<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.3 20210610//EN" "JATS-journalpublishing1-3.dtd">
<article article-type="research-article" dtd-version="1.3" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xml:lang="ru"><front><journal-meta><journal-id journal-id-type="publisher-id">actabiomedica</journal-id><journal-title-group><journal-title xml:lang="ru">Acta Biomedica Scientifica</journal-title><trans-title-group xml:lang="en"><trans-title>Acta Biomedica Scientifica</trans-title></trans-title-group></journal-title-group><issn pub-type="ppub">2541-9420</issn><issn pub-type="epub">2587-9596</issn><publisher><publisher-name>Scientific Centre for Family Health and Human Reproduction Problems</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.29413/ABS.2026-11.4.13</article-id><article-id custom-type="elpub" pub-id-type="custom">actabiomedica-6328</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="ru"><subject>МОРФОЛОГИЯ, ФИЗИОЛОГИЯ И ПАТОФИЗИОЛОГИЯ</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>MORPHOLOGY, PHYSIOLOGY AND PATHOPHYSIOLOGY</subject></subj-group></article-categories><title-group><article-title>Распределение βIII-тубулина в обонятельных луковицах людей разного пола при старении</article-title><trans-title-group xml:lang="en"><trans-title>βIII-tubulin distribution in the human olfactory bulb: effects of aging and sex</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0009-0006-8047-7336</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Мальков</surname><given-names>А. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Malkov</surname><given-names>A. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Мальков Артем Александрович – лаборант-исследователь лаборатории развития нервной системы НИИ морфологии человека им. акад. А.П. Авцына ФГБНУ «Российский научный центр хирургии им. акад. Б.В. Петровского»; студент 5 курса ФГАОУ ВО «Первый Московский государственный медицинский университет имени И.М. Сеченова» Минздрава России (Сеченовский Университет)</p><p>117418, г. Москва, ул. Цюрупы, д. 3,</p><p>119048, г. Москва, ул. Трубецкая, д. 8, стр. 2</p></bio><bio xml:lang="en"><p>Artyom A. Malkov – Laboratory Assistant, Laboratory of Nervous System Development, Avtsyn Research Institute of Human Morphology of the Petrovsky National Research Centre of Surgery; student of the 5th graduate year I.M. Sechenov First Moscow State Medical University (Sechenov University)</p><p>Tsyurupa str., 3, Moscow 117418, </p><p>Trubetskaya str., 8-2, Moscow 119048</p></bio><email xlink:type="simple">i@aamalkov.ru</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-0515-8275</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Прощина</surname><given-names>А. Е.</given-names></name><name name-style="western" xml:lang="en"><surname>Proshchina</surname><given-names>A. E.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Прощина Александра Евгеньевна – доктор биологических наук, доцент, ведущий научный сотрудник лаборатории развития нервной системы </p><p>117418, г. Москва, ул. Цюрупы, д. 3</p></bio><bio xml:lang="en"><p>Alexandra E. Proshchina – Dr. Sci. (Biol.), assistant professor, leading researcher, Laboratory of Nervous System Development</p><p>Tsyurupa str., 3, Moscow 117418</p></bio><email xlink:type="simple">proshchina@yandex.ru</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-9692-3616</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Кривова</surname><given-names>Ю. С.</given-names></name><name name-style="western" xml:lang="en"><surname>Krivova</surname><given-names>Yu. S.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Кривова Юлия Сергеевна – кандидат биологических наук, старший научный сотрудник лаборатории развития нервной системы </p><p>117418, г. Москва, ул. Цюрупы, д. 3</p></bio><bio xml:lang="en"><p>Yuliya S. Krivova – Cand. Sci. (Biol.), senior researcher, Laboratory of Nervous System Development</p><p>Tsyurupa str., 3, Moscow 117418</p></bio><email xlink:type="simple">homulkina@rambler.ru</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-1163-4132</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Харламова</surname><given-names>А. С.</given-names></name><name name-style="western" xml:lang="en"><surname>Kharlamova</surname><given-names>A. S.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Харламова Анастасия Сергеевна – кандидат биологических наук, старший научный сотрудник лаборатории развития нервной системы </p><p>117418, г. Москва, ул. Цюрупы, д. 3</p></bio><bio xml:lang="en"><p>Anastasia S. Kharlamova – Cand. Sci. (Biol.), senior researcher, Laboratory of Nervous System Development</p><p>Tsyurupa str., 3, Moscow 117418</p></bio><email xlink:type="simple">grossulyar@gmail.com</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-9285-7241</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Годовалова</surname><given-names>О. С.</given-names></name><name name-style="western" xml:lang="en"><surname>Godovalova</surname><given-names>O. S.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Годовалова Ольга Сергеевна – кандидат биологических наук, старший научный сотрудник лаборатории развития нервной системы </p><p>117418, г. Москва, ул. Цюрупы, д. 3</p></bio><bio xml:lang="en"><p>Olga S. Godovalova – Cand. Sci. (Biol.), senior researcher, Laboratory of Nervous System Development</p><p>Tsyurupa str., 3, Moscow 117418</p></bio><email xlink:type="simple">asinello@gmail.com</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-7997-8161</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Гулимова</surname><given-names>В. И.</given-names></name><name name-style="western" xml:lang="en"><surname>Gulimova</surname><given-names>V. I.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Гулимова Виктория Игоревна – кандидат биологических наук, старший научный сотрудник лаборатории развития нервной системы </p><p>117418, г. Москва, ул. Цюрупы, д. 3</p></bio><bio xml:lang="en"><p>Victoria I. Gulimova – Cand. Sci. (Biol.), senior researcher, Laboratory of Nervous System Development</p><p>Tsyurupa str., 3, Moscow 117418</p></bio><email xlink:type="simple">gulimova@yandex.ru</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-3580-1492</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Бучака</surname><given-names>А. С.</given-names></name><name name-style="western" xml:lang="en"><surname>Buchaka</surname><given-names>A. S.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Бучака Антон Сергеевич – исполняющий обязанности руководителя референс-центра инфекционной и вирусной онкопатологии </p><p>117418, г. Москва, ул. Цюрупы, д. 3</p></bio><bio xml:lang="en"><p>Anton S. Buchaka – acting leader, Reference Center for Infectious and Viral Oncopathology</p><p>Tsyurupa str., 3, Moscow 117418</p></bio><email xlink:type="simple">abpao62@yandex.ru</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-6844-700X</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Сонин</surname><given-names>Г. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Sonin</surname><given-names>G. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Сонин Глеб Александрович – младший научный сотрудник лаборатории развития нервной системы </p><p>117418, г. Москва, ул. Цюрупы, д. 3</p></bio><bio xml:lang="en"><p>Gleb A. Sonin – junior researcher, Laboratory of Nervous System Development</p><p>Tsyurupa str., 3, Moscow 117418</p></bio><email xlink:type="simple">glebs0nin@yandex.ru</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0009-0005-9196-4888</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Грушецкая</surname><given-names>Е. О.</given-names></name><name name-style="western" xml:lang="en"><surname>Grushetskaya</surname><given-names>E. O.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Грушецкая Евгения Олеговна – лаборант-исследователь лаборатории развития нервной системы </p><p>117418, г. Москва, ул. Цюрупы, д. 3</p></bio><bio xml:lang="en"><p>Evgenia O. Grushetskaya  – laboratory assistant, Laboratory of Nervous System Development</p><p>Tsyurupa str., 3, Moscow 117418</p></bio><email xlink:type="simple">050101j@mail.ru</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-6595-5305</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Савельев</surname><given-names>С. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Saveliev</surname><given-names>S. V.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Савельев Сергей Вячеславович – доктор биологических наук, профессор, заведующий лабораторией развития нервной системы </p><p>117418, г. Москва, ул. Цюрупы, д. 3</p></bio><bio xml:lang="en"><p>Sergey V. Saveliev – Dr. Sci. (Biol.), Professor, Head of Laboratory of Nervous System Development</p><p>Tsyurupa str., 3, Moscow 117418</p></bio><email xlink:type="simple">braincase@yandex.ru</email><xref ref-type="aff" rid="aff-2"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru"><institution>Научно-исследовательский институт морфологии человека им. акад. А.П. Авцына ФГБНУ «Российский научный центр хирургии им. акад. Б.В. Петровского»;&#13;
ФГАОУ ВО Первый Московский государственный медицинский университет им. И.М. Сеченова Минздрава России</institution><country>Россия</country></aff><aff xml:lang="en"><institution>Avtsyn Research Institute of Human Morphology of the Petrovsky National Research Center of Surgery;&#13;
I.M. Sechenov First Moscow State Medical University (Sechenov University)</institution><country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-2"><aff xml:lang="ru"><institution>Научно-исследовательский институт морфологии человека им. акад. А.П. Авцына ФГБНУ «Российский научный центр хирургии им. акад. Б.В. Петровского»</institution><country>Россия</country></aff><aff xml:lang="en"><institution>Avtsyn Research Institute of Human Morphology of the Petrovsky National Research Center of Surgery</institution><country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2026</year></pub-date><pub-date pub-type="epub"><day>17</day><month>10</month><year>2026</year></pub-date><volume>11</volume><issue>4</issue><fpage>126</fpage><lpage>135</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Мальков А.А., Прощина А.Е., Кривова Ю.С., Харламова А.С., Годовалова О.С., Гулимова В.И., Бучака А.С., Сонин Г.А., Грушецкая Е.О., Савельев С.В., 2026</copyright-statement><copyright-year>2026</copyright-year><copyright-holder xml:lang="ru">Мальков А.А., Прощина А.Е., Кривова Ю.С., Харламова А.С., Годовалова О.С., Гулимова В.И., Бучака А.С., Сонин Г.А., Грушецкая Е.О., Савельев С.В.</copyright-holder><copyright-holder xml:lang="en">Malkov A.A., Proshchina A.E., Krivova Y.S., Kharlamova A.S., Godovalova O.S., Gulimova V.I., Buchaka A.S., Sonin G.A., Grushetskaya E.O., Saveliev S.V.</copyright-holder><license xml:lang="ru" license-type="creative-commons-attribution" xlink:href="https://creativecommons.org/licenses/by/4.0/" xlink:type="simple"><license-p>Данная работа распространяется под лицензией Creative Commons Attribution 4.0.</license-p></license><license xml:lang="en" license-type="creative-commons-attribution" xlink:href="https://creativecommons.org/licenses/by/4.0/" xlink:type="simple"><license-p>This work is licensed under a Creative Commons Attribution 4.0 License.</license-p></license></permissions><self-uri xlink:href="https://www.actabiomedica.ru/jour/article/view/6328">https://www.actabiomedica.ru/jour/article/view/6328</self-uri><abstract><sec><title>Обоснование</title><p>Обоснование. Многочисленные исследования связывают нарушения обоняния с ускоренным снижением когнитивных функций при нейродегенеративных заболеваниях, однако механизм участия в этих процессах первичного обонятельного центра, обонятельной луковицы, остается неясным. βIII-тубулин является одним из наиболее часто применяемых нейрональных маркеров, но данные о его распределении в обонятельной луковице ограничены.</p></sec><sec><title>Цель работы</title><p>Цель работы. Провести анализ распределения βIII-тубулина в обонятельных луковицах мужчин и женщин при старении.</p></sec><sec><title>Методы</title><p>Методы. Исследование выполнено на аутопсийном материале. Обонятельные луковицы получены от 10 мужчин и 14 женщин пожилого (60–74 года, n = 11) и старческого (75–89 лет, n = 13) возрастов. Гистологические срезы окрашивали гематоксилином и эозином, по Нисслю и по Маллори для цитоархитектонического анализа. Иммуногистохимический анализ проводили с использованием первичных антител к нейронспецифическому βIII-тубулину. Выполняли морфометрический и статистический анализ.</p></sec><sec><title>Результаты</title><p>Результаты. Иммуногистохимическая реакция на βIII-тубулин выявлена в основном в слое вхождения волокон обонятельного нерва и в клубочковом слое. Были выявлены половые различия: обнаружено значимое (p = 0,006) преобладание площади реакции на βIII-тубулин на срезах обонятельных луковиц, полученных от женщин. Это совпадает с результатами исследований, показывающих, что у женщин в обонятельных луковицах почти в два раза больше нейронов, а способность к распознаванию запахов, как правило, лучше, чем у мужчин. Несмотря на тенденцию к уменьшению уровня βIII-тубулина с возрастом, статистически значимых отличий не выявлено.</p></sec><sec><title>Заключение</title><p>Заключение. Исследование выявило половую и возрастную изменчивость в распределении βIII-тубулина в обонятельной луковице человека. Поэтому необходим обязательный учет этих факторов при проведении функциональной диагностики обоняния.</p></sec></abstract><trans-abstract xml:lang="en"><sec><title>Background</title><p>Background. Numerous studies link olfactory impairment to accelerated cognitive decline in neurodegenerative diseases, yet the mechanism by which the olfactory bulb participates in these processes remains unclear. βIII-tubulin is one of the most commonly used neuronal markers, but data on its distribution in the olfactory bulb are limited.</p></sec><sec><title>Aim</title><p>Aim. To analyze the distribution of βIII-tubulin in the olfactory bulbs of men and women during aging.</p></sec><sec><title>Methods</title><p>Methods. The study was performed on autopsy material. Olfactory bulbs were obtained from 10 men and 14 women of elderly (60–74 years, n = 11) and senile (75–89 years n = 13) ages. Histological sections were stained with hematoxylin and eosin, Nissl stain, and Mallory’s trichrome for cytoarchitectonic analysis. Immunohistochemical analysis was performed using primary antibodies against neuron-specific βIII-tubulin. Morphometric and statistical analyses were carried out.</p></sec><sec><title>Results</title><p>Results. βIII-tubulin immunoreactivity was mainly detected in the olfactory nerve fiber layer and the glomerular layer. Analysis of sex differences revealed a significant (p = 0.006) predominance of the βIII-tubulin reaction area in olfactory bulb sections obtained from women. This aligns with research findings showing that women have nearly twice as many neurons in their olfactory bulbs and typically exhibit better odor identification ability compared to men. Despite a trend towards a decrease in βIII-tubulin levels with age, no statistically significant differences were found.</p></sec><sec><title>Conclusion</title><p>Conclusion. The study revealed sexand age-related variability in the distribution of βIII-tubulin in the human olfactory bulb. Therefore, these factors should be taken into account when performing functional olfactory diagnostics.</p></sec></trans-abstract><kwd-group xml:lang="ru"><kwd>обонятельная луковица</kwd><kwd>клубочковый слой</kwd><kwd>βIII-тубулин</kwd><kwd>старение</kwd><kwd>половая изменчивость</kwd></kwd-group><kwd-group xml:lang="en"><kwd>olfactory bulb</kwd><kwd>glomerular layer</kwd><kwd>βIII-tubulin</kwd><kwd>aging</kwd><kwd>sex-related variability</kwd></kwd-group><funding-group><funding-statement xml:lang="ru">Исследование выполнено в рамках государственного задания FURG-2025-0031 с использованием благотворительного пожертвования ООО «Компания Хеликон»</funding-statement></funding-group></article-meta></front><back><ref-list><title>References</title><ref id="cit1"><label>1</label><citation-alternatives><mixed-citation xml:lang="ru">Dan X, Wechter N, Gray S, Mohanty JG, Croteau DL, Bohr VA. Olfactory dysfunction in aging and neurodegenerative diseases. Ageing Res Rev. 2021; 70: 101416. doi: 10.1016/j.arr.2021.101416</mixed-citation><mixed-citation xml:lang="en">Dan X, Wechter N, Gray S, Mohanty JG, Croteau DL, Bohr VA. Olfactory dysfunction in aging and neurodegenerative diseases. Ageing Res Rev. 2021; 70: 101416. doi: 10.1016/j.arr.2021.101416</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Doty RL. Olfactory dysfunction in neurodegenerative diseases: is there a common pathological substrate? The Lancet Neurology. 2017; 16(6): 478–488. doi: 10.1016/s1474-4422(17)30123-0</mixed-citation><mixed-citation xml:lang="en">Doty RL. Olfactory dysfunction in neurodegenerative diseases: is there a common pathological substrate? The Lancet Neurology. 2017; 16(6): 478–488. doi: 10.1016/s1474-4422(17)30123-0</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Морозова С.В., Савватеева Д.М. Обонятельные расстройства у пациентов с нейродегенеративными и психическими заболеваниями. Русский медицинский журнал. 2014; 9: 673.</mixed-citation><mixed-citation xml:lang="en">Morozova SV, Savvateeva DM. Olfactory disorders in patients with neurodegenerative and mental diseases. Russian Medical Journal. 2014; 9: 673 (In Russ.).</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Mak E, Fought AJ, Wiste HJ, Przybelski SA, Reid RI, Schwarz CG, et al. Sex-Specific Associations of α-Synuclein Pathology with Tau Accumulation. JAMA Netw Open. 2026; 9(3): e260461. doi: 10.1001/jamanetworkopen.2026.0461</mixed-citation><mixed-citation xml:lang="en">Mak E, Fought AJ, Wiste HJ, Przybelski SA, Reid RI, Schwarz CG, et al. Sex-Specific Associations of α-Synuclein Pathology with Tau Accumulation. JAMA Netw Open. 2026; 9(3): e260461. doi: 10.1001/jamanetworkopen.2026.0461</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Kharlamova AS, Godovalova OS, Otlyga EG, Proshchina AE. Primary and secondary olfactory centres in human ontogeny. Neurosci Res. 2023; 190: 1–16. doi: 10.1016/j.neures.2022.12.005</mixed-citation><mixed-citation xml:lang="en">Kharlamova AS, Godovalova OS, Otlyga EG, Proshchina AE. Primary and secondary olfactory centres in human ontogeny. Neurosci Res. 2023; 190: 1–16. doi: 10.1016/j.neures.2022.12.005</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Bhatnagar KP, Kennedy RC, Baron G, Greenberg RA. Number of mitral cells and the bulb volume in the aging human olfactory bulb: a quantitative morphological study. Anat Rec. 1987; 218(1): 73–87. doi: 10.1002/ar.1092180112</mixed-citation><mixed-citation xml:lang="en">Bhatnagar KP, Kennedy RC, Baron G, Greenberg RA. Number of mitral cells and the bulb volume in the aging human olfactory bulb: a quantitative morphological study. Anat Rec. 1987; 218(1): 73–87. doi: 10.1002/ar.1092180112</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Meisami E, Mikhail L, Baim D, Bhatnagar KP. Human olfactory bulb: aging of glomeruli and mitral cells and a search for the accessory olfactory bulb. Ann N Y Acad Sci. 1998; 855: 708–715. doi: 10.1111/j.1749-6632.1998.tb10649.x</mixed-citation><mixed-citation xml:lang="en">Meisami E, Mikhail L, Baim D, Bhatnagar KP. Human olfactory bulb: aging of glomeruli and mitral cells and a search for the accessory olfactory bulb. Ann N Y Acad Sci. 1998; 855: 708–715. doi: 10.1111/j.1749-6632.1998.tb10649.x</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Sama-ul-Haq, Tahir M, Lone KP. Age and gender-related differences in mitral cells of olfactory bulb. J Coll Physicians Surg Pak. 2008; 18(11): 669–673.</mixed-citation><mixed-citation xml:lang="en">Sama-ul-Haq, Tahir M, Lone KP. Age and gender-related differences in mitral cells of olfactory bulb. J Coll Physicians Surg Pak. 2008; 18(11): 669–673.</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Kovács T. Mechanisms of olfactory dysfunction in aging and neurodegenerative disorders. Ageing Res Rev. 2004; 3(2): 215–232. doi: 10.1016/j.arr.2003.10.003</mixed-citation><mixed-citation xml:lang="en">Kovács T. Mechanisms of olfactory dysfunction in aging and neurodegenerative disorders. Ageing Res Rev. 2004; 3(2): 215–232. doi: 10.1016/j.arr.2003.10.003</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Son G, Jahanshahi A, Yoo SJ, Boonstra JT, Hopkins DA, Steinbusch HWM, et al. Olfactory neuropathology in Alzheimer’s disease: a sign of ongoing neurodegeneration. BMB Rep. 2021; 54(6): 295–304. doi: 10.5483/BMBRep.2021.54.6.055</mixed-citation><mixed-citation xml:lang="en">Son G, Jahanshahi A, Yoo SJ, Boonstra JT, Hopkins DA, Steinbusch HWM, et al. Olfactory neuropathology in Alzheimer’s disease: a sign of ongoing neurodegeneration. BMB Rep. 2021; 54(6): 295–304. doi: 10.5483/BMBRep.2021.54.6.055</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Shoff TA, Derbez-Morin M, Cai P, Julian RR. The microtubule nexus linking amyloid beta and tau: A simple and unifying theory for the underlying cause of Alzheimer’s disease. PNAS Nexus. 2026; 5(3): pgag034. doi: 10.1093/pnasnexus/pgag034</mixed-citation><mixed-citation xml:lang="en">Shoff TA, Derbez-Morin M, Cai P, Julian RR. The microtubule nexus linking amyloid beta and tau: A simple and unifying theory for the underlying cause of Alzheimer’s disease. PNAS Nexus. 2026; 5(3): pgag034. doi: 10.1093/pnasnexus/pgag034</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Lee VM, Pixley SK. Age and differentiation-related differences in neuron-specific tubulin immunostaining of olfactory sensory neurons. Brain Res Dev Brain Res. 1994; 83(2): 209–215. doi: 10.1016/0165-3806(94)00139-1</mixed-citation><mixed-citation xml:lang="en">Lee VM, Pixley SK. Age and differentiation-related differences in neuron-specific tubulin immunostaining of olfactory sensory neurons. Brain Res Dev Brain Res. 1994; 83(2): 209–215. doi: 10.1016/0165-3806(94)00139-1</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Santiago-Mujika E, Luthi-Carter R, Giorgini F, Mukaetova-Ladinska EB. Tubulin Isotypes and Posttranslational Modifications in Vascular Dementia and Alzheimer’s disease. J Alzheimers Dis Rep. 2022; 6(1): 739–748. doi: 10.3233/ADR-220068</mixed-citation><mixed-citation xml:lang="en">Santiago-Mujika E, Luthi-Carter R, Giorgini F, Mukaetova-Ladinska EB. Tubulin Isotypes and Posttranslational Modifications in Vascular Dementia and Alzheimer’s disease. J Alzheimers Dis Rep. 2022; 6(1): 739–748. doi: 10.3233/ADR-220068</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Lucas L, Tsoi PS, Quan MD, Choi KJ, Ferreon JC, Ferreon ACM. Tubulin transforms Tau and α-synuclein condensates from pathological to physiological. Nat Commun. 2026; 17(1): 3362. doi: 10.1038/s41467-026-69618-3</mixed-citation><mixed-citation xml:lang="en">Lucas L, Tsoi PS, Quan MD, Choi KJ, Ferreon JC, Ferreon ACM. Tubulin transforms Tau and α-synuclein condensates from pathological to physiological. Nat Commun. 2026; 17(1): 3362. doi: 10.1038/s41467-026-69618-3</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">von Hungen K, Chin RC, Baxter CF. Brain tubulin microheterogeneity in the mouse during development and aging. J Neurochem. 1981; 37(2): 511–514. doi: 10.1111/j.1471-4159.1981.tb00485.x</mixed-citation><mixed-citation xml:lang="en">von Hungen K, Chin RC, Baxter CF. Brain tubulin microheterogeneity in the mouse during development and aging. J Neurochem. 1981; 37(2): 511–514. doi: 10.1111/j.1471-4159.1981.tb00485.x</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Santiago-Mujika E, Luthi-Carter R, Giorgini F, Kalaria RN, Mukaetova-Ladinska EB. Tubulin and Tubulin Posttranslational Modifications in Alzheimer’s disease and Vascular Dementia. Front Aging Neurosci. 2021; 13: 730107. doi: 10.3389/fnagi.2021.730107</mixed-citation><mixed-citation xml:lang="en">Santiago-Mujika E, Luthi-Carter R, Giorgini F, Kalaria RN, Mukaetova-Ladinska EB. Tubulin and Tubulin Posttranslational Modifications in Alzheimer’s disease and Vascular Dementia. Front Aging Neurosci. 2021; 13: 730107. doi: 10.3389/fnagi.2021.730107</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Bédard A, Parent A. Evidence of newly generated neurons in the human olfactory bulb. Brain Res Dev Brain Res. 2004; 151(1–2): 159–168. doi: 10.1016/j.devbrainres.2004.03.021</mixed-citation><mixed-citation xml:lang="en">Bédard A, Parent A. Evidence of newly generated neurons in the human olfactory bulb. Brain Res Dev Brain Res. 2004; 151(1–2): 159–168. doi: 10.1016/j.devbrainres.2004.03.021</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Oliveira-Pinto AV, Santos RM, Coutinho RA, Oliveira LM, Santos GB, Alho AT, et al. Sexual dimorphism in the human olfactory bulb: females have more neurons and glial cells than males. PLoS One. 2014; 9(11): e111733. doi: 10.1371/journal.pone.0111733</mixed-citation><mixed-citation xml:lang="en">Oliveira-Pinto AV, Santos RM, Coutinho RA, Oliveira LM, Santos GB, Alho AT, et al. Sexual dimorphism in the human olfactory bulb: females have more neurons and glial cells than males. PLoS One. 2014; 9(11): e111733. doi: 10.1371/journal.pone.0111733</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Durante MA, Kurtenbach S, Sargi ZB, Harbour JW, Choi R, Kurtenbach S, et al. Single-cell analysis of olfactory neurogenesis and differentiation in adult humans. Nat Neurosci. 2020; 23(3): 323–326. doi: 10.1038/s41593-020-0587-9</mixed-citation><mixed-citation xml:lang="en">Durante MA, Kurtenbach S, Sargi ZB, Harbour JW, Choi R, Kurtenbach S, et al. Single-cell analysis of olfactory neurogenesis and differentiation in adult humans. Nat Neurosci. 2020; 23(3): 323–326. doi: 10.1038/s41593-020-0587-9</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Kunkhyen T, Brechbill TR, Berg SPR, Pothuri P, Rangel AN, Gupta A, et al. Cell type- and layer-specific plasticity of olfactory bulb interneurons following olfactory sensory neuron ablation. Sci Rep. 2024; 14(1): 17771. doi: 10.1038/s41598-024-68649-4</mixed-citation><mixed-citation xml:lang="en">Kunkhyen T, Brechbill TR, Berg SPR, Pothuri P, Rangel AN, Gupta A, et al. Cell type- and layer-specific plasticity of olfactory bulb interneurons following olfactory sensory neuron ablation. Sci Rep. 2024; 14(1): 17771. doi: 10.1038/s41598-024-68649-4</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Fitzek M, Patel PK, Solomon PD, Lin B, Hummel T, Schwob JE, et al. Integrated age-related immunohistological changes occur in human olfactory epithelium and olfactory bulb. J Comp Neurol. 2022; 530(12): 2154–2175. doi: 10.1002/cne.25325</mixed-citation><mixed-citation xml:lang="en">Fitzek M, Patel PK, Solomon PD, Lin B, Hummel T, Schwob JE, et al. Integrated age-related immunohistological changes occur in human olfactory epithelium and olfactory bulb. J Comp Neurol. 2022; 530(12): 2154–2175. doi: 10.1002/cne.25325</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Sorokowski P, Karwowski M, Misiak M, Marczak MK, Dziekan M, Hummel T, et al. Sex Differences in Human Olfaction: A Meta-Analysis. Front Psychol. 2019; 10: 242. doi: 10.3389/fpsyg.2019.00242</mixed-citation><mixed-citation xml:lang="en">Sorokowski P, Karwowski M, Misiak M, Marczak MK, Dziekan M, Hummel T, et al. Sex Differences in Human Olfaction: A Meta-Analysis. Front Psychol. 2019; 10: 242. doi: 10.3389/fpsyg.2019.00242</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Flores-Cuadrado A, Saiz-Sanchez D, Mohedano-Moriano A, Lamas-Cenjor E, Leon-Olmo V, Martinez-Marcos A, et al. Astrogliosis and sexually dimorphic neurodegeneration and microgliosis in the olfactory bulb in Parkinson’s disease. NPJ Parkinsons Dis. 2021; 7(1): 11. doi: 10.1038/s41531-020-00154-7</mixed-citation><mixed-citation xml:lang="en">Flores-Cuadrado A, Saiz-Sanchez D, Mohedano-Moriano A, Lamas-Cenjor E, Leon-Olmo V, Martinez-Marcos A, et al. Astrogliosis and sexually dimorphic neurodegeneration and microgliosis in the olfactory bulb in Parkinson’s disease. NPJ Parkinsons Dis. 2021; 7(1): 11. doi: 10.1038/s41531-020-00154-7</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Yeo BSY, Song HJJMD, Tan BKJ, Suresh A, Ho OTW, Chan JH, et al. Olfactory Impairment and Incident Cognitive Decline: A Systematic Review and Meta-Analysis. Int Forum Allergy Rhinol. 2025; 15(12): 1347–1358. doi: 10.1002/alr.23635</mixed-citation><mixed-citation xml:lang="en">Yeo BSY, Song HJJMD, Tan BKJ, Suresh A, Ho OTW, Chan JH, et al. Olfactory Impairment and Incident Cognitive Decline: A Systematic Review and Meta-Analysis. Int Forum Allergy Rhinol. 2025; 15(12): 1347–1358. doi: 10.1002/alr.23635</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Liu Q, Chen B, Wang Q, Xu D, Yang M, Lin G, et al. Sex differences in the relationship between olfactory and cognitive impairment among subjects with subjective cognitive decline and mild cognitive impairment. Biol Sex Differ. 2025; 16(1): 12. doi: 10.1186/s13293-025-00691-x</mixed-citation><mixed-citation xml:lang="en">Liu Q, Chen B, Wang Q, Xu D, Yang M, Lin G, et al. Sex differences in the relationship between olfactory and cognitive impairment among subjects with subjective cognitive decline and mild cognitive impairment. Biol Sex Differ. 2025; 16(1): 12. doi: 10.1186/s13293-025-00691-x</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Kamath V, Senjem ML, Spychalla AJ, Chen H, Palta P, Mosley TH, et al. The Neuroanatomic Correlates of Olfactory Identification Impairment in Healthy Older Adults and in Persons with Mild Cognitive Impairment. J Alzheimers Dis. 2022; 89(1): 233–245. doi: 10.3233/JAD-220228</mixed-citation><mixed-citation xml:lang="en">Kamath V, Senjem ML, Spychalla AJ, Chen H, Palta P, Mosley TH, et al. The Neuroanatomic Correlates of Olfactory Identification Impairment in Healthy Older Adults and in Persons with Mild Cognitive Impairment. J Alzheimers Dis. 2022; 89(1): 233–245. doi: 10.3233/JAD-220228</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Fatuzzo I, Niccolini GF, Zoccali F, Cavalcanti L, Bellizzi MG, Riccardi G, et al. Neurons, Nose, and Neurodegenerative Diseases: Olfactory Function and Cognitive Impairment. Int J Mol Sci. 2023; 24(3): 2117. doi: 10.3390/ijms24032117</mixed-citation><mixed-citation xml:lang="en">Fatuzzo I, Niccolini GF, Zoccali F, Cavalcanti L, Bellizzi MG, Riccardi G, et al. Neurons, Nose, and Neurodegenerative Diseases: Olfactory Function and Cognitive Impairment. Int J Mol Sci. 2023; 24(3): 2117. doi: 10.3390/ijms24032117</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Prem B, Liu DT, Boehme K, Maurer MT, Renner B, Mueller CA. Factors Associated with Persisting Olfactory Dysfunction After COVID-19. Clin Otolaryngol. 2025; 50(1): 53–61. doi: 10.1111/coa.14233</mixed-citation><mixed-citation xml:lang="en">Prem B, Liu DT, Boehme K, Maurer MT, Renner B, Mueller CA. Factors Associated with Persisting Olfactory Dysfunction After COVID-19. Clin Otolaryngol. 2025; 50(1): 53–61. doi: 10.1111/coa.14233</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Knight JE, Yoneda T, Lewis NA, Muniz-Terrera G, Bennett DA, Piccinin AM. Transitions between Mild Cognitive Impairment, Dementia, and Mortality: The Importance of Olfaction. J Gerontol a Biol Sci Med Sci. 2023; 78(7): 1284– 1291. doi: 10.1093/gerona/glad001</mixed-citation><mixed-citation xml:lang="en">Knight JE, Yoneda T, Lewis NA, Muniz-Terrera G, Bennett DA, Piccinin AM. Transitions between Mild Cognitive Impairment, Dementia, and Mortality: The Importance of Olfaction. J Gerontol a Biol Sci Med Sci. 2023; 78(7): 1284– 1291. doi: 10.1093/gerona/glad001</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Grillo F, Bruzzone M, Pigozzi S, Prosapio S, Migliora P, Fiocca R, et al. Immunohistochemistry on old archival paraffin blocks: is there an expiry date? J Clin Pathol. 2017; 70(11): 988–993. doi: 10.1136/jclinpath-2017-204387</mixed-citation><mixed-citation xml:lang="en">Grillo F, Bruzzone M, Pigozzi S, Prosapio S, Migliora P, Fiocca R, et al. Immunohistochemistry on old archival paraffin blocks: is there an expiry date? J Clin Pathol. 2017; 70(11): 988–993. doi: 10.1136/jclinpath-2017-204387</mixed-citation></citation-alternatives></ref></ref-list><fn-group><fn fn-type="conflict"><p>The authors declare that there are no conflicts of interest present.</p></fn></fn-group></back></article>
