Роль нейротрофического фактора мозга (BDNF) в физической активности (обзор)
DOI:
https://doi.org/10.33910/2687-0223-2022-4-2-124-133Ключевые слова:
мозг, нейротрофические факторы, BDNF, физическая активность, онтогенезАннотация
Обзор посвящен описанию одного из нейротрофических факторов мозга (BDNF) и его роли в физической активности человека. В работе говорится, что это белок, обнаруживаемый во многих областях мозга (например, кора гиппокамп). Приводятся доказательства участия этого белка в процессе развития (например, в дифференцировке клеток, их миграции к месту дислокации, росте дендритов, синаптогенезе), выживания и поддержания активности нейронов. Описываются механизмы синтеза BDNF. Показано, что этот нейротрофический фактор присутствует в клетка микроглии, лимфоцитах, моноцитах, в клетках скелетных мышц, он может преодолевать гематоэнцефалический барьер. Он является критическим компонентом в происхождении связанных с долговременной потенциацией изменений синапсов. BDNF помогает поддерживать высокий уровень концентрации и внимания и, возможно, играет роль в защите мозга от повреждений. Снижение уровня BDNF в крови у новорожденных может повышать риск развития РАС, СДВГ, посттравматического стрессового расстройства, биполярного расстройства. Процессы обучения и памяти связаны со структурными изменениями дендритных шипиков в гиппокампе, коре и миндалевидном теле, и BDNF модулирует количество и структуру дендритных шипиков как во время развития, так и во взрослом мозге. Физическая активность различными путями воздействует на повышение уровня BDNF. Один из таких механизмов связан с лактатом, который появляется вследствие физической активности человека, но одновременно представляет собой важнейшую сигнальную молекулу. Чем выше уровень лактата, тем выше уровень BDNF. Улучшение когнитивной функции после физической активности также объясняется высоким уровнем BDNF в сыворотке.
Библиографические ссылки
Aksu, S., Unlu, G., Kardesler, A. C. et al. (2018) Altered levels of brain-derived neurotrophic factor, proBDNF and tissue plasminogen activator in children with posttraumatic stress disorder. Psychiatry Research, vol. 268, pp. 478–483. https://doi.org/10.1016/j.psychres.2018.07.013 (In English)
Amidfar, M., de Oliveira, J., Kucharska, E. (2020) CREB and BDNF: Neurobiology and treatment of Alzheimer’s disease. Life Sciences, vol. 257, article 118020. https://doi.org/10.1016/j.lfs.2020.118020 (In English)
Andreska, T., Rauskolb, S., Schukraft, N. et al. (2020) Induction of BDNF expression in layer II/III and layer V neurons of the motor cortex is essential for motor learning. The Journal of Neuroscience, vol. 40, no. 33, pp. 6289–6308. https://doi.org/10.1523/jneurosci.0288-20.2020 (In English)
Barbosa, A.G., Pratesi, R., Paz, G. S. C. et al. (2020) Assessment of BDNF serum levels as a diagnostic marker in children with autism spectrum disorder. Scientific Reports, vol. 10, article 17348. https://doi.org/10.1038/s41598-020-74239-x (In English)
Bathina, S., Das, U. N. (2015) Brain-derived neurotrophic factor and its clinical implications. Archives of Medical Science, vol. 11, no. 6, pp. 1164–1178. https://doi.org/10.5114/aoms.2015.56342 (In English)
Baykara, B., Koc, D., Resmi, H. et al. (2020) Brain-derived neurotrophic factor in bipolar disorder: Associations with age at onset and illness duration. Progress in Neuro-Psychopharmacology and Biological Psychiatry, vol. 108, article 10075. https://doi.org/10.1016/j.pnpbp.2020.110075 (In English)
Beltran-Valls, M. R., Adelantado-Renau, M., Moliner-Urdiales, D. (2018) Association between objectively measured physical activity and plasma BDNF in adolescents: DADOS study. Journal of Molecular Neuroscience, vol. 65, pp. 467–471. https://doi.org/10.1007/s12031-018-1122-2 (In English)
Brattico, E., Bonetti, L., Ferretti, G. et al. (2021) Putting cells in motion: Advantages of endogenous boosting of BDNF production. Cells, vol. 10, no. 1, article 183. https://doi.org/10.3390/cells10010183 (In English)
Brigadski, T., Leßmann, V. (2020) The physiology of regulated BDNF release. Cell and Tissue Research, vol. 382, pp. 15–45. https://doi.org/10.1007/s00441-020-03253-2 (In English)
Caffino, L., Mottarlini, F., Fumagalli, F. (2020) Born to protect: Leveraging BDNF against cognitive deficit in Alzheimer’s disease. CNS Drugs, vol. 34, pp. 281–297. https://doi.org/10.1007/s40263-020-00705-9 (In English)
Camuso, S., Rosa, P., Fiorenza, M. T., Canterini, S. (2022) Pleiotropic effects of BDNF on the cerebellum and hippocampus: Implications for neurodevelopmental disorders. Neurobiology of Disease, vol. 163. article 105606. https://doi.org/10.1016/j.nbd.2021.105606 (In English)
Chaddock-Heyman, L., Erickson, K. I., Holtrop, J. L. et al. (2014) Aerobic fitness is associated with greater white matter integrity in children. Frontiers in Human Neuroscience, vol. 8, article 584. https://doi.org/10.3389/fnhum.2014.00584 (In English)
Chau, C. M. Y., Cepeda, I. L., Devlin, A. M. (2017) The Val66Met brain-derived neurotrophic factor gene variant interacts with early pain exposure to predict cortisol dysregulation in 7-year-old children born very preterm: Implications for cognition. Neuroscience, vol. 342, pp. 188–199. https://doi.org/10.1016/j.neuroscience.2015.08.044 (In English)
Choi, S. H., Bylykbashi, E., Chatila, Z. K. et al. (2018) Combined adult neurogenesis and BDNF mimic exercise effects on cognition in an Alzheimer’s mouse model. Science, vol. 361 (6406), article eaan8821. https://doi.org/10.1126/science.aan8821 (In English)
De Azevedo, K. P. M., de Oliveira, V. H., de Medeiros G. C. B. S. et al. (2020) The Effects of exercise on BDNF levels in adolescents: A systematic review with meta-analysis. International Journal of Environmental Research and Public Health, vol. 17, no. 17, article 6056. https://doi.org/10.3390/ijerph17176056 (In English)
Delmotte, Q., Diabira, D., Belaidouni, Y. et al. (2020) Sonic hedgehog signaling agonist (sag) triggers BDNF secretion and promotes the maturation of GABAergic networks in the postnatal rat hippocampus. Frontiers in Cellular Neuroscience, vol. 14, article 98. https://doi.org/10.3389/fncel.2020.00098 (In English)
Dinoff, A., Herrmann, N., Swardfager, W., Lanctôt, K. L. (2017) The effect of acute exercise on blood concentrations of brain-derived neurotrophic factor in healthy adults: A meta-analysis. European Journal of Neuroscience, vol. 46, no. 1, pp. 1635–1646. https://doi.org/10.1111/ejn.13603 (In English)
Di Raimondo, D., Rizzo, G., Musiari, G. et al. (2020) Role of regular physical activity in neuroprotection against acute ischemia. International Journal of Molecular Sciences, vol. 21, no. 23, article 9086. https://doi.org/10.3390/ijms21239086 (In English)
Dougherty, K. D., Dreyfus, C. F., Black, I. B. (2000) Brain-derived neurotrophic factor in astrocytes, oligodendrocytes, and microglia/macrophages after spinal cord injury. Neurobiology of Disease, vol. 7, no. 6, pp. 574–585. https://doi.org/10.1006/nbdi.2000.0318 (In English)
Egan, M. F., Kojima, M., Callicott, J. H. et al. (2003) The BDNF val66met polymorphism affects activity-dependent secretion of BDNF and human memory and hippocampal function. Cell, vol. 112, no. 2, pp. 257–269. https://doi.org/10.1016/s0092-8674(03)00035-7 (In English)
El Hayek, L., Khalifeh, M., Zibara, V. et al. (2019) Lactate mediates the effects of exercise on learning and memory through SIRT1-dependent activation of hippocampal brain-derived neurotrophic factor (BDNF). The Journal of Neuroscience, vol. 39, no. 13, pp. 2369–2382. https://doi.org/10.1523/JNEUROSCI.1661-18.2019 (In English)
Erickson, K. I., Leckie, R. L., Weinstein, A. M. (2014) Physical activity, fitness, and gray matter volume. Neurobiology of Aging, vol. 35, suppl. 2, pp. 20–28. https://doi.org/10.1016/j.neurobiolaging.2014.03.034 (In English)
Erickson, K. I., Prakash, R. S., Voss, M. W. et al. (2010) Brain-derived neurotrophic factor is associated with age-related decline in hippocampal volume. Journal of Neuroscience, vol. 30, no. 15, pp. 5368–5375. https://doi.org/10.1523/jneurosci.6251-09.2010 (In English)
Ferreira, A. F. B., Real, C. C., Rodrigues, A. C. et al. (2011) Short-term, moderate exercise is capable of inducing structural, BDNF-independent hippocampal plasticity. Brain Research, vol. 1425, pp. 111–122. https://doi.org/10.1016/j.brainres.2011.10.004 (In English)
Frühauf-Perez, P. K., Temp, F. R., Pillat, M. M. et al. (2018) Spermine protects from LPS-induced memory deficit via BDNF and TrkB activation. Neurobiology of Learning and Memory, vol. 149, pp. 135–143. https://doi.org/10.1016/j.nlm.2018.02.012 (In English)
Greenberg, M. E., Xu, B., Lu, B., Hempstead, B. L. (2009) New insights in the biology of BDNF synthesis and release: Implications in CNS function. Journal of Neuroscience, vol. 29, no. 41, pp. 12764–12767. https://doi.org/10.1523/JNEUROSCI.3566-09.2009 (In English)
Goda, A., Ohgi, S., Kinpara, K. et al. (2013) Changes in serum BDNF levels associated with moderate-intensity exercise in healthy young Japanese men. SpringerPlus, vol. 2, article 678. https://doi.org/10.1186/2193-1801-2-678. (In English)
Griffin, E. W., Mullally, S., Foley, C. et al. (2011) Aerobic exercise improves hippocampal function and increases BDNF in the serum of young adult males. Physiology & Behavior, vol. 104, no. 5, pp. 934–941. https://doi.org/10.1016/j.physbeh.2011.06.005 (In English)
Griguoli, M., Cherubini, E. (2017) Early correlated network activity in the hippocampus: Its putative role in shaping neuronal circuits. Frontiers in Cellular Neuroscience, vol. 11, article 255. https://doi.org/10.3389/fncel.2017.00255 (In English)
Harward, S. C., Hedrick, N. G., Hall, C. E. et al. (2016) Autocrine BDNF–TrkB signalling within a single dendritic spine. Nature, vol. 538(7623), pp. 99–103. https://doi.org/10.1038/nature19766 (In English).
Hing, B., Sathyaputri, L., Potash, J. B. (2017) A comprehensive review of genetic and epigenetic mechanisms that regulate BDNF expression and function with relevance to major depressive disorder. American Journal of Medical Genetics. Part B: Neuropsychiatric Genetics, vol. 177, no. 2, pp. 143–167. https://doi.org/10.1002/ajmg.b.32616 (In English)
Hwang, J., Brothers, R. M., Castelli, D. M. et al. (2016) Acute high-intensity exercise-induced cognitive enhancement and brain-derived neurotrophic factor in young, healthy adults. Neuroscience Letters, vol. 630, pp. 247–253. https://doi.org/10.1016/j.neulet.2016.07.033 (In English)
Jodeiri Farshbaf, M., Ghaedi, K., Megraw, T. L. et al. (2015) Does PGC1α/FNDC5/BDNF elicit the beneficial effects of exercise on neurodegenerative disorders? NeuroMolecular Medicine, vol. 18, no. 1, pp. 1–15. https://doi.org/10.1007/s12017-015-8370-x (In English)
Johnstone, A., Mobley, W. (2020) Local TrkB signaling: themes in development and neural plasticity. Cell and Tissue Research, vol. 382 (1), pp. 101–111. https://doi.org/10.1007/s00441-020-03278-7 (In English)
Kishi, T., Yoshimura, R., Ikuta, T., Iwata, N. (2018) Brain-derived neurotrophic factor and major depressive disorder: Evidence from meta-analyses. Frontiers in Psychiatry, vol. 8, article 309. https://doi.org/10.3389/fpsyt.2017.00308 (In English)
Klein, A. B., Williamson, R., Santini, M. A. et al. (2010) Blood BDNF concentrations reflect brain-tissue BDNF levels across species. The International Journal of Neuropsychopharmacology, vol. 14, no. 3, pp. 347–353. https://doi.org/10.1017/s1461145710000738 (In English)
Kowiański, P., Lietzau, G., Czuba, E. et al. (2017) BDNF: A key factor with multipotent impact on brain signaling and synaptic plasticity. Cellular and Molecular Neurobiology, vol. 38, no. 3, pp. 579–593. https://doi.org/10.1007/s10571-017-0510-4 (In English)
Li, A., Jing, D., Dellarco, D. V. et al. (2021) Role of BDNF in the development of an OFC-amygdala circuit regulating sociability in mouse and human. Molecular Psychiatry, vol. 26, pp. 955–973. https://doi.org/10.1038/s41380-019-0422-4 (In English)
Lin, P.-Y., Kavalali, E. T., Monteggia, L. M. (2018) Genetic dissection of presynaptic and postsynaptic BDNF-TrkB signaling in synaptic efficacy of CA3-CA1 synapses. Cell Reports, vol. 24, no. 6, pp. 1550–1561. https://doi.org/10.1016/j.celrep.2018.07.020 (In English)
Liu, Y.-K., Gao, H., Jin, S.-B. et al. (2021) Association of neonatal blood levels of brain-derived neurotrophic factor with development of autism spectrum disorder: a systematic review and meta-analysis. World Journal of Pediatrics, vol. 17, no. 2, pp. 164–170. https://doi.org/10.1007/s12519-021-00415-2. (In English)
Lombardi, A., Jedlicka, P., Luhmann, H. J., Kilb, W. (2018) Giant depolarizing potentials trigger transient changes in the intracellular cl- concentration in ca3 pyramidal neurons of the immature mouse hippocampus. Frontiers in Cellular Neuroscience, vol. 12, article 420. https://doi.org/10.3389/fncel.2018.00420 (In English)
Luine, V., Frankfurt, M. (2013) Interactions between estradiol, BDNF and dendritic spines in promoting memory. Neuroscience, vol. 239, pp. 34–45. https://doi.org/10.1016/j.neuroscience.2012.10.019 (In English)
Luo, L., Jiang, X., Cao, G. et al. (2020) Association between BDNF gene polymorphisms and attention deficit hyperactivity disorder in school-aged children in Wuhan, China. Journal of Affective Disorders, vol. 264, pp. 304–309. https://doi.org/10.1016/j.jad.2020.01.017 (In English)
Martínez-Díaz, I. C., Escobar-Muñoz, M. C., Carrasco, L. (2020) Acute effects of high-intensity interval training on brain-derived neurotrophic factor, cortisol and working memory in physical education college students. International Journal of Environmental Research and Public Health, vol. 17, no. 21, article 8216. https://doi.org/10.3390/ijerph17218216 (In English)
Melkonian, E. A., Schury, M. P. (2021) Biochemistry, Anaerobic Glycolysis. Treasure Island: StatPearls Publ. [Online]. Available at: https://www.ncbi.nlm.nih.gov/books/NBK546695/ (accessed 11.06.2022). (In English)
Miao, Z., Wang, Y., Sun, Z. (2020) The relationships between stress, mental disorders, and epigenetic regulation of BDNF. International Journal of Molecular Sciences, vol. 21, no. 4, article 1375. https://doi.org/10.3390/ijms21041375 (In English)
Miguez, M. J., Bueno, D., Espinoza, L. et al. (2020) Among adolescents, BDNF and Pro-BDNF Lasting changes with alcohol use are stage specific. Neural Plasticity, vol. 2020, article 3937627. https://doi.org/10.1155/2020/3937627 (In English)
Mizoguchi, Y., Yao, H., Imamura, Y. et al. (2020) Lower brain-derived neurotrophic factor levels are associated with age-related memory impairment in community-dwelling older adults: The Sefuri study. Scientific Reports, vol. 10, article. 16442. https://doi.org/10.1038/s41598-020-73576-1 (In English)
Moore, D., Loprinzi, P. D. (2020) Exercise influences episodic memory via changes in hippocampal neurocircuitry and long-term potentiation. European Journal of Neuroscience, vol. 54, no. 8, pp. 6960–6971. https://doi.org/10.1111/ejn.14728 (In English)
Müller, P., Duderstadt, Y., Lessmann, V., Müller, N. G. (2020) Lactate and BDNF: Key mediators of exercise induced neuroplasticity? Journal of Clinical Medicine, vol. 9, no. 4, article 1136. https://doi.org/10.3390/jcm9041136 (In English)
Murakami, S., Imbe, H., Morikawa, Y. et al. (2005) Chronic stress, as well as acute stress, reduces BDNF mRNA expression in the rat hippocampus but less robustly. Neuroscience Research, vol. 53, no. 2, pp. 129–139. https://doi.org/10.1016/j.neures.2005.06.008 (In English)
Niculescu, D., Michaelsen-Preusse, K., Güner, Ü. et al. (2018) A BDNF-mediated push-pull plasticity mechanism for synaptic clustering. Cell Reports, vol. 24, no. 8, pp. 2063–2074. https://doi.org/10.1016/j.celrep.2018.07.073 (In English)
Numakawa, T., Odaka, H., Adachi, N. (2018) Actions of brain-derived neurotrophin factor in the neurogenesis and neuronal function, and its involvement in the pathophysiology of brain diseases. International Journal of Molecular Sciences, vol. 19, no. 11, article 3650. https://doi.org/10.3390/ijms19113650 (In English).
Orefice, L. L., Shih, C.-C., Xu, H. et al. (2016) Control of spine maturation and pruning through proBDNF synthesized and released in dendrites. Molecular and Cellular Neuroscience, vol. 71, pp. 66–79. https://doi.org/10.1016/j.mcn.2015.12.010 (In English)
Sakata, K., Woo, N. H., Martinowich, K. et al. (2009) Critical role of promoter IV-driven BDNF transcription in GABAergic transmission and synaptic plasticity in the prefrontal cortex. Proceedings of the National Academy of Sciences, vol. 106, no. 14, pp. 5942–5947. https://doi.org/10.1073/pnas.0811431106 (English)
Sbrini, G., Brivio, P., Fumagalli, M. et al. (2020) Centella asiatica L. phytosome improves cognitive performance by promoting BDNF expression in rat prefrontal cortex. Nutrients, vol. 12, no. 2, article 355. https://doi.org/10.3390/nu12020355 (In English)
Sheikhzadeh, F., Etemad, A., Khoshghadam, S. et al. (2015) Hippocampal BDNF content in response to short- and long-term exercise. Neurological Sciences, vol. 36, pp. 1163–1166. https://doi.org/10.1007/s10072-015-2208-z (In English)
Schiffer, T., Schulte, S., Hollmann, W. et al. (2008) Effects of strength and endurance training on brain-derived neurotrophic factor and insulin-like growth factor 1 in humans. Hormone and Metabolic Research, vol. 41, no.3, pp. 250–254. https://doi.org/10.1055/s-0028-1093322 (In English)
Skogstrand, K., Hagen, C. M., Borbye-Lorenzen, N. et al. (2019) Reduced neonatal brain-derived neurotrophic factor is associated with autism spectrum disorders. Translational Psychiatry, vol. 9, no. 1, article 252. https://doi.org/10.1038/s41398-019-0587-2 (In English)
Skriver, K., Roig, M., Lundbye-Jensen, J. et al. (2014) Acute exercise improves motor memory: Exploring potential biomarkers. Neurobiology of Learning and Memory, vol. 116, pp. 46–58. https://doi.org/10.1016/j.nlm.2014.08.004 (In English)
Sleiman, S. F., Henry, J., Al-Haddad, R. et al. (2016) Exercise promotes the expression of brain derived neurotrophic factor (BDNF) through the action of the ketone body β-hydroxybutyrate. eLife, vol. 5, article 15092. https://doi.org/10.7554/elife.15092 (In English)
Toh, Y. L., Ng, T., Tan, M. et al. (2018) Impact of brain-derived neurotrophic factor genetic polymorphism on cognition: A systematic review. Brain and Behavior, vol. 8, no. 7, article e01009. https://doi.org/10.1002/brb3.1009 (In English)
Vaynman, S., Ying, Z., Gomez-Pinilla, F. (2004) Hippocampal BDNF mediates the efficacy of exercise on synaptic plasticity and cognition. European Journal of Neuroscience, vol. 20, no. 10, pp. 2580–2590. https://doi.org/10.1111/j.1460-9568.2004.03720.x (In English)
Voss, M. W., Chaddock, L., Kim, J. S. et al. (2011) Aerobic fitness is associated with greater efficiency of the network underlying cognitive control in preadolescent children. Neuroscience, vol. 199, pp. 166–176. https://doi.org/10.1016/j.neuroscience.2011.10.009 (In English)
Walsh, E. I., Smith, L., Northey, J. et al. (2020) Towards an understanding of the physical activity-BDNF-cognition triumvirate: A review of associations and dosage. Ageing Research Reviews, vol. 60, article 101044. https://doi.org/10.1016/j.arr.2020.101044 (In English)
Walsh, J. J., Tschakovsky, M. E. (2018) Exercise and circulating BDNF: Mechanisms of release and implications for the design of exercise interventions. Applied Physiology, Nutrition, and Metabolism, vol. 43, no. 11, pp. 1095–1104. https://doi.org/10.1139/apnm-2018-0192 (In English)
Wang, L.-J., Wu, C. C., Lee, M.-J. et al. (2019) Peripheral brain-derived neurotrophic factor and contactin-1 levels in patients with attention-deficit/hyperactivity disorder. Journal of Clinical Medicine, vol. 8, no. 9, article 1366. https://doi.org/10.3390/jcm8091366 (In English)
Wiegert, J. S., Oertner, T. G. (2013) Long-term depression triggers the selective elimination of weakly integrated synapses. Proceedings of the National Academy of Sciences, vol. 110, no. 47, pp. E4510–E4519. https://doi.org/10.1073/pnas.1315926110 (In English)
Winnubst, J., Cheyne, J. E., Niculescu, D., Lohmann, C. (2015) Spontaneous activity drives local synaptic plasticity in vivo. Neuron, vol. 87, no. 2, pp. 399–410. https://doi.org/10.1016/j.neuron.2015.06.029 (In English)
Wrann, C. D., White, J. P., Salogiannnis, J. et al. (2013) Exercise induces hippocampal BDNF through a PGC-1α/ FNDC5 pathway. Cell Metabolism, vol. 18, no. 5, pp. 649–659. https://doi.org/10.1016/j.cmet.2013.09.008 (In English)
Yoshinori, F., Kazushige, I., Mitsugu, H. et al. (2021) Higher serum brain-derived neurotrophic factor levels are associated with a lower risk of cognitive decline: a 2-year follow up study in community-dwelling older adults. Frontiers Behavioral Neuroscience, vol. 15, article 641608. https://doi.org/10.3389/fnbeh.2021.641608 (In English)
Zagrebelsky, M., Tacke, C., Korte, M. (2020) BDNF signaling during the lifetime of dendritic spines. Cell and Tissue Research, vol. 382, pp. 185–199. https://doi.org/10.1007/s00441-020-03226-5 (In English)
Zhang, J., Luo, W., Li, Q. et al. (2018) Peripheral brain-derived neurotrophic factor in attention-deficit/hyperactivity disorder: A comprehensive systematic review and meta-analysis. Journal of Affective Disorders, vol. 227, pp. 298–304. https://doi.org/10.1016/j.jad.2017.11.012 (In English)
Zhao, Z., Zhu, Z., Li, H. et al. (2022) Effects of traumatic stress in adolescence on PTSD-like behaviors, dendrite development, and H3K9me2/BDNF expression in the amygdala of male rats. Journal of Affective Disorders, vol. 296, pp. 388–399. https://doi.org/10.1016/j.jad.2021.09.101 (In English)
Загрузки
Опубликован
Выпуск
Раздел
Лицензия
Copyright (c) 2022 Сутормина Надежда Владимировна
Это произведение доступно по лицензии Creative Commons «Attribution-NonCommercial» («Атрибуция — Некоммерческое использование») 4.0 Всемирная.
Автор предоставляет материалы на условиях публичной оферты и лицензии CC BY-NC 4.0. Эта лицензия позволяет неограниченному кругу лиц копировать и распространять материал на любом носителе и в любом формате, но с обязательным указанием авторства и только в некоммерческих целях. После публикации все статьи находятся в открытом доступе.
Авторы сохраняют авторские права на статью и могут использовать материалы опубликованной статьи при подготовке других публикаций, а также пользоваться печатными или электронными копиями статьи в научных, образовательных и иных целях. Право на номер журнала как составное произведение принадлежит издателю.
