arthyperАртериальная гипертензия"Arterial’naya Gipertenziya" ("Arterial Hypertension")1607-419X2411-8524Antihypertensive League10.18705/1607-419X-2021-27-5-499-508arthyper-2133Research ArticleСтатьиArticlesНейрофизиологическое обоснование гипотезы Г. Ф. Ланга о возникновении гипертонической болезниNeurophysiological justification of the G. F. Lang hypothesis about the occurrence of essential hypertensionhttps://orcid.org/0000-0003-2929-0980ШляхтоЕ. В.ShlyakhtoE. V.

Шляхто Евгений Владимирович — доктор медицинских наук, профессор, академик РАН, генеральный директор

пр. Пархоменко, д. 15, Санкт-Петербург, Россия,194156

Eugene V. Shlyakhto, MD, PhD, DSc, Professor, Academician of the Russian Academy of Sciences, General Director

15 Parkhomenko avenue, St Petersburg, 194156

e.shlyakhto@almazovcentre.ruhttps://orcid.org/0000-0002-7767-8560ЦырлинВ. А.TsyrlinV. A.

Цырлин Виталий Александрович — главный научный сотрудник отдела экспериментальной физиологии и фармакологии Центра доклинических и трансляционных исследований

пр. Пархоменко, д. 15, Санкт-Петербург, Россия,194156

Vitaliy A. Tsyrlin, MD, Ph D. DSc, Professor, Chief Researcher, Department of Experimental Physiology and Pharmacology, Preclinical and Translational Research Centre

15 Parkhomenko avenue, St Petersburg, 194156

tsyrlinva@mail.ruhttps://orcid.org/0000-0001-6027-7325КузьменкоН. В.KuzmenkoN. V.

Кузьменко Наталия Владимировна — старший научный сотрудник отдела экспериментальной физиологии и фармакологии Центра доклинических и трансляционных исследований; младшийнаучный сотрудник лаборатории биофизики кровообращения

пр. Пархоменко, д. 15, Санкт-Петербург, Россия,194156

Natalia V. Kuzmenko, PhD, Senior Researcher, Department of Experimental Physiology and Pharmacology, Preclinical and Translational Research Centre; Researcher, Blood Circulation Biophysics Laboratory

15 Parkhomenko avenue, St Petersburg, 194156

nat.kuzmencko2011@yandex.ruhttps://orcid.org/0000-0002-1515-1616ПлиссМ. Г.PlissM. G.

Плисс Михаил Геннадиевич — заведующий отделом экспериментальной физиологии и фармакологии Центра доклинических и трансляционных исследований; заведующий лабораторией биофизики кровообращения

пр. Пархоменко, д. 15, Санкт-Петербург, Россия,194156

 

Mikhail G. Pliss, PhD, Head, Department of Experimental Physiology and Pharmacology, Preclinical and Translational Research Centre; Head, Blood Circulation Biophysics Laboratory

15 Parkhomenko avenue, St Petersburg, 194156

pliss@niiekf.ruФедеральное государственное бюджетное учреждение «Национальный медицинский исследовательский центр имени В. А. Алмазова» Министерства здравоохранения Российской Федерации; Государственное бюджетное образовательное учреждение высшего профессионального образования «Первый Санкт-Петербургский государственный медицинский университет имени академика И. П. Павлова» Министерства здравоохранения Российской ФедерацииAlmazov National Medical Research Centre; Pavlov UniversityФедеральное государственное бюджетное учреждение «Национальный медицинский исследовательский центр имени В. А. Алмазова» Министерства здравоохранения Российской ФедерацииAlmazov National Medical Research Centre202109102021275499508Copyright © Шляхто Е.В., Цырлин В.А., Кузьменко Н.В., Плисс М.Г., 20222022Шляхто Е.В., Цырлин В.А., Кузьменко Н.В., Плисс М.Г.Shlyakhto E.V., Tsyrlin V.A., Kuzmenko N.V., Pliss M.G.Данная работа распространяется под лицензией Creative Commons Attribution 4.0.This work is licensed under a Creative Commons Attribution 4.0 License.https://htn.almazovcentre.ru/jour/article/view/2133

Настоящая статья представляет собой анализ современных данных о морфологической, функциональной и нейрохимической организации центральных механизмов регуляции кровообращения, нарушение которых может являться причиной развития гипертонической болезни. Приводятся сведения, указывающие на справедливость гипотезы Г.Ф. Ланга о том, что длительный эмоциональный стресс приводит к нарушению нейрональной активности структур гипоталамуса. В статье описаны морфологические связи гипоталамуса с нейронами вентролатеральной области продолговатого мозга и структурами спинного мозга, обеспечивающие нейрогенный тонус сосудов. Представлены доказательства конвергенции аксонов нейронов паравентрикулярного ядра гипоталамуса на преганглионарные нейроны спинного мозга. Рассматриваются нейрохимические процессы в вазомоторном центре продолговатого мозга, указаны морфологические изменения преганглионарных симпатических нейронов при экспериментальной артериальной гипертензии.

This article is an analysis of modern data on the morphological, functional and neurochemical organization of the central mechanisms of blood circulation regulation, the violation of which can cause the development of essential hypertension. The data indicating the validity of the hypothesis of G.F. Lang that prolonged emotional stress leads to a violation of the neuronal activity of the hypothalamus structures are presented. The article describes the morphological connections of the hypothalamus with the neurons of the ventrolateral region of the medulla oblongata and the structures of the spinal cord that provide neurogenic vascular tone. The evidence of convergence of axons of neurons of the paraventricular nucleus of the hypothalamus on the preganglionic neurons of the spinal cord is presented. Neurochemical processes in the vasomotor center of the medulla oblongata are considered, morphological changes of preganglionic sympathetic neurons in experimental arterial hypertension are indicated.

гипертоническая болезньгипоталамусвазомоторный центрпреганглионарные симпатические нейроныглутаматessential hypertensionhypothalamusvasomotor centerpreganglionic sympathetic neuronsglutamateРабота финансирована из средств Государ- ственного задания № 056–00109–21–02.The authors declare no conflict of interest.ReferencesЛанг Г.Ф. Гипертоническая болезнь. Л.,Медгиз, 1950, 496с.Lang G. F. Hypertension. L., Medgiz, 1950, 496 р. In Russian.Шляхто Е.В., Конради А.О., Цырлин В.А. Вегетативная нервная система и артериальная гипертензия. СПб,ООО « Медицинское издательство», 2008, 312 с.Shlyakhto E. V., Konradi A. O., Tsyrlin V. A. Autonomic nervous system and arterial hypertension. St. Petersburg, LLC "Medical publishing house", 2008, 312 p. In Russian.Вальдман А.В. Нейрофармакология центральной регуляции сосудистого тонуса. Л., Медицина, 1976, 326с.Valdman A.V. Neuropharmacology of central vascular regulation tonusa. L., Medicine, 1976, 326 р. In Russian.Schlager G. Selection for blood pressure levels in mice. Genetics 1974;76(3): 537- 549.Schlager G. Selection for blood pressure levels in mice. Genetics 1974;76(3): 537- 549.Okamoto K., Aoki K. Development of a strain of spontaneously hypertensive rats. Jap.circulat.J.,1963;.27: 282-293.Okamoto K., Aoki K. Development of a strain of spontaneously hypertensive rats. Jap.circulat.J.,1963;.27: 282-293.Okamoto K. Spontaneous hypertensive in rats. Exp.path.1969; 7: 227- 269.Okamoto K. Spontaneous hypertensive in rats. Exp.path.1969; 7: 227- 269.Saper C.B., Loewy A.D., Swanson L.W., Cowan W.M. Direct hypothalamo-autonomic connections. Brain Res. 1976; 117: 305–312. doi:10.1016/0006- 8993(76)90738-1Saper C.B., Loewy A.D., Swanson L.W., Cowan W.M. Direct hypothalamo-autonomic connections. Brain Res. 1976; 117: 305–312. doi:10.1016/0006- 8993(76)90738-1Cechetto D.F., Saper C.B. Neurochemical organization of the hypothalamic projection to the spinal cord in the rat. J. Comp. Neurol. 1988; 272(4): 579-604. doi:10.1002/cne.902720410Cechetto D.F., Saper C.B. Neurochemical organization of the hypothalamic projection to the spinal cord in the rat. J. Comp. Neurol. 1988; 272(4): 579-604. doi:10.1002/cne.902720410Strack A.M., Sawyer W.B., Hughes J.H., Platt K.B., Loewy A.D. A general pattern of CNS innervation of the sympathetic outflow demonstrated by transneuronal pseudorabies viral infections. Brain Res. 1989;491: 156–162. doi:10.1016/0006-8993(89)90098-X.[PubMed]Strack A.M., Sawyer W.B., Hughes J.H., Platt K.B., Loewy A.D. A general pattern of CNS innervation of the sympathetic outflow demonstrated by transneuronal pseudorabies viral infections. Brain Res. 1989;491: 156–162. doi:10.1016/0006-8993(89)90098-X.[PubMed]Coote J.H. A role for the paraventricular nucleus of the hypothalamus in the autonomic control of heart and kidney. Exp. Physiol. 2005; 90: 169–173.Coote J.H. A role for the paraventricular nucleus of the hypothalamus in the autonomic control of heart and kidney. Exp. Physiol. 2005; 90: 169–173.Hallbeck M., Larhammar D., Blomqvist A. Neuropeptide expression in rat paraventricular hypothalamic neurons that project to the spinal cord. J. Comp.Neurol .2001; 433: 222–238. doi:10.1002/cne.113711 Hallbeck M., Larhammar D., Blomqvist A. Neuropeptide expression in rat paraventricular hypothalamic neurons that project to the spinal cord. J. Comp.Neurol .2001; 433: 222–238. doi:10.1002/cne.1137Benarroch E.E. Paraventricular nucleus, stress response, and cardiovascular disease. Clin. Auton. Res. 2005; 15: 254–263. doi:10.1007/s10286-005-0290-7Benarroch E.E. Paraventricular nucleus, stress response, and cardiovascular disease. Clin. Auton. Res. 2005; 15: 254–263. doi:10.1007/s10286-005-0290-7Pyner S.J. Neurochemistry of the paraventricular nucleus of the hypothalamus: implications for cardiovascular regulation. Chem. Neuroanat. 2009; 38(3):197-208. doi: 10.1016/j.jchemneu.2009.03.005Pyner S.J. Neurochemistry of the paraventricular nucleus of the hypothalamus: implications for cardiovascular regulation. Chem. Neuroanat. 2009; 38(3):197-208. doi: 10.1016/j.jchemneu.2009.03.005Motawei K., Pyner S., Ranson R.N., Kamel M., Coote J.H. Terminals of paraventricular spinal neurones are closely associated with adrenal medullary sympathetic preganglionic neurones: immunocytochemical evidence for vasopressin as a possible neurotransmitter in this pathway. Exp Brain Res. 1999; 126(1): 68-76. doi:10.1007/s002210050717Motawei K., Pyner S., Ranson R.N., Kamel M., Coote J.H. Terminals of paraventricular spinal neurones are closely associated with adrenal medullary sympathetic preganglionic neurones: immunocytochemical evidence for vasopressin as a possible neurotransmitter in this pathway. Exp Brain Res. 1999; 126(1): 68-76. doi:10.1007/s002210050717Сентаготаи Я. ,Флерко Б., Меш Б., Халас Б. Гипоталамическая регуляция передней части гипофиза. АН Венгрии, Будапешт, 1968, 353 с.Sentagotai Ya., Flerko B., Mesh B., Halas B. Hypothalamic regulation of the anterior pituitary gland. Hungarian Academy of Sciences, Budapest, 1968, 353 p.Dampney R.A., Michelini L.C., Li D.P., Pan H.L. Regulation of sympathetic vasomotor activity by the hypothalamic paraventricular nucleus in normotensive and hypertensive states. Am J Physiol Heart Circ Physiol. 2018;315(5):H1200-H1214. doi: 10.1152/ajpheart.00216.2018Dampney R.A., Michelini L.C., Li D.P., Pan H.L. Regulation of sympathetic vasomotor activity by the hypothalamic paraventricular nucleus in normotensive and hypertensive states. Am J Physiol Heart Circ Physiol. 2018;315(5):H1200-H1214. doi: 10.1152/ajpheart.00216.2018Kunkler P.E., Hwang B.H. Lower GABAA receptor binding in the amygdala and hypothalamus of spontaneously hypertensive rats. Brain Res Bull. 1995;36(1):57-61. doi: 10.1016/0361-9230(94)00164-vKunkler P.E., Hwang B.H. Lower GABAA receptor binding in the amygdala and hypothalamus of spontaneously hypertensive rats. Brain Res Bull. 1995;36(1):57-61. doi: 10.1016/0361-9230(94)00164-vQiao X., Zhou J.J., Li D.P., Pan H.L. Src Kinases Regulate Glutamatergic Input to Hypothalamic Presympathetic Neurons and Sympathetic Outflow in Hypertension. Hypertension. 2017;69(1):154-162. doi: 10.1161/HYPERTENSIONAHA.116.07947Qiao X., Zhou J.J., Li D.P., Pan H.L. Src Kinases Regulate Glutamatergic Input to Hypothalamic Presympathetic Neurons and Sympathetic Outflow in Hypertension. Hypertension. 2017;69(1):154-162. doi: 10.1161/HYPERTENSIONAHA.116.07947Gabor A., Leenen F.H. Central neuromodulatory pathways regulating sympathetic activity in hypertension. J Appl Physiol (1985). 2012;113(8):1294-303. doi: 10.1152/japplphysiol.00553.2012Gabor A., Leenen F.H. Central neuromodulatory pathways regulating sympathetic activity in hypertension. J Appl Physiol (1985). 2012;113(8):1294-303. doi: 10.1152/japplphysiol.00553.2012Koba S., Hanai E., Kumada N., Kataoka N., Nakamura K., Watanabe T. Sympathoexcitation by hypothalamic paraventricular nucleus neurons projecting to the rostral ventrolateral medulla. J Physiol. 2018;596(19):4581-4595. doi: 10.1113/JP276223Koba S., Hanai E., Kumada N., Kataoka N., Nakamura K., Watanabe T. Sympathoexcitation by hypothalamic paraventricular nucleus neurons projecting to the rostral ventrolateral medulla. J Physiol. 2018;596(19):4581-4595. doi: 10.1113/JP276223Basting T., Xu J., Mukerjee S., Epling J., Fuchs R., Sriramula S., Lazartigues E. Glutamatergic neurons of the paraventricular nucleus are critical contributors to the development of neurogenic hypertension. J Physiol. 2018; 596(24):6235-6248. doi: 10.1113/JP276229Basting T., Xu J., Mukerjee S., Epling J., Fuchs R., Sriramula S., Lazartigues E. Glutamatergic neurons of the paraventricular nucleus are critical contributors to the development of neurogenic hypertension. J Physiol. 2018; 596(24):6235-6248. doi: 10.1113/JP276229Soriano J.E., Scott B.A., Rosentreter R.E., Vaseghi B. The sympathetic role of glutamatergic paraventricular nucleus neurons in blood pressure regulation. J Physiol. 2019;597(6):1433-1434. doi: 10.1113/JP277558Soriano J.E., Scott B.A., Rosentreter R.E., Vaseghi B. The sympathetic role of glutamatergic paraventricular nucleus neurons in blood pressure regulation. J Physiol. 2019;597(6):1433-1434. doi: 10.1113/JP277558Ali D.W., Salter M.W. NMDA receptor regulation by Src kinase signalling in excitatory synaptic transmission and plasticity. Curr Opin Neurobiol. 2001;11(3):336-42. doi: 10.1016/s0959-4388(00)00216-6Ali D.W., Salter M.W. NMDA receptor regulation by Src kinase signalling in excitatory synaptic transmission and plasticity. Curr Opin Neurobiol. 2001;11(3):336-42. doi: 10.1016/s0959-4388(00)00216-6Allen A.M. Inhibition of the hypothalamic paraventricular nucleus in spontaneously hypertensive rats dramatically reduces sympathetic vasomotor tone. Hypertension. 2002;39(2):275-80. doi: 10.1161/hy0202.104272Allen A.M. Inhibition of the hypothalamic paraventricular nucleus in spontaneously hypertensive rats dramatically reduces sympathetic vasomotor tone. Hypertension. 2002;39(2):275-80. doi: 10.1161/hy0202.104272Li D.P., Yang Q., Pan H.M., Pan H.L. Pre- and postsynaptic plasticity underlying augmented glutamatergic inputs to hypothalamic presympathetic neurons in spontaneously hypertensive rats. J Physiol. 2008;586(6):1637-47. doi: 10.1113/jphysiol.2007.149732Li D.P., Yang Q., Pan H.M., Pan H.L. Pre- and postsynaptic plasticity underlying augmented glutamatergic inputs to hypothalamic presympathetic neurons in spontaneously hypertensive rats. J Physiol. 2008;586(6):1637-47. doi: 10.1113/jphysiol.2007.149732Li D.P., Pan H.L. Increased group I metabotropic glutamate receptor activity in paraventricular nucleus supports elevated sympathetic vasomotor tone in hypertension. Am J Physiol Regul Integr Comp Physiol. 2010;299(2):R552-61. doi: 10.1152/ajpregu.00195.2010Li D.P., Pan H.L. Increased group I metabotropic glutamate receptor activity in paraventricular nucleus supports elevated sympathetic vasomotor tone in hypertension. Am J Physiol Regul Integr Comp Physiol. 2010;299(2):R552-61. doi: 10.1152/ajpregu.00195.2010Ye Z.Y., Li D.P., Li L., Pan H.L. Protein kinase CK2 increases glutamatergic input in the hypothalamus and sympathetic vasomotor tone in hypertension. J Neurosci. 2011;31(22):8271-9. doi: 10.1523/JNEUROSCI.1147-11.2011Ye Z.Y., Li D.P., Li L., Pan H.L. Protein kinase CK2 increases glutamatergic input in the hypothalamus and sympathetic vasomotor tone in hypertension. J Neurosci. 2011;31(22):8271-9. doi: 10.1523/JNEUROSCI.1147-11.2011Ito S., Komatsu K., Tsukamoto K., Sved A.F. Excitatory amino acids in the rostral ventrolateral medulla support blood pressure in spontaneously hypertensive rats. Hypertension. 2000;35(1 Pt 2):413-7. doi: 10.1161/01.hyp.35.1.413Ito S., Komatsu K., Tsukamoto K., Sved A.F. Excitatory amino acids in the rostral ventrolateral medulla support blood pressure in spontaneously hypertensive rats. Hypertension. 2000;35(1 Pt 2):413-7. doi: 10.1161/01.hyp.35.1.413Ito S., Komatsu K., Tsukamoto K., Sved A.F. Tonic excitatory input to the rostral ventrolateral medulla in Dahl salt-sensitive rats. Hypertension. 2001;37(2):687-91Ito S., Komatsu K., Tsukamoto K., Sved A.F. Tonic excitatory input to the rostral ventrolateral medulla in Dahl salt-sensitive rats. Hypertension. 2001;37(2):687-91Colombari E., Sato M.A., Cravo S.L., Bergamaschi C.T., Campos R.R. Jr, Lopes O.U. Role of the medulla oblongata in hypertension. Hypertension. 2001;38(3 Pt 2):549-54. doi: 10.1161/01.hyp.38.3.549Colombari E., Sato M.A., Cravo S.L., Bergamaschi C.T., Campos R.R. Jr, Lopes O.U. Role of the medulla oblongata in hypertension. Hypertension. 2001;38(3 Pt 2):549-54. doi: 10.1161/01.hyp.38.3.549Bergamaschi C., Campos R.R., Schor N., Lopes O.U. Importance of rostral ventrolateral medulla in rats with Goldblatt hypertension. Fundam Clin Pharmacol. 1997; 11:92S–93SBergamaschi C., Campos R.R., Schor N., Lopes O.U. Importance of rostral ventrolateral medulla in rats with Goldblatt hypertension. Fundam Clin Pharmacol. 1997; 11:92S–93SHuber M.J., Chen Q.H., Shan Z. The Orexin System and Hypertension. Cell Mol Neurobiol. 2018;38(2):385-391. doi: 10.1007/s10571-017-0487-zHuber M.J., Chen Q.H., Shan Z. The Orexin System and Hypertension. Cell Mol Neurobiol. 2018;38(2):385-391. doi: 10.1007/s10571-017-0487-zGeerling J.C., Shin J.W., Chimenti P.C., Loewy A.D. Paraventricular hypothalamic nucleus: axonal projections to the brainstem. J Comp Neurol. 2010;518(9):1460-99. doi: 10.1002/cne.22283Geerling J.C., Shin J.W., Chimenti P.C., Loewy A.D. Paraventricular hypothalamic nucleus: axonal projections to the brainstem. J Comp Neurol. 2010;518(9):1460-99. doi: 10.1002/cne.22283Guyenet P.G., Stornetta R.L., Holloway B.B., Souza G.M.P.R., Abbott S.B.G. Rostral Ventrolateral Medulla and Hypertension. Hypertension. 2018;72(3):559-566. doi: 10.1161/HYPERTENSIONAHA.118.10921Guyenet P.G., Stornetta R.L., Holloway B.B., Souza G.M.P.R., Abbott S.B.G. Rostral Ventrolateral Medulla and Hypertension. Hypertension. 2018;72(3):559-566. doi: 10.1161/HYPERTENSIONAHA.118.10921Li C., Horn J.P. Physiological classification of sympathetic neurons in the rat superior cervical ganglion. J Neurophysiol. 2006;95(1):187-95. doi: 10.1152/jn.00779.2005Li C., Horn J.P. Physiological classification of sympathetic neurons in the rat superior cervical ganglion. J Neurophysiol. 2006;95(1):187-95. doi: 10.1152/jn.00779.2005Minson J.B., Arnolda L.F., Llewellyn-Smith I.J. Neurochemistry of nerve fibers apposing sympathetic preganglionic neurons activated by sustained hypotension. J Comp Neurol. 2002;449(4):307-18. doi: 10.1002/cne.10282Minson J.B., Arnolda L.F., Llewellyn-Smith I.J. Neurochemistry of nerve fibers apposing sympathetic preganglionic neurons activated by sustained hypotension. J Comp Neurol. 2002;449(4):307-18. doi: 10.1002/cne.10282Kumar N.N., Allen K., Parker L., Damanhuri H., Goodchild A.K. Neuropeptide coding of sympathetic preganglionic neurons; focus on adrenally projecting populations. Neuroscience. 2010;170(3):789-99. doi: 10.1016/j.neuroscience.2010.07.047Kumar N.N., Allen K., Parker L., Damanhuri H., Goodchild A.K. Neuropeptide coding of sympathetic preganglionic neurons; focus on adrenally projecting populations. Neuroscience. 2010;170(3):789-99. doi: 10.1016/j.neuroscience.2010.07.047Bernstein-Goral H., Bohn M.C. Ontogeny of adrenergic fibers in rat spinal cord in relationship to adrenal preganglionic neurons. J Neurosci Res. 1988;21(2-4):333-51. doi: 10.1002/jnr.490210226Bernstein-Goral H., Bohn M.C. Ontogeny of adrenergic fibers in rat spinal cord in relationship to adrenal preganglionic neurons. J Neurosci Res. 1988;21(2-4):333-51. doi: 10.1002/jnr.490210226Llewellyn-Smith I.J. Anatomy of synaptic circuits controlling the activity of sympathetic preganglionic neurons. J Chem Neuroanat. 2009;38(3):231-9. doi: 10.1016/j.jchemneu.2009.06.001Llewellyn-Smith I.J. Anatomy of synaptic circuits controlling the activity of sympathetic preganglionic neurons. J Chem Neuroanat. 2009;38(3):231-9. doi: 10.1016/j.jchemneu.2009.06.001McNair C.J., Baxter G.J., Kerr R., Maxwell D.J. Glutamate receptor subunits associated with rat sympathetic preganglionic neurons. Neurosci Lett. 1998;256(1):29-32. doi: 10.1016/s0304-3940(98)00747-2McNair C.J., Baxter G.J., Kerr R., Maxwell D.J. Glutamate receptor subunits associated with rat sympathetic preganglionic neurons. Neurosci Lett. 1998;256(1):29-32. doi: 10.1016/s0304-3940(98)00747-2Hosoya Y., Matsukawa, Okado N., Sugiura Y., Kohno K. Oxytocinergic innervation to the upper thoracic sympathetic preganglionic neurons in the rat. A light and electron microscopical study using a combined retrograde transport and immunocytochemical technique. Exp Brain Res. 1995;107(1):9-16. doi: 10.1007/BF00228011Hosoya Y., Matsukawa, Okado N., Sugiura Y., Kohno K. Oxytocinergic innervation to the upper thoracic sympathetic preganglionic neurons in the rat. A light and electron microscopical study using a combined retrograde transport and immunocytochemical technique. Exp Brain Res. 1995;107(1):9-16. doi: 10.1007/BF00228011Whyment A.D., Blanks A.M., Lee K., Renaud L.P., Spanswick D. Histamine excites neonatal rat sympathetic preganglionic neurons in vitro via activation of H1 receptors. J Neurophysiol. 2006;95(4):2492-500. doi: 10.1152/jn.01135.2004Whyment A.D., Blanks A.M., Lee K., Renaud L.P., Spanswick D. Histamine excites neonatal rat sympathetic preganglionic neurons in vitro via activation of H1 receptors. J Neurophysiol. 2006;95(4):2492-500. doi: 10.1152/jn.01135.2004Minoura Y., Onimaru H., Iigaya K., Homma I., Kobayashi Y. Electrophysiological responses of sympathetic preganglionic neurons to ANG II and aldosterone. Am J Physiol Regul Integr Comp Physiol. 2009;297(3):R699-706. doi: 10.1152/ajpregu.00041.2009Minoura Y., Onimaru H., Iigaya K., Homma I., Kobayashi Y. Electrophysiological responses of sympathetic preganglionic neurons to ANG II and aldosterone. Am J Physiol Regul Integr Comp Physiol. 2009;297(3):R699-706. doi: 10.1152/ajpregu.00041.2009McAllen R.M., Häbler H.J., Michaelis M., Peters O., Jänig W. Monosynaptic excitation of preganglionic vasomotor neurons by subretrofacial neurons of the rostral ventrolateral medulla. Brain Res. 1994;634(2):227-34. doi: 10.1016/0006-8993(94)91925-9McAllen R.M., Häbler H.J., Michaelis M., Peters O., Jänig W. Monosynaptic excitation of preganglionic vasomotor neurons by subretrofacial neurons of the rostral ventrolateral medulla. Brain Res. 1994;634(2):227-34. doi: 10.1016/0006-8993(94)91925-9Rexed B. A cytoarchitectonic atlas of the spinal cord in the cat. J Comp Neurol. 1954;100(2):297-379. doi: 10.1002/cne.901000205Rexed B. A cytoarchitectonic atlas of the spinal cord in the cat. J Comp Neurol. 1954;100(2):297-379. doi: 10.1002/cne.901000205Deuchars S.A. How sympathetic are your spinal cord circuits? Exp Physiol. 2015;100(4):365-71. doi: 10.1113/EP085031Deuchars S.A. How sympathetic are your spinal cord circuits? Exp Physiol. 2015;100(4):365-71. doi: 10.1113/EP085031Moon E.A., Goodchild A.K., Pilowsky P.M. Lateralisation of projections from the rostral ventrolateral medulla to sympathetic preganglionic neurons in the rat. Brain Res. 2002;929(2):181-90. doi: 10.1016/s0006-8993(01)03388-1Moon E.A., Goodchild A.K., Pilowsky P.M. Lateralisation of projections from the rostral ventrolateral medulla to sympathetic preganglionic neurons in the rat. Brain Res. 2002;929(2):181-90. doi: 10.1016/s0006-8993(01)03388-1Ranson R.N., Motawei K., Pyner S., Coote J.H. The paraventricular nucleus of the hypothalamus sends efferents to the spinal cord of the rat that closely appose sympathetic preganglionic neurones projecting to the stellate ganglion. Exp Brain Res. 1998;120(2):164-72. doi: 10.1007/s002210050390Ranson R.N., Motawei K., Pyner S., Coote J.H. The paraventricular nucleus of the hypothalamus sends efferents to the spinal cord of the rat that closely appose sympathetic preganglionic neurones projecting to the stellate ganglion. Exp Brain Res. 1998;120(2):164-72. doi: 10.1007/s002210050390Forehand C.J. Morphology of sympathetic preganglionic neurons in the neonatal rat spinal cord: an intracellular horseradish peroxidase study. J Comp Neurol. 1990;298(3):334-42. doi: 10.1002/cne.902980306Forehand C.J. Morphology of sympathetic preganglionic neurons in the neonatal rat spinal cord: an intracellular horseradish peroxidase study. J Comp Neurol. 1990;298(3):334-42. doi: 10.1002/cne.902980306Wu L., Chang H.H., Havton L.A. The soma and proximal dendrites of sympathetic preganglionic neurons innervating the major pelvic ganglion in female rats receive predominantly inhibitory inputs. Neuroscience. 2012;217:32-45. doi: 10.1016/j.neuroscience.2012.05.005Wu L., Chang H.H., Havton L.A. The soma and proximal dendrites of sympathetic preganglionic neurons innervating the major pelvic ganglion in female rats receive predominantly inhibitory inputs. Neuroscience. 2012;217:32-45. doi: 10.1016/j.neuroscience.2012.05.005Tang F.R., Tan C.K., Ling E.A. A comparative study by retrograde neuronal tracing and substance P immunohistochemistry of sympathetic preganglionic neurons in spontaneously hypertensive rats and Wistar-Kyoto rats. J Anat. 1995;186 ( Pt 1):197-207Tang F.R., Tan C.K., Ling E.A. A comparative study by retrograde neuronal tracing and substance P immunohistochemistry of sympathetic preganglionic neurons in spontaneously hypertensive rats and Wistar-Kyoto rats. J Anat. 1995;186 ( Pt 1):197-207Folkow B., Rubinstein E.H. Cardiovascular effects of acute and chronic stimulations of the hypothalamus defence area in the rat. Acta physiol. scand.1966; 68: 48 – 57Folkow B., Rubinstein E.H. Cardiovascular effects of acute and chronic stimulations of the hypothalamus defence area in the rat. Acta physiol. scand.1966; 68: 48 – 57Cудаков К.В. Эмоциональный стресс как ведущий фактор патогенеза артериальной гипертензии. Патол. физиол. экспер. терап. 1975; 1: 3-12.Sudakov K.V. Emotional stress as a leading factor of pathogenesis arterial hypertension. Pathol. physiol. exp. therap. 1975; 1: 3-12. In Russian.

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