Neocort. pyramidal cells subthreshold somatic voltage controls spike propagation (Munro Kopell 2012)

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Accession:136309
There is suggestive evidence that pyramidal cell axons in neocortex may be coupled by gap junctions into an ``axonal plexus" capable of generating Very Fast Oscillations (VFOs) with frequencies exceeding 80 Hz. It is not obvious, however, how a pyramidal cell in such a network could control its output when action potentials are free to propagate from the axons of other pyramidal cells into its own axon. We address this problem by means of simulations based on 3D reconstructions of pyramidal cells from rat somatosensory cortex. We show that somatic depolarization enables propagation via gap junctions into the initial segment and main axon, while somatic hyperpolarization disables it. We show further that somatic voltage cannot effectively control action potential propagation through gap junctions on minor collaterals; action potentials may therefore propagate freely from such collaterals regardless of somatic voltage. In previous work, VFOs are all but abolished during the hyperpolarization phase of slow-oscillations induced by anesthesia in vivo. This finding constrains the density of gap junctions on collaterals in our model and suggests that axonal sprouting due to cortical lesions may result in abnormally high gap junction density on collaterals, leading in turn to excessive VFO activity and hence to epilepsy via kindling.
Reference:
1 . Munro E, Kopell N (2012) Subthreshold somatic voltage in neocortical pyramidal cells can control whether spikes propagate from the axonal plexus to axon terminals: a model study. J Neurophysiol 107:2833-52 [PubMed]
Model Information (Click on a link to find other models with that property)
Model Type: Realistic Network; Neuron or other electrically excitable cell; Axon;
Brain Region(s)/Organism: Neocortex;
Cell Type(s): Neocortex L5/6 pyramidal GLU cell; Neocortex U1 L2/6 pyramidal intratelencephalic GLU cell;
Channel(s): I Na,t; I K; I Sodium; I Potassium;
Gap Junctions: Gap junctions;
Receptor(s):
Gene(s):
Transmitter(s):
Simulation Environment: NEURON; MATLAB;
Model Concept(s): Oscillations; Detailed Neuronal Models; Axonal Action Potentials; Epilepsy;
Implementer(s): Munro, Erin [ecmun at math.bu.edu];
Search NeuronDB for information about:  Neocortex L5/6 pyramidal GLU cell; Neocortex U1 L2/6 pyramidal intratelencephalic GLU cell; I Na,t; I K; I Sodium; I Potassium;
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Munro_Kopell_corticalcontrol
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D3D_g0611P1.out
D3D_g0612P1.out
D3D_g0684P1.out
D3D_g0685P1.out
D3D_g0692P1.out
D3D_g0697P1.out
D3D_g0698P1.out
D3D_g0699P1.out
D3D_g0700P1.out
D3D_g0701P1.out
D3D_g0702P1.out
D3D_g0703P1.out
D3D_g0704P1.out
D3D_g0705P1.out
D3D_g0706P1.out
D3D_g0707P1.out
D3D_g0709P1.out
D3D_g0715_b.out
D3D_g0716_b.out
D3D_g0717_b.out
D3D_g0718_b.out
D3D_g0719_b.out
D3D_g0720_b.out
D3D_g0721_b.out
D3D_g0722_b.out
D3D_g0724P1.out
D3D_g0746_b.out
D3D_g0758_b.out
D3D_g0759_b2.out
D3D_g0760_b.out
D3D_g0761_b.out
D3D_g0762_b.out
D3D_g0763_b.out
D3D_GA01112c2-a.out
D3D_GA01703c2-a.out
D3D_GA01n06c1.out
D3D_GA01o29c1n2-2.out
D3D_GA02209c1n2-1.out
D3D_GA02212c2n3-1.out
D3D_GA02212c2n3-2.out
D3D_GA02219c1n2-1.out
D3D_GA02325c2n3-1.out
D3D_GA02326c1n2-1.out
D3D_GA02402c1n2-1.out
D3D_GA02403c2n3-1.out
D3D_GA02403c3n3-2.out
D3D_GA02531-C4.out
D3D_GA02605-3.out
D3D_gR001P1.out
D3D_gR002P1.out
D3D_gR003P1.out
D3D_I0319_3A.out
D3D_I0319_4A.out
D3D_I0319_5A.out
D3D_I0319_5B.out
D3D_I03192A.out
D3D_I03464.out
D3D_I03471.out
D3D_I03472.out
D3D_I03474.out
D3D_I03481.out
D3D_I03482.out
D3D_I03485.out
D3D_i03535.out
D3D_i03537.out
D3D_I03538.out
D3D_I03539.out
D3D_I03541.out
D3D_I03542.out
D3D_I03544.out
D3D_I03555.out
D3D_i03557.out
D3D_i03558.out
D3D_I03564.out
D3D_I03566.out
D3D_I03569.out
D3D_IF1_190301.out
D3D_IF1_210301.out
D3D_IF2_080501.out
D3D_IF3_040400.out
D3D_IF3_150300.out
D3D_IF3_261199.out
D3D_IF4_140201.out
D3D_JM011004-10-1.out
D3D_JM011004-10-2.out
D3D_JM011304-10-1.out
D3D_JM011504-10.out
D3D_JM011504-10-1.out
D3D_JM012004-30.out
D3D_JM012704-40-1.out
D3D_JM012804-40-2.out
D3D_JM020304-10-1.out
D3D_JM020404-30-1.out
D3D_JM072403-30.out
D3D_JM072803-10.out
D3D_JM072803-20.out
D3D_JM080703-30-1.out
D3D_JM082003-30-1.out
D3D_JM082003-30-2.out
D3D_JM082503-10-1.out
D3D_JM082503-10-2.out
D3D_JM082503-20.out
D3D_JM091503-20-1.out
D3D_JM091503-20-2.out
D3D_JM091503-20-3.out
D3D_JM091703-30-2.out
D3D_JM092303-10.out
D3D_JM092903-10-2.out
D3D_JMcell1or7-2.out
D3D_JMcell7or1-1.out
D3D_JMcell8or11-2.out
D3D_JMn030105-10-1.out
D3D_JMn030105-10-2.out
D3D_JMn030105-20-1.out
D3D_JMn030105-20-2.out
D3D_JMn032205-10-1.out
D3D_JMn091304-10-1.out
D3D_KP060107-2.out
D3D_KP060807p14-s1complete.out
D3D_KP062507complete.out
D3D_M050305SL1.out
D3D_M053005SL1.out
D3D_M060405SL1.out
D3D_MD070401-2.out
D3D_MD070424-4.out
D3D_MD070426-4.out
D3D_MD070456-4.out
D3D_MD070501-1.out
D3D_MD070501-2.out
D3D_MD070503-2-2.out
D3D_MD070517-2.out
D3D_MD070521-1.out
D3D_MD070521-2.out
D3D_MD070620-1.out
D3D_MD070629-1.out
D3D_MD232.out
D3D_MJ021506SL2.out
D3D_MJ030105-2-1.out
D3D_MJ033005-4.out
D3D_MJ042605-1.out
D3D_MJ050305-3.out
D3D_MJ061404-2-1.out
D3D_MJ061404-2-2.out
D3D_MJ061404-3.out
D3D_MJ070504-c1.out
D3D_MJ070504-c2.out
D3D_MJ070604-2.out
D3D_MJ072004-1.out
D3D_MJ080904-1.out
D3D_MJ090804-1-1.out
D3D_MJ090804-1-2.out
D3D_MJ090804-1-3.out
D3D_MJ090804-2-1.out
D3D_MJ090804-2-2.out
D3D_MJ090804-2-3.out
D3D_MJ090804-2-5.out
D3D_RA080304-1.out
D3D_RA080304-3.out
D3D_RosaG282A.out
D3D_RosaY131B.out
D3D_RosaY142A-incomplete.out
D3D_RosaY151B.out
D3D_TS060407s1c1complete.out
D3D_TS070804.out
D3D_Vn01062006-0-1-A.out
D3D_Vn01062006-0-1-B.out
D3D_Vn01132006-0-A.out
D3D_Vn01132006-0-C.out
D3D_Vn01162006-0-A.out
D3D_Vn01162006-0-B.out
D3D_Vn01162006-0-D.out
D3D_Vn01162006-1-C.out
D3D_Vn01232006-0-A.out
D3D_Vn01232006-0-B.out
D3D_Vn01232006-0-C.out
D3D_Vn02042006-0-C.out
D3D_Vn02042006-0-D.out
D3D_Vn02062006-1-B.out
D3D_Vn02062006-1-C.out
D3D_Vn02062006-1-D.out
D3D_Vn02162006-0-D.out
D3D_Vn02222006-0-D.out
D3D_Vn03082006-0-B.out
D3D_Vn03082006-0-C.out
D3D_Vn03082006-0-D.out
D3D_Vn03202006-1-A.out
D3D_Vn03202006-1-D.out
D3D_VN033105-1.out
D3D_VN033105-2.out
D3D_Vn03312005-1.out
D3D_Vn04012005-0-1.out
D3D_Vn04042006-0-A.out
D3D_Vn04042006-0-C.out
D3D_Vn04042006-0-D.out
D3D_VN041305-0-1.out
D3D_VN041305-0-2.out
D3D_VN041305-0-3.out
D3D_Vn04172006-0-D.out
D3D_VN04172006-1unfinished.out
D3D_Vn04182006-0-B.out
D3D_Vn04182006-0-D.out
D3D_Vn04192006-0-D.out
D3D_VN042105-0.out
D3D_Vn04212005-0-0.out
D3D_VN051305-a.out
D3D_VN051305-b.out
D3D_Vn05132005-1.out
D3D_Vn05192006-0-1.out
D3D_Vn05192006-1-B.out
D3D_Vn05302006-0-D.out
D3D_VN06012004-0.out
D3D_Vn06202006-0-B.out
D3D_Vn06202006-0-D.out
D3D_VN062404-1.out
D3D_Vn07132006-0-C.out
D3D_VN07142006-0-C.out
D3D_Vn07142006-1-C.out
D3D_Vn07142006-1-D.out
D3D_Vn07172006-0-B.out
D3D_VN071904.out
D3D_Vn07202006-0-D.out
D3D_Vn11302005-0-A.out
D3D_Vn12052005-0-A.out
D3D_Vn12052005-0-D.out
D3D_Vn12052005-1-A.out
D3D_Vn12052005-1-D.out
D3D_Vn12262005-1-C.out
D3D_Vn12262005-1-D.out
Markram_cells.txt
Markram_L5_cells.txt
rat_layer23.txt
rat_layer23_diams.txt
rat_layer4.txt
rat_layer4_diams.txt
rat_layer5.txt
rat_layer5_diams.txt
Staiger_cells.txt
Svoboda_cells.txt
Yuste_cells.txt
                            
0	0	88.6435	1.27082	-0.113	0.759	0.5	52.64	-69.42	0	-1	2	210	1
1	88.6435	15.4227	0.302481	52.64	-69.42	0	63.29	-80.51	-0.5	0	2	191	2
2	104.066	33.3122	0.25	63.29	-80.51	-0.5	86.3	-104.13	-1	1	2	190	3
3	137.378	31.2838	0.25	86.3	-104.13	-1	104.37	-129.44	-0.5	2	2	185	4
4	168.662	72.4431	0.25	104.37	-129.44	-0.5	144.47	-189.23	-0.5	3	2	178	5
5	241.105	7.25207	0.25	144.47	-189.23	-0.5	149.42	-194.53	-0.5	4	2	131	6
6	248.357	8.68367	0.25	149.42	-194.53	-0.5	154.86	-201.28	-1	5	2	34	7
7	257.041	76.811	0.25	154.86	-201.28	-1	191.49	-267.09	-1	6	2	29	8
8	333.852	38.1564	0.25	191.49	-267.09	-1	220.45	-291.19	0	7	2	28	9
9	372.008	27.8783	0.25	220.45	-291.19	0	231.34	-316.02	-1	8	2	23	10
10	399.887	70.9982	0.25	231.34	-316.02	-1	264.75	-377.25	1.5	9	2	20	11
11	470.885	9.81918	0.25	264.75	-377.25	1.5	270.44	-385.2	1.5	10	2	19	12
12	480.704	58.5358	0.25	270.44	-385.2	1.5	304.6	-428.11	1.5	11	2	14	13
13	539.24	1.74999	0.25	304.6	-428.11	1.5	305.09	-426.43	1.5	12	0	-1	-1
14	539.24	4.6354	0.25	304.6	-428.11	1.5	307.57	-431.49	0.5	12	2	18	15
15	543.875	394.166	0.25	307.57	-431.49	0.5	450.87	-676.16	26	14	2	17	16
16	938.041	252.193	0.25	450.87	-676.16	26	320.93	-743.9	44	15	0	-1	-1
17	938.041	6.33575	0.25	450.87	-676.16	26	448.89	-673.27	23.5	15	0	-1	-1
18	543.875	8.29102	0.25	307.57	-431.49	0.5	304.35	-430.76	0	14	0	-1	-1
19	480.704	1.7596	0.25	270.44	-385.2	1.5	269.95	-386.89	1.5	11	0	-1	-1
20	470.885	9.50873	0.25	264.75	-377.25	1.5	270.69	-381.35	0.5	10	2	22	21
21	480.394	32.2798	0.25	270.69	-381.35	0.5	257.33	-404.25	1	20	0	-1	-1
22	480.394	58.9026	0.25	270.69	-381.35	0.5	266.73	-377.73	-4.5	20	0	-1	-1
23	399.887	76.2434	0.25	231.34	-316.02	-1	210.55	-325.9	11	9	2	25	24
24	476.13	41.2241	0.25	210.55	-325.9	11	212.78	-339.16	18.5	23	0	-1	-1
25	476.13	301.759	0.25	210.55	-325.9	11	365.48	-181.99	13	23	2	27	26
26	777.889	200.746	0.25	365.48	-181.99	13	312.76	-81.23	-7	25	0	-1	-1
27	777.889	33.0258	0.25	365.48	-181.99	13	368.45	-151.14	12.5	25	0	-1	-1
28	372.008	85.2971	0.284295	220.45	-291.19	0	274.4	-324.46	-7	8	0	-1	-1
29	333.852	126.571	0.25	191.49	-267.09	-1	158.57	-337.96	7	7	2	33	30
30	460.422	2.32569	0.25	158.57	-337.96	7	156.35	-338.44	6.5	29	2	32	31
31	462.748	189.214	0.25	156.35	-338.44	6.5	58.09	-356.52	35	30	0	-1	-1
32	462.748	416.682	0.25	156.35	-338.44	6.5	-120.6	-41.94	6.5	30	0	-1	-1
33	460.422	171.529	0.25	158.57	-337.96	7	279.6	-351.46	26	29	0	-1	-1
34	257.041	60.6377	0.25	154.86	-201.28	-1	212.78	-216.71	-2.5	6	2	130	35
35	317.679	68.2127	0.25	212.78	-216.71	-2.5	272.18	-236.47	-4.5	34	2	127	36
36	385.891	47.982	0.25	272.18	-236.47	-4.5	313.75	-257.44	-3	35	2	102	37
37	433.873	89.2846	0.25	313.75	-257.44	-3	389.49	-296.25	-5	36	2	39	38
38	523.158	2.37608	0.25	389.49	-296.25	-5	390.48	-294.09	-5	37	0	-1	-1
39	523.158	29.5005	0.25	389.49	-296.25	-5	414.73	-307.34	-4.5	37	2	97	40
40	552.658	43.0768	0.25	414.73	-307.34	-4.5	439.98	-303.49	-3	39	2	46	41
41	595.735	2.34786	0.25	439.98	-303.49	-3	441.22	-301.56	-2.5	40	2	45	42
42	598.083	21.0637	0.25	441.22	-301.56	-2.5	458.54	-307.1	-3.5	41	2	44	43
43	619.147	1.0017	0.25	458.54	-307.1	-3.5	458.79	-308.07	-3.5	42	0	-1	-1
44	619.147	69.7203	0.25	458.54	-307.1	-3.5	515.71	-307.1	-6.5	42	0	-1	-1
45	598.083	28.5333	0.25	441.22	-301.56	-2.5	462.75	-294.09	-2	41	0	-1	-1
46	595.735	58.5521	0.25	439.98	-303.49	-3	457.8	-278.18	4	40	2	48	47
47	654.287	63.2451	0.25	457.8	-278.18	4	504.58	-287.09	7	46	0	-1	-1
48	654.287	64.5701	0.25	457.8	-278.18	4	485.77	-257.69	9.5	46	2	96	49
49	718.857	72.9527	0.25	485.77	-257.69	9.5	544.18	-224.42	10.5	48	2	51	50
50	791.81	8.94812	0.25	544.18	-224.42	10.5	546.65	-231.65	10	49	0	-1	-1
51	791.81	447.353	0.25	544.18	-224.42	10.5	836.47	34.96	1.5	49	2	57	52
52	1239.16	18.4593	0.25	836.47	34.96	1.5	849.34	44.12	-0.5	51	2	54	53
53	1257.62	2.13159	0.25	849.34	44.12	-0.5	850.83	45.56	-1	52	0	-1	-1
54	1257.62	27.4205	0.25	849.34	44.12	-0.5	869.14	56.89	-2	52	2	56	55
55	1285.04	1.45	0.25	869.14	56.89	-2	869.14	58.34	-2	54	0	-1	-1
56	1285.04	107.343	0.25	869.14	56.89	-2	900.57	87.75	-9.5	54	0	-1	-1
57	1239.16	49.2757	0.25	836.47	34.96	1.5	834.99	78.35	-1.5	51	2	59	58
58	1288.44	2.16535	0.25	834.99	78.35	-1.5	836.97	77.63	-1	57	0	-1	-1
59	1288.44	42.5323	0.25	834.99	78.35	-1.5	823.85	118.85	-2.5	57	2	63	60
60	1330.97	54.1713	0.25	823.85	118.85	-2.5	815.19	164.16	-5.5	59	2	62	61
61	1385.14	9.24032	0.25	815.19	164.16	-5.5	816.18	155.49	-6	60	0	-1	-1
62	1385.14	89.0045	0.25	815.19	164.16	-5.5	808.26	243.47	-9	60	0	-1	-1
63	1330.97	170.302	0.25	823.85	118.85	-2.5	716.93	243.95	2	59	2	91	64
64	1501.27	7.59627	0.25	716.93	243.95	2	711.73	248.78	2.5	63	2	82	65
65	1508.87	29.806	0.25	711.73	248.78	2.5	689.21	267.34	1.5	64	2	67	66
66	1538.68	0.882092	0.25	689.21	267.34	1.5	688.47	266.86	1.5	65	0	-1	-1
67	1538.68	8.91216	0.25	689.21	267.34	1.5	682.28	272.88	1	65	2	69	68
68	1547.59	0.707107	0.25	682.28	272.88	1	681.78	272.88	1.5	67	0	-1	-1
69	1547.59	65.3779	0.25	682.28	272.88	1	639.21	319.41	-0.5	67	2	81	70
70	1612.97	84.6515	0.25	639.21	319.41	-0.5	585.76	382.08	-2.5	69	2	76	71
71	1697.62	65.8254	0.25	585.76	382.08	-2.5	549.37	432.7	-3.5	70	2	73	72
72	1763.44	9.54053	0.25	549.37	432.7	-3.5	551.11	431.98	-5.5	71	0	-1	-1
73	1763.44	29.8475	0.25	549.37	432.7	-3.5	531.8	454.88	-4.5	71	2	75	74
74	1793.29	25.6552	0.25	531.8	454.88	-4.5	524.13	449.82	-5.5	73	0	-1	-1
75	1793.29	11.5183	0.25	531.8	454.88	-4.5	526.6	464.76	-5	73	0	-1	-1
76	1697.62	8.70935	0.25	585.76	382.08	-2.5	583.53	389.55	-3.5	70	2	78	77
77	1706.33	9.27963	0.25	583.53	389.55	-3.5	583.03	386.42	-3	76	0	-1	-1
78	1706.33	23.4098	0.25	583.53	389.55	-3.5	575.36	405.7	-4.5	76	2	80	79
79	1729.74	25.8789	0.25	575.36	405.7	-4.5	574.87	428.12	-6	78	0	-1	-1
80	1729.74	33.6005	0.25	575.36	405.7	-4.5	559.52	420.65	-6	78	0	-1	-1
81	1612.97	121.515	0.25	639.21	319.41	-0.5	577.09	373.16	-6.5	69	0	-1	-1
82	1508.87	95.6344	0.25	711.73	248.78	2.5	657.53	324.71	5.5	64	2	90	83
83	1604.5	9.04962	0.25	657.53	324.71	5.5	653.57	332.66	6	82	2	89	84
84	1613.55	145.839	0.25	653.57	332.66	6	558.04	438.49	12	83	2	88	85
85	1759.39	151.391	0.25	558.04	438.49	12	476.36	476.58	31	84	2	87	86
86	1910.78	34.6937	0.25	476.36	476.58	31	447.16	470.79	31.5	85	0	-1	-1
87	1910.78	100.373	0.25	476.36	476.58	31	463.99	433.19	43.5	85	0	-1	-1
88	1759.39	66.2769	0.25	558.04	438.49	12	504.58	474.89	12	84	0	-1	-1
89	1613.55	27.8983	0.25	653.57	332.66	6	663.47	354.12	5	83	0	-1	-1
90	1604.5	152.667	0.25	657.53	324.71	5.5	647.13	455.36	15	82	0	-1	-1
91	1501.27	8.27153	0.25	716.93	243.95	2	718.91	250.22	3.5	63	2	95	92
92	1509.55	2.79645	0.25	718.91	250.22	3.5	719.65	252.87	4	91	2	94	93
93	1512.34	277.236	0.25	719.65	252.87	4	574.87	391.24	33.5	92	0	-1	-1
94	1512.34	293.682	0.25	719.65	252.87	4	596.64	285.66	45	92	0	-1	-1
95	1509.55	40.7745	0.25	718.91	250.22	3.5	718.66	246.37	14.5	91	0	-1	-1
96	718.857	257.088	0.25	485.77	-257.69	9.5	499.38	-153.55	46	48	0	-1	-1
97	552.658	68.5899	0.25	414.73	-307.34	-4.5	411.02	-364.96	-2	39	2	99	98
98	621.248	25.3094	0.25	411.02	-364.96	-2	431.32	-373.39	-3.5	97	0	-1	-1
99	621.248	34.4384	0.25	411.02	-364.96	-2	404.59	-391.23	1	97	2	101	100
100	655.687	4.99598	0.25	404.59	-391.23	1	403.6	-395.81	1	99	0	-1	-1
101	655.687	140.471	0.25	404.59	-391.23	1	434.78	-500.19	10	99	0	-1	-1
102	433.873	43.5591	0.25	313.75	-257.44	-3	343.45	-239.85	-2.5	36	2	104	103
103	477.432	5.13823	0.25	343.45	-239.85	-2.5	342.71	-234.79	-2	102	0	-1	-1
104	477.432	12.9333	0.25	343.45	-239.85	-2.5	354.59	-240.33	-1.5	102	2	106	105
105	490.366	145.408	0.25	354.59	-240.33	-1.5	391.72	-357.72	4	104	0	-1	-1
106	490.366	221.188	0.25	354.59	-240.33	-1.5	527.59	-179.58	8	104	2	108	107
107	711.554	52.9413	0.25	527.59	-179.58	8	542.2	-219.12	12.5	106	0	-1	-1
108	711.554	80.2146	0.25	527.59	-179.58	8	596.65	-160.3	7.5	106	2	124	109
109	791.769	538.307	0.25	596.65	-160.3	7.5	887.21	42.19	9	108	2	117	110
110	1330.08	65.9542	0.25	887.21	42.19	9	874.83	105.59	12	109	2	114	111
111	1396.03	117.725	0.25	874.83	105.59	12	859.98	214.3	10.5	110	2	113	112
112	1513.75	3.29955	0.25	859.98	214.3	10.5	863.2	213.58	10.5	111	0	-1	-1
113	1513.75	280.195	0.25	859.98	214.3	10.5	826.82	440.18	-8.5	111	0	-1	-1
114	1396.03	237.537	0.25	874.83	105.59	12	735.24	265.41	21	110	2	116	115
115	1633.57	20.3048	0.25	735.24	265.41	21	728.07	282.04	25.5	114	0	-1	-1
116	1633.57	78.7157	0.25	735.24	265.41	21	778.31	289.51	9	114	0	-1	-1
117	1330.08	102.807	0.25	887.21	42.19	9	824.84	87.51	3	109	2	123	118
118	1432.88	78.6888	0.25	824.84	87.51	3	775.09	136.44	-1	117	2	122	119
119	1511.57	270.771	0.25	775.09	136.44	-1	670.4	376.54	-2	118	2	121	120
120	1782.34	84.6535	0.25	670.4	376.54	-2	660.25	447.65	1.5	119	0	-1	-1
121	1782.34	46.2181	0.25	670.4	376.54	-2	665.45	420.65	-0.5	119	0	-1	-1
122	1511.57	198.166	0.25	775.09	136.44	-1	686.98	289.27	-8	118	0	-1	-1
123	1432.88	47.805	0.25	824.84	87.51	3	827.31	54.48	-1	117	0	-1	-1
124	791.769	70.4634	0.25	596.65	-160.3	7.5	591.45	-113.05	0.5	108	2	126	125
125	862.232	3.00857	0.25	591.45	-113.05	0.5	588.48	-112.57	0.5	124	0	-1	-1
126	862.232	85.422	0.25	591.45	-113.05	0.5	587.74	-72.31	-6	124	0	-1	-1
127	385.891	24.0565	0.25	272.18	-236.47	-4.5	286.28	-233.34	-6	35	2	129	128
128	409.948	10.3475	0.25	286.28	-233.34	-6	284.55	-240.09	-7	127	0	-1	-1
129	409.948	23.1547	0.25	286.28	-233.34	-6	299.4	-233.82	-7.5	127	0	-1	-1
130	317.679	101.15	0.25	212.78	-216.71	-2.5	232.08	-143.18	-8	34	0	-1	-1
131	248.357	167.507	0.25	149.42	-194.53	-0.5	125.66	-319.64	10.5	5	2	163	132
132	415.865	29.1425	0.25	125.66	-319.64	10.5	125.16	-344.95	12.5	131	2	162	133
133	445.007	7.02932	0.25	125.16	-344.95	12.5	127.39	-351.46	12	132	2	153	134
134	452.036	53.2305	0.25	127.39	-351.46	12	162.29	-390.03	13	133	2	136	135
135	505.267	2.31203	0.25	162.29	-390.03	13	164.02	-388.58	13.5	134	0	-1	-1
136	505.267	148.186	0.25	162.29	-390.03	13	281.83	-387.86	5.5	134	2	152	137
137	653.453	36.2675	0.25	281.83	-387.86	5.5	311.28	-378.94	5	136	2	139	138
138	689.721	3.64907	0.25	311.28	-378.94	5	310.78	-375.56	5	137	0	-1	-1
139	689.721	53.3919	0.25	311.28	-378.94	5	347.41	-369.3	0.5	137	2	141	140
140	743.113	1.75285	0.25	347.41	-369.3	0.5	347.91	-370.98	0.5	139	0	-1	-1
141	743.113	18.6707	0.25	347.41	-369.3	0.5	364.74	-367.61	-0.5	139	2	151	142
142	761.783	40.1651	0.25	364.74	-367.61	-0.5	393.45	-360.86	-2	141	2	150	143
143	801.948	66.1979	0.25	393.45	-360.86	-2	449.88	-346.39	-4	142	2	145	144
144	868.146	18.1992	0.25	449.88	-346.39	-4	461.76	-344.95	-3	143	0	-1	-1
145	868.146	37.6165	0.25	449.88	-346.39	-4	476.61	-336.99	-5.5	143	2	147	146
146	905.763	2.29272	0.25	476.61	-336.99	-5.5	475.87	-339.16	-5.5	145	0	-1	-1
147	905.763	6.2913	0.25	476.61	-336.99	-5.5	482.05	-334.82	-5.5	145	2	149	148
148	912.054	2.473	0.25	482.05	-334.82	-5.5	481.81	-332.41	-6	147	0	-1	-1
149	912.054	128.189	0.25	482.05	-334.82	-5.5	563.23	-271.43	-6.5	147	0	-1	-1
150	801.948	81.1878	0.25	393.45	-360.86	-2	406.81	-299.15	-7	142	0	-1	-1
151	761.783	64.2956	0.25	364.74	-367.61	-0.5	399.64	-361.58	4	141	0	-1	-1
152	653.453	138.651	0.25	281.83	-387.86	5.5	328.6	-358.93	-8	136	0	-1	-1
153	452.036	92.3419	0.25	127.39	-351.46	12	150.41	-428.11	10.5	133	2	155	154
154	544.378	179.347	0.25	150.41	-428.11	10.5	206.59	-475.12	-8	153	0	-1	-1
155	544.378	1.0017	0.25	150.41	-428.11	10.5	150.16	-429.08	10.5	153	2	157	156
156	545.38	45.4094	0.25	150.16	-429.08	10.5	114.02	-412.69	5	155	0	-1	-1
157	545.38	12.9175	0.25	150.16	-429.08	10.5	158.08	-430.52	12.5	155	2	159	158
158	558.298	9.50827	0.25	158.08	-430.52	12.5	165.01	-425.7	12.5	157	0	-1	-1
159	558.298	23.2155	0.25	158.08	-430.52	12.5	176.15	-436.31	14	157	2	161	160
160	581.513	192.43	0.25	176.15	-436.31	14	282.57	-563.83	18.5	159	0	-1	-1
161	581.513	11.7177	0.25	176.15	-436.31	14	174.66	-440.89	15.5	159	0	-1	-1
162	445.007	2.87124	0.25	125.16	-344.95	12.5	123.18	-346.39	14	132	0	-1	-1
163	415.865	254.225	0.25	125.66	-319.64	10.5	334.54	-250.94	18.5	131	2	177	164
164	670.09	45.984	0.25	334.54	-250.94	18.5	367.96	-226.35	17.5	163	2	176	165
165	716.074	79.1997	0.25	367.96	-226.35	17.5	426.86	-192.6	18.5	164	2	167	166
166	795.274	3.17932	0.25	426.86	-192.6	18.5	429.34	-194.53	18.5	165	0	-1	-1
167	795.274	39.9421	0.25	426.86	-192.6	18.5	450.62	-165.12	20	165	2	175	168
168	835.216	124.346	0.25	450.62	-165.12	20	538.48	-88.95	19	167	2	170	169
169	959.562	2.88195	0.25	538.48	-88.95	19	537.49	-86.29	19.5	168	0	-1	-1
170	959.562	297.163	0.25	538.48	-88.95	19	781.28	-48.21	2	168	2	172	171
171	1256.73	2.51618	0.25	781.28	-48.21	2	783.75	-48.69	2	170	0	-1	-1
172	1256.73	75.5084	0.25	781.28	-48.21	2	823.35	0	4	170	2	174	173
173	1332.23	11.735	0.25	823.35	0	4	830.28	5.55	4.5	172	0	-1	-1
174	1332.23	26.3825	0.25	823.35	0	4	841.42	15.43	2	172	0	-1	-1
175	835.216	33.9221	0.25	450.62	-165.12	20	479.83	-166.32	24.5	167	0	-1	-1
176	716.074	141.94	0.25	367.96	-226.35	17.5	429.58	-188.74	2	164	0	-1	-1
177	670.09	164.94	0.25	334.54	-250.94	18.5	390.73	-127.27	24.5	163	0	-1	-1
178	241.105	336.763	0.25	144.47	-189.23	-0.5	-144.61	-107.51	18	4	2	180	179
179	577.868	1.94413	0.25	-144.61	-107.51	18	-145.6	-108.95	18	178	0	-1	-1
180	577.868	205.31	0.25	-144.61	-107.51	18	-207.48	74.97	12.5	178	2	182	181
181	783.178	42.8694	0.25	-207.48	74.97	12.5	-210.69	68.71	6.5	180	0	-1	-1
182	783.178	45.2832	0.25	-207.48	74.97	12.5	-219.11	105.11	9	180	2	184	183
183	828.461	10.1962	0.25	-219.11	105.11	9	-221.34	112.58	9	182	0	-1	-1
184	828.461	28.7531	0.25	-219.11	105.11	9	-212.92	130.42	8.5	182	0	-1	-1
185	168.662	232.793	0.25	104.37	-129.44	-0.5	167.48	80.04	-2	3	2	187	186
186	401.455	3.11157	0.25	167.48	80.04	-2	169.71	77.87	-2	185	0	-1	-1
187	401.455	140.011	0.25	167.48	80.04	-2	195.45	202.98	-3	185	2	189	188
188	541.467	17.8944	0.25	195.45	202.98	-3	201.14	212.62	-4.5	187	0	-1	-1
189	541.467	98.0104	0.25	195.45	202.98	-3	240.25	273.12	-1.5	187	0	-1	-1
190	137.378	112.033	0.25	86.3	-104.13	-1	88.53	-83.64	10.5	2	0	-1	-1
191	104.066	9.26197	0.25	63.29	-80.51	-0.5	63.29	-79.3	-1.5	1	2	197	192
192	113.328	13.3264	0.25	63.29	-79.3	-1.5	64.03	-76.41	-2.5	191	2	194	193
193	126.654	29.69	0.25	64.03	-76.41	-2.5	61.31	-67.73	-6	192	0	-1	-1
194	126.654	170.765	0.25	64.03	-76.41	-2.5	137.04	42.19	-5	192	2	196	195
195	297.42	11.6869	0.25	137.04	42.19	-5	145.46	48.22	-5.5	194	0	-1	-1
196	297.42	24.7884	0.25	137.04	42.19	-5	147.68	54.48	-6	194	0	-1	-1
197	113.328	9.81476	0.25	63.29	-79.3	-1.5	54.13	-79.3	-1.5	191	2	203	198
198	123.143	187.328	0.25	54.13	-79.3	-1.5	84.82	-260.1	-5	197	2	200	199
199	310.471	3.43984	0.25	84.82	-260.1	-5	88.04	-258.89	-5	198	0	-1	-1
200	310.471	38.4409	0.25	84.82	-260.1	-5	90.51	-297.46	-5	198	2	202	201
201	348.911	3.70369	0.25	90.51	-297.46	-5	88.53	-300.59	-5	200	0	-1	-1
202	348.911	5.85226	0.25	90.51	-297.46	-5	93.23	-301.8	-5	200	0	-1	-1
203	123.143	42.1127	0.25	54.13	-79.3	-1.5	17.99	-61.46	-0.5	197	2	209	204
204	165.256	164.38	0.25	17.99	-61.46	-0.5	-119.12	20.74	3	203	2	206	205
205	329.635	137.12	0.25	-119.12	20.74	3	-117.63	81.24	15.5	204	0	-1	-1
206	329.635	51.3434	0.25	-119.12	20.74	3	-156.99	38.82	0.5	204	2	208	207
207	380.979	71.4141	0.25	-156.99	38.82	0.5	-196.83	72.56	-2	206	0	-1	-1
208	380.979	80.527	0.25	-156.99	38.82	0.5	-203.52	98.12	5	206	0	-1	-1
209	165.256	54.94	0.25	17.99	-61.46	-0.5	12.3	-42.9	-4.5	203	0	-1	-1
210	88.6435	6.14709	0.49	52.64	-69.42	0	48.19	-71.59	-0.5	0	2	250	211
211	94.7905	4.77263	0.299246	48.19	-71.59	-0.5	44.48	-72.79	-0.5	210	2	247	212
212	99.5632	20.014	0.25	44.48	-72.79	-0.5	29.63	-62.43	1.5	211	2	242	213
213	119.577	22.544	0.25	29.63	-62.43	1.5	10.07	-53.03	3	212	2	241	214
214	142.121	34.0831	0.25	10.07	-53.03	3	-18.88	-41.22	5	213	2	238	215
215	176.204	104.377	0.25	-18.88	-41.22	5	-106	-2.16	10	214	2	217	216
216	280.581	69.7166	0.25	-106	-2.16	10	-161.94	26.04	6	215	0	-1	-1
217	280.581	37.2454	0.25	-106	-2.16	10	-127.53	-7.71	13	215	2	223	218
218	317.827	13.0271	0.25	-127.53	-7.71	13	-136.2	1.45	13.5	217	2	220	219
219	330.854	56.2561	0.25	-136.2	1.45	13.5	-141.64	53.04	13	218	0	-1	-1
220	330.854	120.103	0.25	-136.2	1.45	13.5	-180	88.23	16	218	2	222	221
221	450.957	32.5705	0.25	-180	88.23	16	-173.32	116.68	16	220	0	-1	-1
222	450.957	314.059	0.25	-180	88.23	16	-38.68	67.5	45	220	0	-1	-1
223	317.827	126.529	0.25	-127.53	-7.71	13	-75.06	-111.6	13	217	2	237	224
224	444.356	3.77876	0.25	-75.06	-111.6	13	-71.85	-113.53	13.5	223	2	236	225
225	448.134	169.66	0.25	-71.85	-113.53	13.5	10.32	-259.37	12	224	2	235	226
226	617.795	71.893	0.25	10.32	-259.37	12	34.58	-324.94	9.5	225	2	228	227
227	689.688	195.243	0.25	34.58	-324.94	9.5	131.35	-478.01	7.5	226	0	-1	-1
228	689.688	80.0737	0.25	34.58	-324.94	9.5	51.9	-366.16	0.5	226	2	232	229
229	769.761	8.87694	0.25	51.9	-366.16	0.5	51.16	-371.95	0.5	228	2	231	230
230	778.638	5.23403	0.25	51.16	-371.95	0.5	52.4	-376.53	0	229	0	-1	-1
231	778.638	74.8564	0.25	51.16	-371.95	0.5	55.86	-410.52	-4.5	229	0	-1	-1
232	769.761	86.8381	0.25	51.9	-366.16	0.5	113.28	-363.75	-5	228	2	234	233
233	856.6	13.3911	0.25	113.28	-363.75	-5	120.46	-353.87	-4	232	0	-1	-1
234	856.6	26.6321	0.25	113.28	-363.75	-5	129.37	-360.86	-5.5	232	0	-1	-1
235	617.795	302.607	0.25	10.32	-259.37	12	-69.37	-512	18.5	225	0	-1	-1
236	448.134	67.2639	0.25	-71.85	-113.53	13.5	-41.16	-75.45	15	224	0	-1	-1
237	444.356	128.621	0.25	-75.06	-111.6	13	-113.67	-221.53	17.5	223	0	-1	-1
238	176.204	106.966	0.25	-18.88	-41.22	5	-81.25	-112.81	8	214	2	240	239
239	283.171	7.64028	0.25	-81.25	-112.81	8	-78.78	-120.04	8	238	0	-1	-1
240	283.171	133.371	0.25	-81.25	-112.81	8	-122.83	-202.48	21.5	238	0	-1	-1
241	142.121	47.0411	0.25	10.07	-53.03	3	19.23	-94.97	9	213	0	-1	-1
242	119.577	126.082	0.25	29.63	-62.43	1.5	-67.39	10.37	0.5	212	2	244	243
243	245.659	6.7223	0.25	-67.39	10.37	0.5	-67.39	11.09	0	242	0	-1	-1
244	245.659	132.359	0.25	-67.39	10.37	0.5	-171.09	56.17	-0.5	242	2	246	245
245	378.018	47.4056	0.25	-171.09	56.17	-0.5	-193.12	69.19	-3	244	0	-1	-1
246	378.018	74.7725	0.25	-171.09	56.17	-0.5	-229.75	89.92	1	244	0	-1	-1
247	99.5632	108.244	0.25	44.48	-72.79	-0.5	84.82	5.07	4	211	2	249	248
248	207.807	235.353	0.25	84.82	5.07	4	60.56	214.06	-4	247	0	-1	-1
249	207.807	18.0737	0.25	84.82	5.07	4	89.27	11.09	13	247	0	-1	-1
250	94.7905	93.4352	0.269768	48.19	-71.59	-0.5	16.01	12.54	-2.5	210	2	252	251
251	188.226	106.747	0.25	16.01	12.54	-2.5	-18.64	85.58	-3.5	250	0	-1	-1
252	188.226	87.9241	0.25	16.01	12.54	-2.5	67.25	67.02	-4	250	2	254	253
253	276.15	42.182	0.25	67.25	67.02	-4	44.48	99.56	-4.5	252	0	-1	-1
254	276.15	60.3537	0.25	67.25	67.02	-4	111.8	85.82	-5	252	2	256	255
255	336.504	2.79645	0.25	111.8	85.82	-5	112.54	88.47	-4.5	254	0	-1	-1
256	336.504	18.3768	0.25	111.8	85.82	-5	122.44	97.87	-5	254	0	-1	-1

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