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_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	63.8367	1.38106	-1.338	0.16	1	1.21	-52.13	8.5	-1	2	2	1
1	63.8367	1.37902	0.49	1.21	-52.13	8.5	0.22	-53.09	8.5	0	0	-1	-1
2	63.8367	25.2712	0.49	1.21	-52.13	8.5	1.96	-75.03	10.5	0	2	162	3
3	89.108	12.3691	0.311474	1.96	-75.03	10.5	7.15	-74.06	4.5	2	2	75	4
4	101.477	2.62118	0.25	7.15	-74.06	4.5	7.4	-71.65	5.5	3	2	30	5
5	104.098	14.0949	0.25	7.4	-71.65	5.5	10.37	-60.32	4.5	4	2	7	6
6	118.193	38.205	0.25	10.37	-60.32	4.5	7.9	-25.61	1.5	5	0	-1	-1
7	118.193	14.9931	0.25	10.37	-60.32	4.5	21.51	-64.42	3	5	2	11	8
8	133.186	21.4419	0.25	21.51	-64.42	3	16.56	-69.48	-0.5	7	2	10	9
9	154.628	31.7183	0.25	16.56	-69.48	-0.5	12.35	-89.49	-3.5	8	0	-1	-1
10	154.628	32.5075	0.25	16.56	-69.48	-0.5	-5.22	-65.39	-3.5	8	0	-1	-1
11	133.186	18.3502	0.25	21.51	-64.42	3	35.37	-68.52	2.5	7	2	21	12
12	151.536	15.8295	0.25	35.37	-68.52	2.5	37.6	-54.06	1	11	2	20	13
13	167.366	85.4172	0.25	37.6	-54.06	1	82.15	5.48	3	12	2	19	14
14	252.783	6.56909	0.25	82.15	5.48	3	86.85	9.82	3.5	13	2	18	15
15	259.352	10.7078	0.25	86.85	9.82	3.5	89.57	19.47	2.5	14	2	17	16
16	270.06	34.4188	0.25	89.57	19.47	2.5	77.2	45.98	-0.5	15	0	-1	-1
17	270.06	21.9342	0.25	89.57	19.47	2.5	90.31	35.38	2	15	0	-1	-1
18	259.352	2.45416	0.25	86.85	9.82	3.5	84.87	11.27	3.5	14	0	-1	-1
19	252.783	1.79535	0.25	82.15	5.48	3	83.88	5	3	13	0	-1	-1
20	167.366	0.746726	0.25	37.6	-54.06	1	37.1	-54.3	1.5	12	0	-1	-1
21	151.536	44.6238	0.25	35.37	-68.52	2.5	65.56	-88.29	-1	11	2	29	22
22	196.16	99.3827	0.25	65.56	-88.29	-1	137.83	-125.17	-3	21	2	28	23
23	295.543	17.0811	0.25	137.83	-125.17	-3	150.46	-125.17	-3	22	2	27	24
24	312.624	23.2314	0.25	150.46	-125.17	-3	169.27	-130.95	-3.5	23	2	26	25
25	335.855	14.685	0.25	169.27	-130.95	-3.5	181.64	-133.85	-3	24	0	-1	-1
26	335.855	2.57562	0.25	169.27	-130.95	-3.5	171.74	-131.68	-3.5	24	0	-1	-1
27	312.624	2.57015	0.25	150.46	-125.17	-3	151.2	-122.76	-3.5	23	0	-1	-1
28	295.543	4.30666	0.25	137.83	-125.17	-3	141.55	-127.34	-3	22	0	-1	-1
29	196.16	3.00853	0.25	65.56	-88.29	-1	62.59	-88.77	-1	21	0	-1	-1
30	104.098	36.0576	0.25	7.4	-71.65	5.5	-5.72	-41.76	6.5	4	2	74	31
31	140.156	25.6024	0.25	-5.72	-41.76	6.5	-14.63	-18.38	7.5	30	2	59	32
32	165.758	69.7432	0.25	-14.63	-18.38	7.5	-48.53	-9.7	15.5	31	2	58	33
33	235.501	19.7679	0.25	-48.53	-9.7	15.5	-59.67	-8.01	17.5	32	2	39	34
34	255.269	50.983	0.25	-59.67	-8.01	17.5	-71.3	37.55	20.5	33	2	38	35
35	306.252	6.20534	0.25	-71.3	37.55	20.5	-74.03	43.09	20	34	2	37	36
36	312.458	84.9697	0.25	-74.03	43.09	20	-129.22	99.5	22	35	0	-1	-1
37	312.458	2.09611	0.25	-74.03	43.09	20	-72.79	44.78	20	35	0	-1	-1
38	306.252	2.4993	0.25	-71.3	37.55	20.5	-72.54	35.38	20.5	34	0	-1	-1
39	255.269	23.2451	0.25	-59.67	-8.01	17.5	-76.25	-12.11	20.5	33	2	41	40
40	278.514	188.728	0.25	-76.25	-12.11	20.5	-46.06	-0.06	42.5	39	0	-1	-1
41	278.514	12.1168	0.25	-76.25	-12.11	20.5	-85.16	-18.38	20	39	2	57	42
42	290.631	28.5181	0.25	-85.16	-18.38	20	-104.47	-37.66	21.5	41	2	56	43
43	319.149	144.239	0.25	-104.47	-37.66	21.5	-204.46	-128.54	17	42	2	55	44
44	463.389	15.331	0.25	-204.46	-128.54	17	-217.57	-132.4	17.5	43	2	54	45
45	478.72	9.35519	0.25	-217.57	-132.4	17.5	-224.26	-135.77	16.5	44	2	53	46
46	488.075	4.21975	0.25	-224.26	-135.77	16.5	-224.26	-139.63	15.5	45	2	52	47
47	492.295	119.067	0.25	-224.26	-139.63	15.5	-316.33	-145.18	8.5	46	2	51	48
48	611.362	30.0509	0.25	-316.33	-145.18	8.5	-340.33	-145.18	6.5	47	2	50	49
49	641.413	15.4255	0.25	-340.33	-145.18	6.5	-354.19	-144.69	5.5	48	0	-1	-1
50	641.413	10.4657	0.25	-340.33	-145.18	6.5	-349	-148.79	4.5	48	0	-1	-1
51	611.362	2.99827	0.25	-316.33	-145.18	8.5	-314.59	-144.21	8.5	47	0	-1	-1
52	492.295	14.3311	0.25	-224.26	-139.63	15.5	-231.19	-150.96	15	46	0	-1	-1
53	488.075	5.79679	0.25	-224.26	-135.77	16.5	-222.52	-139.87	19	45	0	-1	-1
54	478.72	5.33093	0.25	-217.57	-132.4	17.5	-217.57	-129.03	18.5	44	0	-1	-1
55	463.389	2.08213	0.25	-204.46	-128.54	17	-205.69	-126.86	17	43	0	-1	-1
56	319.149	6.55353	0.25	-104.47	-37.66	21.5	-103.97	-31.64	21.5	42	0	-1	-1
57	290.631	3.48698	0.25	-85.16	-18.38	20	-82.69	-20.79	20.5	41	0	-1	-1
58	235.501	2.79644	0.25	-48.53	-9.7	15.5	-47.79	-12.35	15	32	0	-1	-1
59	165.758	36.0492	0.25	-14.63	-18.38	7.5	-21.31	15.61	9	31	2	61	60
60	201.807	153.598	0.25	-21.31	15.61	9	36.85	93.23	27.5	59	0	-1	-1
61	201.807	6.54772	0.25	-21.31	15.61	9	-22.05	21.88	7.5	59	2	73	62
62	208.355	75.2585	0.25	-22.05	21.88	7.5	-35.17	94.19	5.5	61	2	72	63
63	283.614	70.5349	0.25	-35.17	94.19	5.5	-4.48	151.32	5.5	62	2	71	64
64	354.148	14.9135	0.25	-4.48	151.32	5.5	4.68	161.21	6.5	63	2	70	65
65	369.062	84.0126	0.25	4.68	161.21	6.5	65.56	186.76	3	64	2	69	66
66	453.075	71.0665	0.25	65.56	186.76	3	121.25	211.83	3	65	2	68	67
67	524.141	19.0042	0.25	121.25	211.83	3	136.84	215.69	2.5	66	0	-1	-1
68	524.141	2.65001	0.25	121.25	211.83	3	121.25	209.18	3	66	0	-1	-1
69	453.075	14.455	0.25	65.56	186.76	3	74.23	198.09	2.5	65	0	-1	-1
70	369.062	2.34659	0.25	4.68	161.21	6.5	3.94	163.38	7	64	0	-1	-1
71	354.148	3.11801	0.25	-4.48	151.32	5.5	-1.76	149.88	5	63	0	-1	-1
72	283.614	6.54294	0.25	-35.17	94.19	5.5	-36.16	99.26	6	62	0	-1	-1
73	208.355	2.11029	0.25	-22.05	21.88	7.5	-24.03	21.15	7.5	61	0	-1	-1
74	140.156	3.44866	0.25	-5.72	-41.76	6.5	-2.75	-40.08	6	30	0	-1	-1
75	101.477	5.10685	0.25	7.15	-74.06	4.5	9.13	-74.31	6	3	2	87	76
76	106.584	66.3343	0.25	9.13	-74.31	6	16.56	-11.15	7.5	75	2	80	77
77	172.918	25.4002	0.25	16.56	-11.15	7.5	27.2	10.79	7	76	2	79	78
78	198.318	9.89303	0.25	27.2	10.79	7	23.74	18.98	3.5	77	0	-1	-1
79	198.318	80.6705	0.25	27.2	10.79	7	77.2	27.18	6	77	0	-1	-1
80	172.918	30.451	0.25	16.56	-11.15	7.5	11.86	18.02	7.5	76	2	86	81
81	203.369	53.119	0.25	11.86	18.02	7.5	10.12	70.09	7	80	2	85	82
82	256.488	96.9734	0.25	10.12	70.09	7	-27	156.15	8	81	2	84	83
83	353.462	11.3815	0.25	-27	156.15	8	-30.96	166.75	8.5	82	0	-1	-1
84	353.462	2.33967	0.25	-27	156.15	8	-25.76	158.07	7.5	82	0	-1	-1
85	256.488	252.298	0.25	10.12	70.09	7	108.88	276.92	19	81	0	-1	-1
86	203.369	1.76409	0.25	11.86	18.02	7.5	13.34	18.98	7.5	80	0	-1	-1
87	106.584	12.2079	0.25	9.13	-74.31	6	15.32	-71.89	7	75	2	109	88
88	118.792	18.5818	0.25	15.32	-71.89	7	15.82	-54.54	9	87	2	104	89
89	137.374	24.9178	0.25	15.82	-54.54	9	21.51	-31.88	12	88	2	101	90
90	162.291	70.1541	0.25	21.51	-31.88	12	8.39	34.65	17	89	2	100	91
91	232.446	25.6761	0.25	8.39	34.65	17	11.11	58.28	17.5	90	2	93	92
92	258.122	321.536	0.25	11.11	58.28	17.5	132.39	76.11	44	91	0	-1	-1
93	258.122	124.636	0.25	11.11	58.28	17.5	-32.94	152.77	25.5	91	2	99	94
94	382.758	44.0486	0.25	-32.94	152.77	25.5	-52.74	185.55	27.5	93	2	98	95
95	426.806	19.7915	0.25	-52.74	185.55	27.5	-59.67	203.63	28	94	2	97	96
96	446.598	9.91071	0.25	-59.67	203.63	28	-62.39	210.62	29.5	95	0	-1	-1
97	446.598	3.14479	0.25	-59.67	203.63	28	-60.91	200.74	28	95	0	-1	-1
98	426.806	2.0791	0.25	-52.74	185.55	27.5	-54.23	186.52	27.5	94	0	-1	-1
99	382.758	3.09666	0.25	-32.94	152.77	25.5	-29.97	153.49	25	93	0	-1	-1
100	232.446	3.15888	0.25	8.39	34.65	17	6.41	32.24	17.5	90	0	-1	-1
101	162.291	86.0909	0.25	21.51	-31.88	12	33.14	48.39	7.5	89	2	103	102
102	248.382	161.328	0.25	33.14	48.39	7.5	39.08	191.82	7.5	101	0	-1	-1
103	248.382	12.7362	0.25	33.14	48.39	7.5	32.89	58.04	5	101	0	-1	-1
104	137.374	17.5977	0.25	15.82	-54.54	9	3.19	-45.62	8.5	88	2	108	105
105	154.971	121.193	0.25	3.19	-45.62	8.5	-73.53	-15.01	22.5	104	2	107	106
106	276.164	92.4844	0.25	-73.53	-15.01	22.5	-19.58	2.59	33.5	105	0	-1	-1
107	276.164	45.1042	0.25	-73.53	-15.01	22.5	-98.28	-26.58	27	105	0	-1	-1
108	154.971	2.28013	0.25	3.19	-45.62	8.5	2.7	-47.79	9	104	0	-1	-1
109	118.792	4.34602	0.25	15.32	-71.89	7	18.29	-73.34	7.5	87	2	117	110
110	123.138	22.7967	0.25	18.29	-73.34	7.5	14.83	-92.38	6	109	2	116	111
111	145.935	4.66081	0.25	14.83	-92.38	6	13.59	-96.72	6.5	110	2	115	112
112	150.595	192.217	0.25	13.59	-96.72	6.5	-23.29	-217.25	27	111	2	114	113
113	342.812	14.6511	0.25	-23.29	-217.25	27	-25.52	-229.79	28	112	0	-1	-1
114	342.812	8.05651	0.25	-23.29	-217.25	27	-24.53	-223.76	28	112	0	-1	-1
115	150.595	109.61	0.25	13.59	-96.72	6.5	27.94	-194.83	6.5	111	0	-1	-1
116	145.935	3.24871	0.25	14.83	-92.38	6	18.04	-92.38	6.5	110	0	-1	-1
117	123.138	20.61	0.25	18.29	-73.34	7.5	34.13	-70.45	9.5	109	2	123	118
118	143.748	176.507	0.25	34.13	-70.45	9.5	-90.36	-165.43	11	117	2	122	119
119	320.255	24.2017	0.25	-90.36	-165.43	11	-105.21	-172.42	13.5	118	2	121	120
120	344.456	154.778	0.25	-105.21	-172.42	13.5	-184.16	-219.9	32	119	0	-1	-1
121	344.456	13.8192	0.25	-105.21	-172.42	13.5	-115.85	-176.03	14.5	119	0	-1	-1
122	320.255	1.90032	0.25	-90.36	-165.43	11	-89.12	-166.87	11	118	0	-1	-1
123	143.748	27.0031	0.25	34.13	-70.45	9.5	51.95	-69	12	117	2	139	124
124	170.751	26.4623	0.25	51.95	-69	12	66.55	-48.03	12	123	2	132	125
125	197.213	15.5675	0.25	66.55	-48.03	12	65.56	-33.57	12.5	124	2	131	126
126	212.781	46.2808	0.25	65.56	-33.57	12.5	38.83	-26.33	5.5	125	2	130	127
127	259.062	36.8367	0.25	38.83	-26.33	5.5	41.8	-15.01	0	126	2	129	128
128	295.898	21.5261	0.25	41.8	-15.01	0	38.59	-3.92	-3	127	0	-1	-1
129	295.898	2.46982	0.25	41.8	-15.01	0	43.54	-16.69	-0.5	127	0	-1	-1
130	259.062	1.55245	0.25	38.83	-26.33	5.5	38.59	-27.78	5	126	0	-1	-1
131	212.781	123.698	0.25	65.56	-33.57	12.5	88.58	61.41	1	125	0	-1	-1
132	197.213	112.285	0.25	66.55	-48.03	12	122.1	30.31	22.5	124	2	136	133
133	309.498	133.369	0.25	122.1	30.31	22.5	187.44	69.6	38.5	132	2	135	134
134	442.867	147.113	0.25	187.44	69.6	38.5	286.43	14.4	45.5	133	0	-1	-1
135	442.867	5.99344	0.25	187.44	69.6	38.5	187.93	75.15	37.5	133	0	-1	-1
136	309.498	95.952	0.25	122.1	30.31	22.5	139.07	36.1	35.5	132	2	138	137
137	405.45	62.1414	0.25	139.07	36.1	35.5	144.27	61.17	42	136	0	-1	-1
138	405.45	37.771	0.25	139.07	36.1	35.5	146.99	14.16	41.5	136	0	-1	-1
139	170.751	35.6998	0.25	51.95	-69	12	75.22	-76.23	16	123	2	161	140
140	206.451	13.4675	0.25	75.22	-76.23	16	85.36	-76.72	18	139	2	154	141
141	219.918	63.6581	0.25	85.36	-76.72	18	82.64	-136.74	18.5	140	2	151	142
142	283.576	11.903	0.25	82.64	-136.74	18.5	79.92	-147.59	19	141	2	150	143
143	295.479	23.1223	0.25	79.92	-147.59	19	77.94	-170.01	20.5	142	2	149	144
144	318.602	30.9486	0.25	77.94	-170.01	20.5	72.99	-198.21	21.5	143	2	148	145
145	349.55	207.595	0.25	72.99	-198.21	21.5	27.2	-363.82	47	144	2	147	146
146	557.146	124.906	0.25	27.2	-363.82	47	10.62	-448.43	59	145	0	-1	-1
147	557.146	5.57361	0.25	27.2	-363.82	47	24.73	-366.23	48.5	145	0	-1	-1
148	349.55	3.62263	0.25	72.99	-198.21	21.5	75.46	-195.56	21.5	144	0	-1	-1
149	318.602	3.54783	0.25	77.94	-170.01	20.5	76.95	-173.38	20	143	0	-1	-1
150	295.479	0.989998	0.25	79.92	-147.59	19	78.93	-147.59	19	142	0	-1	-1
151	283.576	37.9857	0.25	82.64	-136.74	18.5	99.47	-167.84	21.5	141	2	153	152
152	321.562	30.5537	0.25	99.47	-167.84	21.5	110.11	-194.59	21.5	151	0	-1	-1
153	321.562	6.10263	0.25	99.47	-167.84	21.5	97.99	-168.32	22	151	0	-1	-1
154	219.918	2.57691	0.25	85.36	-76.72	18	87.84	-76.23	18.5	140	2	156	155
155	222.495	139.285	0.25	87.84	-76.23	18.5	47	50.08	25	154	0	-1	-1
156	222.495	33.7124	0.25	87.84	-76.23	18.5	112.84	-76.72	22	154	2	160	157
157	256.208	44.8408	0.25	112.84	-76.72	22	139.07	-78.64	26.5	156	2	159	158
158	301.048	328.456	0.25	139.07	-78.64	26.5	353.4	-119.38	59.5	157	0	-1	-1
159	301.048	1.6985	0.25	139.07	-78.64	26.5	139.32	-76.96	26.5	157	0	-1	-1
160	256.208	1.55403	0.25	112.84	-76.72	22	112.59	-75.27	21.5	156	0	-1	-1
161	206.451	3.10484	0.25	75.22	-76.23	16	76.21	-79.13	16.5	139	0	-1	-1
162	89.108	35.4202	0.25963	1.96	-75.03	10.5	1.96	-107.81	12	2	2	184	163
163	124.528	15.3203	0.25	1.96	-107.81	12	-1.66	-110.67	5.5	162	2	179	164
164	139.848	2.64161	0.25	-1.66	-110.67	5.5	-1.41	-108.51	4	163	2	168	165
165	142.49	21.8029	0.25	-1.41	-108.51	4	2.79	-91.87	1	164	2	167	166
166	164.293	57.4134	0.25	2.79	-91.87	1	7	-57.4	-3	165	0	-1	-1
167	164.293	65.0653	0.25	2.79	-91.87	1	-30.37	-90.91	-4	165	0	-1	-1
168	142.49	26.6139	0.25	-1.41	-108.51	4	-3.89	-82.47	5.5	164	2	178	169
169	169.104	169.947	0.25	-3.89	-82.47	5.5	-84.08	39.26	2	168	2	177	170
170	339.051	8.58921	0.25	-84.08	39.26	2	-89.28	44.08	2.5	169	2	176	171
171	347.64	56.3944	0.25	-89.28	44.08	2.5	-125.16	67.95	1	170	2	175	172
172	404.035	62.4136	0.25	-125.16	67.95	1	-166.25	42.64	-0.5	171	2	174	173
173	466.448	14.9621	0.25	-166.25	42.64	-0.5	-175.16	33.72	-1	172	0	-1	-1
174	466.448	18.2188	0.25	-166.25	42.64	-0.5	-180.36	34.68	-1	172	0	-1	-1
175	404.035	38.0158	0.25	-125.16	67.95	1	-129.37	101.7	0.5	171	0	-1	-1
176	347.64	3.33306	0.25	-89.28	44.08	2.5	-86.8	46.25	3	170	0	-1	-1
177	339.051	20.0915	0.25	-84.08	39.26	2	-100.91	46.01	3	169	0	-1	-1
178	169.104	2.03735	0.25	-3.89	-82.47	5.5	-1.91	-81.99	5.5	168	0	-1	-1
179	139.848	36.8282	0.25	-1.66	-110.67	5.5	-27.4	-114.77	1.5	163	2	183	180
180	176.677	69.2702	0.25	-27.4	-114.77	1.5	-62.3	-123.21	-3	179	2	182	181
181	245.947	29.7729	0.25	-62.3	-123.21	-3	-67.74	-147.07	-3	180	0	-1	-1
182	245.947	6.66381	0.25	-62.3	-123.21	-3	-60.81	-125.38	-4	180	0	-1	-1
183	176.677	2.11116	0.25	-27.4	-114.77	1.5	-25.67	-115.98	1.5	179	0	-1	-1
184	124.528	18.8093	0.25	1.96	-107.81	12	3.44	-125.65	14	162	2	196	185
185	143.337	7.62596	0.25	3.44	-125.65	14	4.82	-125.62	6.5	184	2	191	186
186	150.963	53.4629	0.25	4.82	-125.62	6.5	9.77	-127.07	-1.5	185	2	190	187
187	204.426	43.8926	0.25	9.77	-127.07	-1.5	-20.92	-108.75	-2.5	186	2	189	188
188	248.319	48.568	0.25	-20.92	-108.75	-2.5	-56.07	-95.25	-3	187	0	-1	-1
189	248.319	2.92577	0.25	-20.92	-108.75	-2.5	-22.16	-106.1	-2.5	187	0	-1	-1
190	204.426	25.7183	0.25	9.77	-127.07	-1.5	20.41	-136.71	-4.5	186	0	-1	-1
191	150.963	37.5298	0.25	4.82	-125.62	6.5	31.3	-146.35	5	185	2	193	192
192	188.493	118.79	0.25	31.3	-146.35	5	131.53	-156	8.5	191	0	-1	-1
193	188.493	306.797	0.25	31.3	-146.35	5	222.37	-336.79	-4	191	2	195	194
194	495.29	22.2995	0.25	222.37	-336.79	-4	232.76	-340.4	-5.5	193	0	-1	-1
195	495.29	3.32216	0.25	222.37	-336.79	-4	220.14	-336.55	-4	193	0	-1	-1
196	143.337	20.432	0.25	3.44	-125.65	14	1.21	-144.21	16	184	2	248	197
197	163.769	75.9259	0.25	1.21	-144.21	16	-5.47	-217.25	20.5	196	2	199	198
198	239.695	1.7596	0.25	-5.47	-217.25	20.5	-5.96	-218.94	20.5	197	0	-1	-1
199	239.695	59.8859	0.25	-5.47	-217.25	20.5	3.69	-269.56	29.5	197	2	229	200
200	299.581	56.339	0.25	3.69	-269.56	29.5	0.72	-323.32	35.5	199	2	208	201
201	355.92	59.0012	0.25	0.72	-323.32	35.5	-2	-377.56	43	200	2	205	202
202	414.921	61.8899	0.25	-2	-377.56	43	-2.99	-427.21	52.5	201	2	204	203
203	476.811	42.944	0.25	-2.99	-427.21	52.5	1.21	-459.76	59	202	0	-1	-1
204	476.811	143.326	0.25	-2.99	-427.21	52.5	-37.15	-399.73	159.5	202	0	-1	-1
205	414.921	19.6231	0.25	-2	-377.56	43	-8.44	-390.33	39	201	2	207	206
206	434.544	55.7308	0.25	-8.44	-390.33	39	10.87	-399.97	24.5	205	0	-1	-1
207	434.544	322.59	0.25	-8.44	-390.33	39	-198.52	-532.31	35	205	0	-1	-1
208	355.92	11.9345	0.25	0.72	-323.32	35.5	8.64	-328.38	38.5	200	2	228	209
209	367.855	2.33523	0.25	8.64	-328.38	38.5	10.87	-328.86	38	208	2	219	210
210	370.19	96.0193	0.25	10.87	-328.86	38	87.1	-327.66	46	209	2	216	211
211	466.209	121.425	0.25	87.1	-327.66	46	181.64	-356.1	45	210	2	213	212
212	587.634	1.97955	0.25	181.64	-356.1	45	181.39	-354.41	44	211	0	-1	-1
213	587.634	2.83581	0.25	181.64	-356.1	45	184.11	-357.07	44	211	2	215	214
214	590.469	3.32129	0.25	184.11	-357.07	44	186.34	-354.66	44.5	213	0	-1	-1
215	590.469	2.70591	0.25	184.11	-357.07	44	186.59	-358.03	44.5	213	0	-1	-1
216	466.209	135.917	0.25	87.1	-327.66	46	44.53	-294.87	7	210	2	218	217
217	602.126	69.1277	0.25	44.53	-294.87	7	4.18	-320.67	4.5	216	0	-1	-1
218	602.126	41.6947	0.25	44.53	-294.87	7	46.26	-285.95	-2.5	216	0	-1	-1
219	370.19	60.3206	0.25	10.87	-328.86	38	42.3	-354.41	49	209	2	221	220
220	430.51	3.5153	0.25	42.3	-354.41	49	45.02	-352.24	48.5	219	0	-1	-1
221	430.51	15.4121	0.25	42.3	-354.41	49	54.92	-360.92	49.5	219	2	225	222
222	445.922	6.11469	0.25	54.92	-360.92	49.5	57.64	-365.26	50	221	2	224	223
223	452.037	30.2699	0.25	57.64	-365.26	50	74.72	-385.03	52	222	0	-1	-1
224	452.037	25.6433	0.25	57.64	-365.26	50	64.82	-361.4	58.5	222	0	-1	-1
225	445.922	41.0288	0.25	54.92	-360.92	49.5	71.75	-335.85	54	221	2	227	226
226	486.951	61.3107	0.25	71.75	-335.85	54	121	-341.88	53.5	225	0	-1	-1
227	486.951	54.3036	0.25	71.75	-335.85	54	101.7	-311.99	65	225	0	-1	-1
228	367.855	69.29	0.25	8.64	-328.38	38.5	40.32	-329.83	57.5	208	0	-1	-1
229	299.581	69.4655	0.25	3.69	-269.56	29.5	-25.27	-312.95	39.5	199	2	231	230
230	369.047	19.7465	0.25	-25.27	-312.95	39.5	-39.38	-316.57	36.5	229	0	-1	-1
231	369.047	9.30393	0.25	-25.27	-312.95	39.5	-30.96	-317.77	42	229	2	237	232
232	378.351	28.1984	0.25	-30.96	-317.77	42	-52.25	-318.5	37	231	2	234	233
233	406.549	2.22687	0.25	-52.25	-318.5	37	-51.75	-316.33	37	232	0	-1	-1
234	406.549	5.62662	0.25	-52.25	-318.5	37	-56.21	-319.46	35.5	232	2	236	235
235	412.176	33.6413	0.25	-56.21	-319.46	35.5	-81.2	-322.35	32	234	0	-1	-1
236	412.176	199.124	0.25	-56.21	-319.46	35.5	-189.11	-305.48	29.5	234	0	-1	-1
237	378.351	38.9263	0.25	-30.96	-317.77	42	-51.75	-346.94	47	231	2	239	238
238	417.277	254.536	0.25	-51.75	-346.94	47	-35.66	-437.34	151	237	0	-1	-1
239	417.277	23.2714	0.25	-51.75	-346.94	47	-64.87	-360.44	50.5	237	2	247	240
240	440.548	201.915	0.25	-64.87	-360.44	50.5	-109.17	-416.61	124.5	239	2	242	241
241	642.463	59.1076	0.25	-109.17	-416.61	124.5	-110.66	-416.37	147	240	0	-1	-1
242	642.463	12.9468	0.25	-109.17	-416.61	124.5	-109.67	-418.05	112.5	240	2	246	243
243	655.41	154.128	0.25	-109.67	-418.05	112.5	-46.06	-485.07	75.5	242	2	245	244
244	809.538	13.4269	0.25	-46.06	-485.07	75.5	-38.88	-479.52	69	243	0	-1	-1
245	809.538	42.4345	0.25	-46.06	-485.07	75.5	-41.85	-487.24	55	243	0	-1	-1
246	655.41	77.5734	0.25	-109.67	-418.05	112.5	-86.65	-427.45	145.5	242	0	-1	-1
247	440.548	110.929	0.25	-64.87	-360.44	50.5	-120.06	-317.77	59.5	239	0	-1	-1
248	163.769	12.1606	0.25	1.21	-144.21	16	-4.97	-137.22	18	196	2	250	249
249	175.93	2.76203	0.25	-4.97	-137.22	18	-2.25	-136.74	18	248	0	-1	-1
250	175.93	13.5718	0.25	-4.97	-137.22	18	-9.68	-127.82	20.5	248	2	256	251
251	189.502	159.044	0.25	-9.68	-127.82	20.5	-92.34	-12.11	26	250	2	253	252
252	348.546	5.89196	0.25	-92.34	-12.11	26	-92.09	-6.57	26	251	0	-1	-1
253	348.546	124.569	0.25	-92.34	-12.11	26	-140.6	93.95	24.5	251	2	255	254
254	473.115	67.8029	0.25	-140.6	93.95	24.5	-180.7	139.27	27.5	253	0	-1	-1
255	473.115	27.0398	0.25	-140.6	93.95	24.5	-155.95	111.31	22	253	0	-1	-1
256	189.502	54.986	0.25	-9.68	-127.82	20.5	-3.98	-98.65	35.5	250	2	262	257
257	244.488	195.328	0.25	-3.98	-98.65	35.5	8.14	75.87	50	256	2	259	258
258	439.816	7.42945	0.25	8.14	75.87	50	3.44	77.08	54	257	0	-1	-1
259	439.816	100.778	0.25	8.14	75.87	50	-0.77	156.87	56.5	257	2	261	260
260	540.594	5.57239	0.25	-0.77	156.87	56.5	3.19	160.24	57.5	259	0	-1	-1
261	540.594	14.0036	0.25	-0.77	156.87	56.5	-0.02	165.55	57.5	259	0	-1	-1
262	244.488	349.783	0.25	-3.98	-98.65	35.5	43.78	80.94	137.5	256	0	-1	-1

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