinsic rhythms and fluctuations
entrained by sinusoidal injected current. Exp Brain Res. 1993; 97 (2): 325-33., 16.
72. Gartside I. B. Mechanism of substained increases of firing rate of neurones
in the rat cerebral cortex after polarization: role of proteins synthesis. Nature,
1968, v. 220, p. 383-384., 3.
73. Geisler C., Brunel N. and Xiao-Jing Wang. Contributions of intrinsic
membrane dynamics to fast network oscillations with irregular neuronal discharges
// Neurophysiol. 2005. 94: 43444361., 25.
74. Geisler Caroline,1,2 Brunel Nicolas,3 and Wang Xiao-Jing Contributions
of intrinsic membrane dynamics with irregular neuronal discharges // Neurophysiol
94: 43444361, 2005., 29.
75. Gmez-Beldarrain M, Garca-Monc JC, Rubio B, Pascual-Leone A Effect
of focal cerebellar lesions on procedural learning in the serial reaction time task //
Exp Brain Res. 1998, May; 120(1): 25-30., 25, 29.

423

76. Gonze D, Bernard S, Waltermann C, Kramer A, Herzel H. Spontaneous

synchronization of coupled circadian oscillators // Biophys J. 2005, Jul; 89 (1):
120-9., 25, 33, 38.
77. Grtz HD, Kuhlmann HW, Mllenbeck M, Tiedtke A, Kusch J, Schmidt HJ,
Miyake A. Intra- and intercellular communication systems in ciliates. Naturwissenschaften.
1999, Sep; 86 (9): 422-34., 3, 33, 38.
78. Guevorkian K, Valles JM Jr Swimming Paramecium in magnetically
simulated enhanced, reduced, and inverted gravity environments Proc Natl Acad
Sci U S A. 2006 Aug 29; 103 (35): 13051-6. Epub 2006, Aug 17., 33, 38.
79. Guilding C, Piggins HD. Challenging the omnipotence of the suprachiasmatic
timekeeper: are circadian oscillators present throughout the mammalian brain? Eur
J Neurosci. 2007, Jun; 25 (11): 3195-216., 3.
80. Habas C. Rev Neurol (Paris). 2001 Dec; 157 (12): 1471-97 [The cerebellum:
from motor coordination to cognitive function], 4.
81. Hahn J.,1. Kullmann P., Horn J. P. and. Levitan E.S D2 autoreceptors
chronically enhance dopamine neuron pacemaker activity // The Journal of
Neuroscience, May 10, 2006 26 (19): 52405247., 25, 29, 36.
82. Harris-Warrick RM Voltage-sensitive ion channels in rhythmic motor
systems. Curr Opin Neurobiol. 2002, Dec; 12 (6): 646-51., 21.
83. Harvey AW, Bovell NK Spontaneous alternation behavior in Paramecium
Learn Behav. 2006 Nov; 34 (4): 361-5, 27, 33, 38.
84. Hiroshi Yoshimura* The Potential of Caffeine for Functional Modification
from Cortical Synapses to Neuron Networks in the Brain Current Neuropharmacology,
2005, 3, 309-316 309, 27.
85. Honma S, Nakamura W, Shirakawa T, Honma K. Diversity in the circadian
periods of single neurons of the rat suprachiasmatic nucleus depends on nuclear
structure and intrinsic period. Neurosci Lett. 2004, Apr 1; 358 (3): 173-6., 4.
86. Honma S, Shirakawa T, Nakamura W, Honma K Synaptic communication
of cellular oscillations in the rat suprachiasmatic neurons.. Neurosci Lett. 2000,
Nov 17; 294 (2): 113-6.
87. Hsieh TF, Pan JT. Extracellular single-unit studies of suprachiasmatic
neurons in brain slices. Effects of serotonin, dopamine, carbachol and LHRH. Chin
J Physiol. 1990; 33 (3): 255-68., 10.
88. Hughes RN The value of spontaneous alternation behavior (SAB) as a test
of retention in pharmacological investigations of memory Neurosci Biobehav Rev.
2004, Sep; 28 (5): 497-505, 33, 38.
89. Ishii K., Hirose K. and Iino M. Ca2 shuttling between endoplasmic reticulum
and mitochondria underlying Ca2 oscillations // EMBO reports, 2006. V. 7. N. 4.,
25, 29.
90. Ito H, Kageyama H, Mutsuda M, Nakajima M, Oyama T, Kondo T.Nat
Struct Mol Biol. 2007 Nov; 14 (11): 1084-8. Epub 2007, Oct 28 Autonomous
synchronization of the circadian KaiC phosphorylation rhythm., 4.

424

91. Iwadate Y, Nakaoka Y Calcium regulates independently ciliary beat and

cell contraction in Paramecium cells Cell Calcium. 2008, Jan 5 [Epub ahead of
print, 33, 38, 39.
92. Iwasaki, H., and Kondo, T. (2004). Circadian timing mechanism in the
prokaryotic clock system of cyanobacteria. J. Biol. Rhythms 19, 436444., 3.
93. Jacklet JW, Barnes S.. Photoresponsive pacemaker neurons from the
dissociated retina of Aplysia. Neuroreport. 1993, Dec 13; 5 (3): 209-12, 19.
94. Johnson B. R.,. Schneider L. R, Nadim F.,2 and Harris-Warrick R. M.
Dopamine modulation of phasing of activity in a rhythmic motor network: contribution
of synaptic and intrinsic modulatory actions // J Neurophysiol. 2005. 94: 3101
3111, 30, 37.
95. Johnson B. R.,. Schneider L. R, Nadim F., 2 and Harris-Warrick R. M.
Dopamine modulation of phasing of activity in a rhythmic motor network: contribution
of synaptic and intrinsic modulatory actions // J. Neurophysiol. 2005. 94: 3101
3111,, 1, 25.
96. Johnson CH, Golden SS, Ishiura M, Kondo T. Mol Microbiol. 1996, Jul; 21
(1): 5-11. Circadian clocks in prokaryotes., 5.
97. Johnson Stephen M., Wiegel Liana M., and Majewski David J. Are pacemaker
properties required for respiratory rhythm generation in adult turtle brain stems in
vitro? Am J Physiol Regul Integr Comp Physiol 293: R901R910, 2007., 28.
98. Kageyama H, Nishiwaki T, Nakajima M, Iwasaki H, Oyama T, Kondo
T.Mol Cell. 2006 Jul 21; 23 (2): 161-71. Cyanobacterial circadian pacemaker: Kai
protein complex dynamics in the KaiC phosphorylation cycle in vitro., 5.
99. Kashiwakura Y., Hee Cheol Cho, PhD; Andreas S. Barth, MD Ezana
Azene, PhD; Eduardo Marbn, MD, PhD Pacemaker Channel Into the Heart Gene
Transfer of a Synthetic A Novel Strategy for Biological Pacing Circulation. 2006;
114: 1682-1686.), 15.
100. Kelso SR, Perlmutter MN, Boulant JA. Am J Physiol. 1982, Jan; 242 (1):
R77-84. Thermosensitive single-unit activity of in vitro hypothalamic slices., 5.
101. Khalsa S.B., Michel S, Block G.D. The role of extracellular sodium in the
mechanism of a neuronal in vitro circadian pacemaker // Chronobiol Int. 1997,
Jan+ADs-14 (1): p. 1., 25.
102. Klisch C, Mahr S, Meissl H Circadian activity rhythms and phase-shifting
of cultured neurons of the rat suprachiasmatic nucleus. Chronobiol Int. 2006; 23
(1-2): 181-90, 11.
103. Kondo T. A cyanobacterial circadian clock based on the Kai oscillator.
Cold Spring Harb Symp Quant Biol.Kondo T. Cold Spring Harb Symp Quant Biol.
2007; 72: 47-55. A cyanobacterial circadian clock based on the Kai oscillator., 5.
104. Kononenko NI, Kuehl-Kovarik MC, Partin KM, Dudek FE Circadian
difference in firing rate of isolated rat suprachiasmatic nucleus neurons Neurosci
Lett. 2008, May 16; 436 (3): 314-6., 23.
105. Kononenko NI, Osipenko ON. Monosynaptic pathway responsible for
generation of burst activity in neuron PPa1 of Helix pomatia. Neirofiziologiia.
1987; 19 (1): 20-8, 15.

425

106. Kononenko NI, Osipenko ON. Postsynaptic mechanisms of the initiation

of burst activity in neuron PPa1 of Helix pomatia in response to an interneuron]
Neirofiziologiia. 1987; 19 (1): 20-8, 15.
107. Kononenko NI, Storozhuk MV. Dopamine inhibits transmission between
the interneuron initiating pacemaker activity in a bursting neuron and the bursting
neuron on the snail Helix pomatia. Comp Biochem Physiol C. 1992, May; 102 (1):
17-22., 15.
108. Kononenko NI, Storozhuk MV.. Dopamine inhibits transmission between
the interneuron initiating pacemaker activity in a bursting neuron and the bursting
neuron on the snail Helix pomatia. Comp Biochem Physiol C. 1992, May; 102 (1):
17-22, 5.
109. Kononenko NI. Role of the axodendritic tree in the of helix bursting
neurons: generation of pacemaker activity functioning and propagation of action
potentials along the axon Neuroscience. 2000; 96 (2): 399-406, 6.
110. Koshiya N, Smith JC.Neuronal pacemaker for breathing visualized in
vitro Nature. 1999, Jul. 22; 400 (6742): 360-3, 28.
111. Kramer RH. Axonal contribution to subthreshold currents in Aplysia bursting
pacemaker neurons. Cell Mol Neurobiol. 1986, Sep.; 6 (3): 239-53., 17.
112. Kristan WB Jr, Eisenhart FJ, Johnson LA, French KA Development of
neuronal circuits and behaviors in the medicinal leech. Brain Res Bull. 2000, Nov.
15; 53 (5): 561-70, 13.
113. Kuhlmann H.-W. Photomovements in ciliated protozoa // Naturwiss.
1998. V. 85. P. 143-154, 33, 39.
114. Lakin-Thomas PL, Brody S Circadian rhythms in microorganisms: new
complexities. Annu Rev Microbiol. 2004, 58: 489519, 6.
115. Lam YW, Cohen LB, Wachowiak M, Zochowski MR. Odors elicit three
different oscillations in the turtle olfactory bulb. J. Neurosci. 2000, Jan. 15; 20 (2):
749-62, 21.
116. Lee B, Almad A, Butcher GQ, Obrietan K. Protein kinase C modulates the
phase-delaying effects of light in the mammalian circadian clock. Eur J Neurosci.
2007. Jul; 26 (2): 451-62, 19.
117. Lewis DV. Calcium-activated inward spike after-currents in bursting neurone
R15 of Aplysia. J Physiol. 1988. Jan.; 395: 285-302, 17.
118. Leznik E and Llins R. Role of gap junctions in synchronized neuronal
oscillations in the inferior olive. // J. Neurophysiol. 2005, 94: 24472456,., 25, 29
119. Liu C, Reppert SM GABA synchronizes clock cells within the
suprachiasmatic circadian clock. Neuron. 2000, 25: 123128, 12.
120. Llinas R, Sugimori M. Electrophysiological properties of in vitro Purkinje
cell somata in mammalian cerebellar slices. J. Physiol. 1980, Aug.; 305: 171-95, 23.
121. Llinas R. O. and Yarom Y. Properties and distribution of ionic conductances
generating electroresponsiveness of mammalian inferior olivary neurones in vitro
J. Physiol. (1981), 315, pp. 569-584 569, 22.
122. Lundkvist G, Block G (2005) Role of neuronal membrane events in circadian
rhythm generation. Methods Enzymol 393: 62342., 12.

426

123. Lundkvist GB, Block GD.Role of neuronal membrane events in circadian

rhythm generation. Methods Enzymol. 2005; 393: 623-42., 33, 39.
124. Maccaferri G. and McBain The hyperpolarization-activated current (Ih) and
its contribution to pacemaker activity in rat CAI hippocampal stratum oriens-alveus
interneurones // Journal of Physiology. 1996, 497. 1, pp. 1 19-130, 26, 30, 36.
125. Maratou E., Theophilidis G. An axon pacemaker: diversity in the mechanism
of generation and conduction of action potentials in snail neurons // Neuroscience,
2000+ADs-96(1): p. 1, 26.
126. Margolis David J.1 and Detwiler Peter B. Different Mechanisms Generate
Maintained Activity in ON and OFF Retinal Ganglion Cells The Journal of
Neuroscience, May 30, 2007 27 (22): 59946005, 21.
127. Marin-Burgin A, Eisenhart FJ, Kristan WB Jr, French KA. Embryonic
electrical connections appear to pre-figure a behavioral circuit in the leech CNS. J
Comp Physiol A Neuroethol Sens Neural Behav Physiol. 2006, Feb.; 192 (2): 12333, 14.
128. Marin-Burgin A, WB Jr, French KA From synapses to behavior:
Development of a sensory-motor circuit in the leech.. Dev Neurobiol. 2008, Mar.
28; 68 (6): 779-787.
129. Marshall Sarah P. and Lang Eric J. Inferior Olive Oscillations Gate
Transmission of Motor Cortical Activity to the Cerebellum The Journal of
Neuroscience, December 15, 2004 24 (50): 1135611367, 20.
130. Martin KC, Barad M, Kandel ER. Local protein synthesis and its role in
synapse-specific plasticity // Curr Opin Neurobiol. 2000, Oct.; 10 (5): 587-92, 34, 39.
131. Martin KC, Barad M, Kandel ER. Local protein synthesis and its role in
synapse-specific plasticity // Curr Opin Neurobiol. 2000, Oct.; 10 (5): 587-92, 26, 37.
132. Maywood E, Reddy A, Wong G, ONeill J, OBrien J, et al. Synchronization
and maintenance of timekeeping in suprachiasma suprachiasmatic circadian clock
cells by neuropeptidergic signaling. Curr Biol. 2006. 16: 599605, 12.
133. McClung C. Robertson Plant Circadian Rhythms Plant Cell. 2006. April;
18 (4): 792803, 30.
134. McDougall A, Sardet C. Function and characteristics of repetitive calcium
waves associated with meiosis.. Curr Biol. 1995, Mar. 1; 5 (3): 318-28, 20.
135. Medanic M, Gillette M. U. Serotonin regulates the phase of the rat
suprachiasmatic circadian pacemaker in vitro only during the subjective day // J.
Physiol (Lond), 1992, May; 450: 629., 26.
136. Meijer J. H., Rietveld W. J. Neurophysiology of the suprachiasmatic
circadian pacemaker in rodents // Physiol. Rev., 1989, Jul; 69 (3): 671., 26, 29.
137. Merrow Martha, Spoelstra1 Kamiel & Till Roenneberg The circadian
cycle: daily rhythms from behaviour to genes First in the Cycles Review Series
EMBO reports. VOL 6. NO 10. 2005 930-934, 6.
138. Mieda M., Williams S. C., Richardson J. A., Tanaka K. and Yanagisawa M.
The dorsomedial hypothalamic nucleus as a putative food-entrainable circadian
pacemaker // PNAS 2006. 8, vol. 103, no. 321215012155, 26, 29.

427

139. Mimikakis JL, Nelson DL, Preston RR Biochem J. 47 Oscillating response

to a purine nucleotide disrupted by mutation in Paramecium tetraurelia. 1998, Feb.
15; 330 (Pt 1): 139, 34, 39.
140. Mirmiran M, Koster-Van Hoffen GC, Bos NP. Circadian rhythm generation
in the cultured suprachiasmatic nucleus. Brain Res Bull. 1995; 38 (3): 275-83., 10
141. Monsivais P, Clark BA, Roth A, Husser M Determinants of action potential
propagation in cerebellar Purkinje cell axons. J. Neurosci. 2005, Jan. 12; 25 (2):
464-72, 23.
142. Moortgat K.T., Bullock T.H., Sejnowski T..J Precision of the pacemaker
nucleus in a weakly electric fish: network versus cellular influences // J.
Neurophysiol. 2000, Feb.; 83 (2) p. 971, 26, 34, 39.
143. Moortgat KT, Bullock TH, Sejnowski TJ. Precision of the pacemaker
nucleus in a weakly electric fish: network versus cellular influences. J. Neurophysiol.
2000, Feb; 83 (2): 971-83, 7.
144. Moortgat KT, Keller CH, Bullock TH, Sejnowski TJ. Submicrosecond
pacemaker precision is behaviorally modulated: the gymnotiform electromotor
pathway. Proc Natl Acad Sci U S A. 1998, Apr. 4; 95 (8): 4684-9R, 6.
145. Mori T. and Johnson C. H. Independence of circadian timing from cell
division in cyanobacteria Journal of Bacteriology, Apr., 2001, p. 24392444 Vol.
183, 8, 6.
146. Nahm SS, Farnell YZ, Griffith W, Earnest DJ. Circadian regulation and
function of voltage-dependent calcium channels in the suprachiasmatic nucleus. J.
Neurosci. 2005, Oct. 5; 25 (40): 9304-8, 7.
147. Nahm SS, Farnell YZ, Griffith W, Earnest DJ. J Neurosci. 2005, Oct. 5;
25 (40): 9304-8 Circadian regulation and function of voltage-dependent calcium
channels in the suprachiasmatic nucleus, 7.
148. Nakamura W, Honma S, Shirakawa T, Honma K Eur J Neurosci. 2001,
Aug; 14 (4): 666-74. Regional pacemakers composed of multiple oscillator neurons
in the rat suprachiasmatic nucleus, 7.
149. Nazarenko L. V., Andreev Igor M., Lyukevich Alexander A Pisareva
Tatiana V. and. Los Dmitry A Calcium release from Synechocystis cells induced by
depolarization of the plasma membrane: MscL as an outward Ca2+ channel
Microbiology (2003), 149, 11471153, 7.
150. Niktarash A. H. Transmission of the subthalamic nucleus oscillatory activity
to the cortex: a computational approach // Journal of Computational Neuroscience,
2003, 15, 223232, 29.
151. Niktarash A. H. Transmission of the subthalamic nucleus oscillatory activity
to the cortex: a computational approach // Journal of Computational Neuroscience,
2003, 15, 223232, 26.
152. Oami K. Membrane potentials responses controlling chemodispersal of
Paramecium caudatum from quinine. \\ J Comp Physiol A (1996) 178: 307-316
(), 34, 39.

428

153. Pape HC, Driesang RB Ionic mechanisms of intrinsic oscillations in neurons

of the basolateral amygdaloid complex. J Neurophysiol. 1998, Jan.; 79 (1): 217-26, 8.
154. Parnas I., Sturmwasser Z. Prolonget excitatory and inhibitory synaptic
modulation of a bursting pacemaker neuron // J. Neurophys. 1975, v. 37, p. 594., 26.
155. Passano L M. and McCullough C. B. The light response and the rhythmic
potentials of hydra // Zoology: 1962, vol. 48, 1376-1382, 37.
156. Passano L M. and McCullough C. B. The light response and the rhythmic
potentials of hydra // Zoology: 1962, vol. 48, 1376-1382., 26.
157. Pedroarena Ch. and Llinas R. Dendritic calcium conductances generate
high-frequency oscillation in thalamocortical neurons // Proc. Natl. Acad. Sci.
USA. 1997, Vol. 94, pp. 724-728., 26, 35.
158. Pena1 Fernando,2 and Ramirez Jan-Marino 1 Substance P-Mediated
Modulation of Pacemaker Properties in the Mammalian Respiratory Network The
Journal of Neuroscience, August 25, 2004 24 (34): 75497556 7549, 28.
159. Perkel DH, Schulman JH, ullock TH, Moore GP, Segundo JP. Pacemaker
neurons: effects of regularly spaced synaptic input. Science. 1964, Jul. 3; 145: 61-3, 8.
160. Pittendrigh C. S. Circadian systems, i. the driving oscillation and its assa y
in drosophila pseudoobscura* PROC. N. A. S. 1967. VOL. 58, Physiology: 1762-8.
161. Pittendrigh, C.S. (1993). Temporal organization: reflections of a Darwinian
clock-watcher. Annu. Rev. Physiol. 55, 1654, 8.
162. Pitts GR, Ohta H, McMahon DG.Brain Res. Daily rhythmicity of largeconductance Ca2+ -activated K+ currents in suprachiasmatic nucleus neurons.
2006 Feb 3;1071(1):54-62. Epub. 2006, Jan. 17, 10.
163. Precht J.C., Bullock T. H., and Kleinfeld D. Direct evidence for local
oscillatory current sources and intracortical phase gradients in turtle visual cortex
PNAS 18, 2000, vol. 97, no. 2, 877882, 20.
164. Preston RR, Usherwood PN. L-glutamate-induced membrane
hyperpolarization and behavioural responses in Paramecium tetraurelia. J. Comp
Physiol [A]. 1988, Nov.; 164 (1): 75-82., 34, 39.
165. Preston RR, Van Houten JL Localization of the chemoreceptive properties
of the surface membrane of Paramecium tetraurelia. J. Comp Physiol [A]. 1987,
Apr.; 160 (4): 537-41, 34, 39.
166. Prosser R.A., McArthur A.J., Gillette M.U. cGMP induces phase shifts of
a mammalian circadian pacemaker at night, in antiphase to cAMP effects //
Proc. Natl. Acad. Sci. U S A, 1989, Sep.; 86 (17): 6812., 26.
167. Rabbah P, Nadim F. Distinct synaptic dynamics of heterogeneous pacemaker
neurons in an oscillatory network // J. Neurophysiol. 2007, Jan. 3; 26, 29.
168. Rabbah P, Nadim F. Distinct synaptic dynamics of heterogeneous pacemaker
neurons in an oscillatory network J. Neurophysiol. 2007, Jan. 3; [Epub ahead of
print, 34, 39.
169. Ramirez J. M., Tryba A. K., Pena F. Pacemaker neurons and neuronal
networks: an integrative view // Curr Opin Neurobiol. 2004, Dec.; 14 (6): 665-74, 26.

429

170. Ramirez JM, Garcia Iii A. Medullary Pacemaker Neurons are Essential
for Both Eupnea and Gasping in Mammals- // The Last Word J Appl Physiol.
2007, Apr. 26, 26, 29.
171. Roenneberg Till 1 and Merrow Martha 2 Circadian Clocks: Translation
Lost Current Biology. Vol 15, No 12, PP., 8.
172. Rose T, Gras H, Hrner M. Activity-dependent suppression of spontaneous
spike generation in the Retzius neurons of the leech Hirudo medicinalis L. Invert
Neurosci. 2006, Dec; 6 (4): 169-76. Epub. 2006, Oct. 31, 14.
173. Ruta V, Jiang Y, Lee A, Chen J, MacKinnon R.. Functional analysis of an
archaebacterial voltage-dependent K+ channel. Nature. 2003, Mar. 13; 422 (6928):
180.
174. Sardet C, Roegiers F, Dumollard R, Rouviere C, McDougall A. Calcium
waves and oscillations in eggs.. Biophys Chem. 1998, May. 5; 72 (1-2): 131-40., 20.
175. Scott IM, Boulant JA.Brain Res. 1984, Jul. 23; 306 (1-2): 157-63.
Dopamine effects on thermosensitive neurons in hypothalamic tissue slices.
2007; 72: 47-55., 9.
176. Shibata S, Moore RY Electrical and metabolic activity of suprachiasmatic
nucleus neurons in hamster hypothalamic slices. Brain Res. 1988, Jan. 12; 438
(1-2): 374-8.
177. Simon S. M. and Llinas R.R. Compartmentalization of the submembrane
calcium activity during calcium influx and its significance in transmitter release //
Biophysical journal, 1985. V. 48 p. 485-497.
178. Soto-Trevio C, Rabbah P, Marder E, Nadim F Computational model of
electrically coupled, intrinsically distinct pacemaker neurons J Neurophysiol. 2005,
Jul.; 94 (1): 590-604. Epub. 2005, Feb. 23., 34, 40.
179. Sotty F., Danik M., Manseau F., Laplante F., Quirion R.and Williams
S.Distinct electrophysiological properties of glutamatergic, cholinergic and
GABAergic rat septohippocampal neurons: novel implications for hippocampal
rhythmicity J. Physiol (2003), 551.3, pp. 927943, 14.
180. Sugino K, Tominaga T., Allen R.D. and Naitoh Y. Electrical properties
and fusion dynamics of in vitro membrane vesicles derived from separate parts of
the contractile vacuole complex of Paramecium multimicronucleatum // The Journal
of Experimental Biology. 208, 3957-3969, 34, 40.
181. Sujino M, Masumoto KH, Yamaguchi S, van der Horst GT, Okamura H,
Inouye ST. Suprachiasmatic nucleus grafts restore circadian behavioral rhythms of
genetically arrhythmic mice. // Curr Biol. 2003, Apr. 15; 13 (8): 664-8, 26.
182. Taddei-Ferretti C, Cordella L.Modulation of Hydra attenuata rhythmic
activity: phase response curve // J Exp Biol. 1976, Dec.; 65 (3): 737-51., 26.
183. Thompson RF Are memory traces localized or distributed? Neuropsychologia.
1991; 29 (6): 571-82, 26.
184. Toller GL, Nagy E, Horvath RA, Klausz B, Rekasi Z Circadian expression
of Bmal1 and serotonin-N-acetyltransferase mRNAs in chicken retina cells and
pinealocytes in vivo and in vitro. J Mol Neurosci. 2006; 28 (2): 143-50., 21.

430

185. Tomida T, Hirose K, Takizawa A, Shibasaki F, Iino M (2003) NFAT

functions as a working memory of Ca2+ signals in decoding Ca2+ oscillation //
EMBO. J. 22: 38253832, 26, 29.
186. Urbano FJ, Simpson JI, Llins RR. Somatomotor and oculomotor inferior
olivary neurons have distinct electrophysiological phenotypes. Proc Natl Acad Sci
USA. 2006, Oct 31; 103 (44): 16550-5. Epub. 2006, Oct. 18, 22.
187. Van Houten J. Chemosensory transduction in Paramecium // Europ. J.
Protostiol. 1998. V. 34. P. 301-307, 2, 35, 40.
188. Vansteensel MJ, Michel S, Meijer JH. Organization of cell and tissue
circadian.
189. Varga V, Hangya B, Krnitz K, Ludnyi A, Zemankovics R, Katona I,
Shigemoto R, Freund TF, Borhegyi Z The presence of pacemaker HCN channels
identifies theta rhythmic GABAergic neurons in the medial septum. J. Physiol.
2008, Aug 15; 586 (16): 3893-915. Epub. 2008, Jun. 19, 16.
190. Victor A. Maltsev, PhD and Edward G. Lakatta, MD Normal heart rhythm
is initiated and regulated by an intracellular Calcium clock within pacemaker cells
Heart Lung Circ. 2007. October; 16 (5): 335348., 22.
191. Wassenberg JJ, Clark KD, Nelson DL. Effect of SERCA pump inhibitors
on chemoresponses in Paramecium // J. Eukaryot Microbiol. 1997, Nov.-Dec.; 44
(6): 574-81, 27.
192. Wassenberg JJ, Clark KD, Nelson DL. Effect of SERCA pump inhibitors
on chemoresponses in Paramecium // J. Eukaryot Microbiol. 1997, Nov.-Dec.; 44
(6): 574-81. 47, 28.
193. Watanabe A, Shibata S, Watanabe S. Circadian rhythm of spontaneous
neuronal activity in the suprachiasmatic nucleus of old hamster in vitro. Brain Res.
1995, Oct. 16; 695 (2): 237-9, 10.
194. Welsh DK, Logothetis DE, Meister M, Reppert SM (1995) Individual
neurons dissociated from rat suprachiasmatic nucleus express independently phased
circadian firing rhythms. Neuron. 14: 697706, 9.
195. Wittner L, Miles R Factors defining a pacemaker region for synchrony in the
hippocampus. J. Physiol. 2007, Nov. 1; 584 (Pt 3): 867-83. Epub. 2007, Sep. 6, 16.
196. Wong R K and M Stewart Different firing patterns generated in dendrites
and somata of CA1 pyramidal neurones in guinea-pig hippocampus. J. Physiol.
1992; 457; 675-687.
197. Woolum J. C. and Strumwasser F. The differential effects of ionizing
radiation on the circadian oscillator and other functions in the eye of Aplysia
(x-rays/molluskan eye) Proc Natl Acad Sci U S A. 1980, Sep.; 77 (9): 5542-6.
198. Wright CL, Burgoon PW, Bishop GA, Boulant JA.Am J Physiol Regul
Integr Comp Physiol. 2008, May; 294 (5): R1704-15. Epub. 2008, Mar. 5. Cyclic
GMP alters the firing rate and thermosensitivity of hypothalamic neurons., 9.
199. Yamaguchi S, Isejima H, Matsuo T, Okura R, Yagita K. Synchronization of
cellular clocks in the suprachiasmatic nucleus. Science. 2003, 302: 14081412, 12.

431

200. Yoshimura Hiroshi The Potential of Caffeine for Functional Modification

from Cortical Synapses to Neuron Networks in the Brain Current Neuropharmacology,
2005, 3, 309-316, 17.
201. Young MW, Kay SA. Time zones: a comparative genetics of circadian
clocks Nat Rev Genet. 2001, Sep.; 2 (9): 702-15, 29.
202. Young, M. W., and Kay, S.A. (2001). Time zones: a comparative genetics
of circadian clocks. Nat. Rev. Genet. 2, 702715., 9.
203. Zhao Y, Boulant JA. Temperature effects on neuronal membrane potentials
and inward currents in rat hypothalamic tissue slices. J Physiol. 2005, Apr. 1; 564
(Pt 1): 245-57., 24.
204. Zhi-Gen Jiang*, YuQin Yang*, Zhao-Ping Liut and Charles N. Al
Membrane properties and synaptic inputs of suprachiasmatic nucleus neurons in
rat brain slices Journal of Physiology (1997), 499.1, pp. 141-159, 9.
205. Zimmerman N. and Menakert M. The pineal gland: A pacemaker within
the circadian system of the house sparrow Proc. Natl. Acad. Sci. USA. Vol. 76,
No. 2, pp. 999-1003, February 1979.

432

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, , , , , .

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. .: , 2003. 671 .
4. . . . .: , 1978. 214 .
5. . . . .: , 2005. 368 .
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, 1997. 396 .

1. Astion M. L., Chavatal A., Orkand R.K. Further studies of electrogenic Na+/
HCO3 cotransport in glial cells of Necturus optic nerve: Regulation of pH // Glia.
1991. Vol. 4. Issue 4. PP. 461-468.
2. Billups B., Attwell D. Modulation of non-vesicular glutamate release by pH
// Nature. 1999. No 379. PP. 171-174.
463

3. Boehler M. D., Wheeler B. C., Brewer G. J. Added astroglia promote greater

synapse density and higher activity in neuronal networks // Neuron Glia Biology.
2007. Vol. 3. Issue 02. PP. 127-140.
4. Broeke J. Oligodendrocyte transfected with GFP (Green Fluorescent Protein)
recorded using a 63x objective 3 days in vitro mouse // . : http: // commons.wikimedia.org.
5. Butt A. M., Ransom B. R. Morphology of astrocytes and oligodendrocytes
during development in the intact rat optic nerve // J. Compar. Neurol. 1993. Vol.
338. Issue 1. PP. 141-158. Brown H. C., Perry V. H. Differential adhesion of
macrophages to white and grey matter in an in vitro assay // Glia. 1998. Vol. 23.
Issue 4. PP. 361-373.
6. David S., Aguayo A. J. Axonal elongation into peripheral nervous system
bridges after central nervous system injury in adult rats // Science. 1981. No
214. PP. 931-933.
7. Ekblom P., Timpl P. (Eds.). The laminins // Amsterdam. Harwood Academic
Publishers. 1996.
8. Fields R. D. Advances in understanding neuronglia interactions // Neuron
Glia Biology. 2006. Vol. 2. PP. 2326.
9. Haber M., Murai K. K. Reshaping neuronglial communication at hippocampal
synapses // Neuron Glia Biology. 2006. Vol. 2. PP. 5966.
10. Hatten V. E. Central nervous system neuronal migration // Annu Rev
Neurosci. 1999. Vol. 22. PP. 511-39.
11. Henn F. A., Hamberger A. Glial Cell Function: Uptake of Transmitter
Substances (rabbit brain/-y-aminobutyric acid/serotonin/dopamine/norepinephrine)
// Proc. Nat. Acad. Sci. USA. 1971. Vol. 68, No. 11, pp. 2686-2690.
12. Kuffler S. W, Nicholls J. G. The physiology of neuroglial cells // Ergeb
Physiol. 1966. Vol. 57. PP. 190.
13. Kuffler S. W., Nicholls J. G., and Orkand R. K. Physiological properties of
glial cells in the central nervous system of amphibia // J. Neurophysiol. 1966.
Vol. 29. PP. 768-787.
14. McGlade-McCulloh E., Morrissey A. M., Norona F., and Muller K. J.
Individual microglia move rapidly and directly to nerve lesions in the leech central
nervous system // Proc. Natl. Acad. Sci. USA. 1989. Vol. 86. PP. 1093-1097.
15. Nagler K., Mauch D. H., Pfrieger F. W. Glia-derived signals induce synapse
formation in neurones of the rat central nervous system // J. Physiol. 2001. Vol.
533. PP. 665-679.
16. Matsui K., Hosoi N., Tachibana M. Active Role of Glutamate Uptake in the
Synaptic Transmission from Retinal Nonspiking Neurons // J. Neurosci. 1999.
Vol. 19. PP. 6755-6766.
17. Newman E. A. Membrane physiology of retinal glial (Muller) cells // J. of
Neuroscience. 1985. Vol. 5. PP. 2225-2239.
18. Newman E. A. Electrophysiology of Retinal Glial Cells // Progress in
Retinal Research. 1988. Vol. 8. PP. 153-171.

464

19. Newman E. A., Reichenbach A. The Mller cell: a functional element of the
retina // Trends Neurosci. 1996. Vol. 19. PP. 307-312.
20. Newman E. A., Zahs K. R. Modulation of neuronal activity by glial cells in
the retina // J. Neurosci. 1998. Vol. 18. PP. 4022-4028.
21. Nicholson C. Volume transmission in the year 2000 // Prog. Brain Res.
2000. Vol. 125. PP. 437-446.
22. Nishida H, Okabe S. Visualization of synapse-glia dynamics // Brain
Nerve. 2007. Jul. 59 (7). PP. 755-61.
23. Nishiyama H., Knopfel T., Endo S., and Itohara S. Glial protein S100B
modulates long-term neuronal synaptic plasticity // PNAS USA. 2002. Vol. 99.
No 6. PP. 4037-4042.
24. Notterpek L., Shooter E. M., Snipes G. J. Upregulation of the EndosomalLysosomal Pathway in the Trembler-J Neuropathy // J. Neurosci. 1997. Vol. 17.
PP. 4190-4200.
25. Odette L. L., Newman E. A. Model of potassium dynamics in the central
nervous system // Glia. 1988. Vol. 1. PP. 198-210.
26. Orkand R. K., Nicholls J. G., and Kuffler S. W. Effect of nerve impulses on
the membrane potential of glial cells in the central nervous system of amphibia /
/ J. Neurophysiol. 1966. Vol. 29. PP. 788-806.
27. Paterniti M. Driving Mr. Albert: A Trip across America with Einsteins
Brain // Publisher: Delta. 2001.
28. Paulson O. B., Newman E. A. Does the release of potassium from astrocyte
endfeet regulate cerebral blood flow? // Science. 1987. Vol. 237. PP. 896-898.
29. Pfrieger F. W., Barres B. A. New views on synapse-glia interactions //
Curr. Opin. Neurobiol. 1996. Vol. 6. PP. 615-621.
30. Piet R., Vargova L., Sykova E., Poulain D. A., Oliet St.H.R. Physiological
contribution of the asrtocytic environment of neurons to intersynaptic crosstalk /
/ PNAS. 2004. Vol. 101. No 7. PP. 2151-2155.
31. Porter J. T., Mccarthy K. D. Astrocytic neurotransmitter receptors in situ
and in vivo // Prog. Neurobiol. 1997. Vol. 51. Issue 4. PP. 439-455.
32. Rakic P. Neuronalglial interaction during brain development // Trends
in Neurosci. 1981. Vol. 4. PP. 184-187.
33. Ransom B. R., Fern R. Does astrocytic glycogen benefit axon function and
survival in CNS white matter during glucose deprivation? // Glia. 1997. Vol. 21.
Issue 1. PP. 134-141.
34. Rasband M. N., Peles E., Trimmer J. S., Levinson S. R., Lux S. E., Shrager
P. Dependence of nodal sodium channel clustering on paranodal contact in the
developing CNS // J. Neurosci. 1999. Vol. 19. PP. 7516-7528.
35. Ritchie J. M., Black J. A., Waxman S. G., and Angelides K. J. Sodium
channels in the cytoplasm of Schwann cells // Proc. Natl. Acad. Sci. USA. 1990.
Vol. 87. PP. 9290-9294.
36. Roitbak T., Sykova E. Diffusion barriers evoked in the rat cortex by reactive
astrogliosis // Glia. 1999. Vol. 28. PP. 40-48.

465

37. Smith P. J., Howes E. A., Treherne J. E. Mechanisms of glial regeneration

in an insect central nervous system // J. Exp. Biol. 1987. Vol. 132. PP. 59-78.
38. Sherwood Ch. C., Stimpson C. D., Raghanti M. A., Wildman D.E., Uddin
M. et.al. Evolution of increased glianeuron ratios in the human frontal cortex //
PNAS. 2006. Vol. 103. PP. 13606-13611.
39. Shrager P., Chiu S. Y., Ritchie J. M. Voltage-dependent sodium and
potassium channels in mammalian cultured Schwann cells // Proc. Natl. Acad.
Sci. USA. 1985. Vol. 82. PP. 948-952.
40. Son Y. J., Thompson W. J. Schwann cell processes guide regeneration of
peripheral axons // Neuron. 1995 a. No 14. PP. 125-132.
41. Son Y. J., Thompson W. J. Nerve sprouting in muscle is induced and guided
by processes extended by Schwann cells // Neuron. 1995 b. No 14. PP. 133-141.
42. Stevens B., Porta S., Haak L. L., Gallo V., Fields R. D. Adenosine: A NeuronGlial Transmitter Promoting Myelination in the CNS in Response to Action Potentials
// Neuron. 2002. Vol. 36. No 5. PP. 855-868.
43. Sul J. Y., Orosz G., Givens R. S., Haydon Ph. G. Astrocytic Connectivity
in the Hippocampus // Neuron Glia Bilogy. 2004. Vol. 1. PP. 3-11.
44. Szatkowski M., Barbour B., Attwell D. Non-vesicular release of glutamate
from glial cells by reversed electrogenic glutamate uptake // Nature. 1990. Vol.
348. PP. 443-446.
45. Sykova E. Glial diffusion barriers during aging and pathological states //
Prog. Brain Res. 2001. Vol. 132. PP. 339-363.
46. Takahashi M., Billups B., Rossi D., Sarantis M., Hamann M., and Attwell
D. The role of glutamate transporters in glutamate homeostasis in the brain //
J. Exp. Biol. 1997. Vol. 200. PP. 401-409.
47. Tao-Cheng J. H., Nagy Z., Brightman M. W. Astrocytic orthogonal arrays
of intramembranous particle assemblies are modulated by brain endothelial cells in
vitro // J. Neurocytol. 1990. Vol. 19. PP. 143-153.
48. Todd K. J., Serrano A., ., Robitaille R. Glial cells in synaptic plasticity //
J. Physiol. Paris. 2006. Mar-May. Vol. 99(2-3). PP. 75-83.
49. Willbold E., Reinicke M., Lance-Jones C., Lagenaur C., Lemmon V., Layer
P.G. Mller Glia Stabilizes Cell Columns During Retinal Development: Lateral Cell
Migration but not Neuropil Growth is Inhibited in Mixed Chick-Quail Retinospheroids
// Eur. J. Neurosci. 1995. Vol. 7. PP. 2277-2284.
50. Journal Neuron-Glia Biology: www.journals.cambridge.org/jid-NGB.

466

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2008). 1943 .
.
, , - - , .
1943 ., , ,
. . -
,
(A Logical Calculus of the Ideas Immanent
in Nervous Activity).
, . -

480

 - (MCP). , : ,
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, .
, ,
, . .
, ,
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. (1982 .),
-
(1982 .), . ,
, . (1987 .), -

482

 - (1987 .)
. . -, , - , (. . , 19-38).
,
. : NeuroSolutions
NeuroDimension, NeuralWorks
NeuralWare, Process Advisor AI Ware,
NeuroShell Word Systems Group, BrainMaker Pro
California Scientific SoftWare, Stanistica Neural Networks StatSoft, -, .
. ,

. ,
, ,
Nihon Denki Fujitsu Neuro-07 FMR 50 (1988 .),
. . (1989 .),
. . (1992 .),
Intel Ni1000, 1024 (1994 .),
SCX-1,
() (FPGA)
.. ( . ., 2000).
2008 . Hewlett-Packard
, . , , . Hewlett-Packard
, 1971 . . Hewlett-Packard .

483

 .
, ,
. , , ,
. , RRAM (resistive random-access memory),
,
(Strukov D. B. t. al., 2008).
,
,
,
.
15.4.
, ,
.

--. , ( 1967 ., . .
).
, , .
.
, .
, , 1980- , . .
,
. , 1980 . , ,
,
... , ,
, ,
484

 ( ., 2008). 1993 .
,

(Kohonen T., 1993). ,
(. .
, 41-42).
, , ( ) . .
. ,
, . , , : (1) ,
; (2) ,
, , ( . .,
2006); (3) ; (4) . ,
, , .
.
, . , , , ,
, .
-
, ,
, , .
, in silico. 1996 . E-CELL, . , Institute
for Advanced Biosciences at Keio University (http://www.e-cell.org/ecell).

485

 2005 .
IBM , Blue Brain Project.

10 . .
( ), :
1. .
2. .
:
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- .
1.2.
.
. ,

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, .

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486

 ( , )
,

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,
,
,
.
( ), :
(1) (2)
.
(1.1.) (1.2.)
.

1. : / . . . . , .. . .: , 2001. 840 .
2. . . .: . , 2008. 655 .
3. . . . .: , 2000. 528 .
4. . ., . ., . ., . .
. .: , 2003. 672 .
5. .. . // : , . 2006. 4-5. . 4-14.

1. . ., . ., . ., . . . .: , 1988. . 22.
2. . ., . ., . . . .: , 2002. 480 .
487

3. / . . . .
.: , 2003. 368 .
4. / . . . . . . .: , 2003. 224 .
5. / .
. . . . . .: , 2003. 192 .
6. . . . .:
, 2007. 224 .
7. Hebb D. O. The Organisation of Behavior: A Neuropsychological Theory.
New York: Wiley, 1949. p. 62-63.
8. . . // . .:
, : , 1996 . 245.
9. . . . .: , 1995.
. 250-252.
10. . , , . .: , 1990. . 198-225.
11. An Enriched Environment Stimulates An Increase In The Number Of
Nerve Cells In Brains Of Older Mice // Press releases The Salk Institute for
Biological Studies, April 24, 1997.
12. Running Boosts Number Of Brain Cells, According To New Salk Study //
Press releases The Salk Institute for Biological Studies, February 22, 1999.
13. Exercise Makes Mice Smarter, Salk Scientists Demonstrate // Press
releases The Salk Institute for Biological Studies, November 08, 1999.
14. Salk Scientists Demonstrate For The First Time That Newly Born Brain
Cells Are Functional In The Adult Brain // Press releases The Salk Institute for
Biological Studies, February 27, 2002.
15. Neural Progenitor Cells Recovered From Postmortem And Adult Tissue //
Press releases The Salk Institute for Biological Studies, May 02, 2001.
16. Jumping genes contribute to the uniqueness of individual brains //
Press releases The Salk Institute for Biological Studies, June 14, 2005.
17. . ., . ., . . . .:
:, 2008. . 60.
18. - ., . ,
// : / .
. . . , . . . .: , 2001. . 5-22.
19. . : . .: , 2006. 1104 .
20. . ., . , .
. .: , 2000. 382 .
21. . . : , .
.: , 2001. 256 .
22. ., ., . . .:
, 2002. 272 .
23. . . . .: , 2002. 448 .

488

24. . ., . . . Matlab 6. .: , 2002. 496 .

25. . . . .: , 2003.
144 .
26. . . :
. .: , 2003. 864 .
27. . . .:
, 2003. 288 .
28. . . .: , 2004. 344 .
29. . . . . .: ,
2005. 256 .
30. ., ., . , . .: , 2006. 452 .
31. . . . .:
-, 2006. 192 .
32. . . , , : . .: , 2006. 224 .
33. . . Data Mining. .: - ; . , 2006. 382 .
34. . . . .: -
; . , 2007. 352 .
35. . . . .: ; . , 2008.
316 .
36. . ., . . . .: -
. . . , 2004. 400 .
37. . . . .:
, 2004. 221 .
38. . ., . ., . ., . . : . .:
, 2008. 152 .
39. Strukov D. B., Snider G. S., Stewart D. R.,Williams R. S. The missing
memristor found // Nature. 453. pp. 80-83.
40. Kohonen T. Neural Networks. 1993. 6. p. 895.
41. . ., . . . .: , 2003. . 241-243.
42. . ., . .
// . 2001. 2. . 692-698.
43. . . .
.: - , 2006. . 110.
44. http://www.e-cell.org/ecell

489

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1. - ., . ,
// : / .
. . . , . . . .: , 2001. . 522.
2. ., .. // : / . . . . , . . . .: , 2001. . 58-73.
3. . : . .: , 2006. 1104 .
4. ., . : .
.: , 2006. 1408 .
5. .., ., .
. .: , 2000. 382 .
6. .. : , . .:
, 2001. 256 .
7. ., ., . . .:
, 2002. 272 .
8. . . .:
, 2003. 288 .
9. . . .:
, 2004. 344 .

1. . . // http://www.intuit.ru/
department/expert/neuro/2/2.html
2. . . . . // http://www.intuit.ru/department/expert/neuro/
2/3.html
3. . . . . // http://www.intuit.ru/department/expert/neuro/
2/5.html
4. . . . . // http://www.intuit.ru/department/expert/neuro/
2/4.html
5. . . . .: , 2000. 528 .
6. Tseng Kuei Y., ODonnell Patricio. DopamineGlutamate Interactions
Controlling Prefrontal Cortical Pyramidal Cell Excitability Involve Multiple Signaling
Mechanisms // The Journal of Neuroscience. 2004. June 2. 24(22). P. 5131-5139.

511

 17.

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. (, ).
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 ,
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(1) ,

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.

1. . ., . ., . ., . .
. .: , 2003. 672 .
2. . , , . .: , 1990. 239 .
533

3. . . . .: , 1995. 384 c.
4. . ., . ., . ., . .
. .: , 1988. 96 c.
5. . . . .: . , 2008. 359 .

1. Wang Z., Edwards J. G., Riley N., Provance D. W., Karcher R., Li X., Davison I. G.,
Ikebe M, Mercer J. A., Kauer J. A., Ehlers M. D. Myosin Vb Mobilizes Recycling
Endosomes and AMPA Receptors for Postsynaptic Plasticity // Cell. 2008. 31
October. Volume 135, Issue 3. P. 535-548.
2. Gelbard-Sagiv H., Mukamel R., Harel M., Malach R., Fried I. Internally Generated
Reactivation of Single Neurons in Human Hippocampus During Free Recall
// Science. 2008. October, 3. Vol. 322. 5898. P. 96-101.

534

 18.

. - ,
. ,


, .
( , in silico)
. , :

;
;
,
, , ,
, , ,
;
,
, .

, ,
.
( . .,
. ., 2007, . 348). ,
535

 , - .
, , , - .
C ,

( . ., 2007, . 361-376).

(E-CELL),
1996 . Keio University (http:/
/www.e-cell.org/ecell)
(Institute for
Advanced Biosciences at Keio University)
(Masaru Tomita). (E-Neuron)
(.: , 4).

,
E-Cell. - ,

. (.: , 5). ,
, . , .
, , E.coli . .

Virtual Cell (VCell), , The National
Resource for Cell Analysis and Modeling (NRCAM). , -

536

 , , , (http://vcell.org). , ,
. , (European
Molecular Biology Laboratory EMBL), (Claude Antony), ,

(Hg J. et al., pp. 349361). .

, .
- (.: , 6-24) ( .. ., 2005, . 232-261;
http://www.mathcell.ru) .

,
. SBML (Systems
Biology Markup Language), , , .
SBML. MathSBML
SBML-.
SBML,
Mathematica, ( .
., 2007).

Blue Brain Project,
2005 .

537

 (EPFL) IBM ( (Henry Markram) EPFL). Blue Brain Project

(NCC).
, NCC, 0,5
2 .
10 . ,
. , ,
.
Blue Brain Project ,
(. 18.1., .: , 26),
-. , , ,
,
, ,

.
, ,
.
Blue Brain
Project Blue Gene IBM 8192 , .
100 ,
,
150 , .

538

 18.1. (BBP/
EPFL, 2007 .).

. 18.2. .
(BBP/EPFL,
2007 .).

539

 2007 . (. 18.2., .: , 27),

(http://bluebrain.epfl.ch).
:
1)
, ;
2)

;
3) ,
.
,

-.
Blue Brain Project ,
, ,
, , . , Blue Brain
Project , , , (http:/
/bluebrain.epfl.ch).
Blue Brain Project IBM . IBM , IBM

540

Almaden Research center, IBM . . .

, IBM , ,
, .
2007 . (James
Frye), (Rajagopal Ananthanarayanan)
. (Dharmendra S. Modha) IBM Almaden Research
center, BlueGene/L 4096
,
8 . ,
6300 (Frye J. et al., 2007).
, Blue Brain Project,

,
2008 . IBM , ,
. IBM , (Cognitive Computing via Synaptronics and
Supercomputing (C2S2).
IBM , ,
(Dharmendra Modha), (IBMs cognitive computing initiative)
. IBM Almaden Research center 2005 .,

2006 . - .


(Kwabena Boahen), .-. (H.-S. Philip Wong)
(Brian Wandell) ,
(Gulio Tononi) , (Rajit Manohar) , (Stefano Fusi) -

541


(Christopher Kello) . IBM
Almaden Research center (Stuart Parkin), (Bulent Kurdi), .
(J. Campbell Scott), (Paul Maglio), (Simone
Raoux), (Rajagopal Ananthanarayanan)
(Raghavendra Singh). IBM . . .
(Chung Lam) (Bipin Rajendran).

.
(Defense Advanced
Research Projects Agency (DARPA)
DARPA (DARPAs Systems of Neuromorphic Adaptive
Plastic Scalable Electronics (SyNAPSE) initiative). DARPA IBM
,
, (.: , 30).
, IBM : 1956 . 512 . ,
, , , .
C2S2 IBM
, 60%, .
, -
.

542

 , IBM
, , , .
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, 40- . , ,
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,
.
2008-09 ., 9
C2S2, , -

544


, . ,
, -

.
IBM
, , , , ,

,
(http://p9.hostingprod.com/@modha.org).
, , , ,
, .
,
, .
, , .
,

. , , , - ,
, , ,
( . ., . ., 2007, . 355).
, , .

545


(
, in silico) .
,
, , ,
,
, .

, , .

, ,
- .

,
.

1. . ., . . - //
. 2007. 2. 2. . 347-360.
2. ., ., ., . . Mathcell // . 2007. . 2. 2. . 361-376.
3. . ., . ., . ., . , .
: // . 2005. . 9. . 232-261.

1. http://www.e-cell.org/ecell
2. http://sciencelinks.jp/j-east/article/200517/000020051705A0633148.php

546

3. http://ecell3d.iab.keio.ac.jp/features.html#section4
4. http://vcell.org
5. Hg J., Schwartz C. Noon A., OToole E., Mastronarde D., McIntosh J.,
Antony C. Organization of Interphase Microtubules in Fission Yeast Analyzed by
Electron Tomography. Developmental Cell. Volume 12. Issue 3. P. 349361.
6. E-cell (http://www.e-cell.org).
7. CyberCell (http://redpoll.pharmacy.ualberta.ca/CCDB).
8. SiliconCell (http://www.bio.vu.nl/hwconf/Silicon/index.html).
9. Systems Biology Research Group (http://gcrg.ucsd.edu).
10. Arkin Laboratory for Dynamical Genomics (http://gobi.lbl.gov).
11. Kitano Symbiotic Systems Project (http://www.symbio.jst.go.jp).
12. VTT Biotechnology (http://sysbio.vtt.fi).
13. Biosystems Group at UCSF (http://biosystems.ucsf.edu).
14. Institute for Systems Biology (http://www.systemsbiology.org).
15. Systems Biology Resource (http://www.systems-biology.org).
16. NRCAM: The National Resource for Cell Analysis and Modeling (http://
www.nrcam.uchc.edu).
17. Plant Systems Biology Resource (http://www.psb.rug.ac.be/).
18. Cellular Networks at Notre Dame (http://www.nd.edu/~networks/cell).
19. BioPathways Consortium (http://www.biopathways.org).
20. SBML: The Systems Biology Markup Language (http://www.sbml.org).
21. CellML (http://www.cellml.org).
22. Cellnomatica (http://www.cellnomica.com).
23. Languages for Systems Biology (http://www.luca.demon.co.uk/
BioComputing.htm).
24. Bio-IT World (http://www.bio-itworld.com/resources/systems).
25. (http://www.mathcell.ru).
26. http://mediatheque.epfl.ch/sv/modules.php?include=view_photo.php&file=
index&name=gallery&op=modload&id=gridNeuron_B3_WM&set_albumName=
Blue_brain_project
27. http://mediatheque.epfl.ch/sv/modules.php?include=view_photo.php&file=
index&name=gallery&op=modload&id=WholeColumn_wm&set_albumName=
Blue_brain_project
28. http://bluebrain.epfl.ch
29. Frye J., Ananthanarayanan R.,Modha D.S. Towards Real-Time, MouseScale Cortical Simulations // Computational and Systems Neuroscience (COSINE).
2007. http://eugen.leitl.org/postbiota/tt/2007-April/000349.html
30. http://www-03.ibm.com/press/us/en/pressrelease/26123.wss
31. http://p9.hostingprod.com/@modha.org

547

 . ., . ., . .,
. ., . ., . .,
. ., . ., . .,
. ., . ., . .

.
.
.

. .
. .
. .

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