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<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">anatomy</journal-id><journal-title-group><journal-title xml:lang="ru">Журнал анатомии и гистопатологии</journal-title><trans-title-group xml:lang="en"><trans-title>Journal of Anatomy and Histopathology</trans-title></trans-title-group></journal-title-group><issn pub-type="ppub">2225-7357</issn><publisher><publisher-name>N.N. Burdenko Voronezh State Medical University</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.18499/2225-7357-2021-10-2-73-79</article-id><article-id custom-type="elpub" pub-id-type="custom">anatomy-1315</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>ОБЗОРНЫЕ СТАТЬИ</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>REVIEW ARTICLES</subject></subj-group></article-categories><title-group><article-title>Функциональная морфология бокаловидных  клеток тонкой кишки при действии  различных факторов</article-title><trans-title-group xml:lang="en"><trans-title>Functional Morphology of Goblet Cells of the Small Intestine under the Influence of  Various Factors</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Шарапов</surname><given-names>И. Ю.</given-names></name><name name-style="western" xml:lang="en"><surname>Sharapov</surname><given-names>I. Yu.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Шарапов Илья Юрьевич</p><p>  ул.  Студенческая,  10,  Воронеж, 394036</p></bio><bio xml:lang="en"><p>Il'ya Sharapov</p><p> ul.  Studencheskaya, 10, Voronezh, 394036</p></bio><email xlink:type="simple">anat-vrn@yandex.ru</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Кварацхелиия</surname><given-names>А. Г.</given-names></name><name name-style="western" xml:lang="en"><surname>Kvaratskheliiya</surname><given-names>A. G.</given-names></name></name-alternatives><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Болгучева</surname><given-names>М. Б.</given-names></name><name name-style="western" xml:lang="en"><surname>Bolgucheva</surname><given-names>M. B.</given-names></name></name-alternatives><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Коротких</surname><given-names>К. Н.</given-names></name><name name-style="western" xml:lang="en"><surname>Korotkikh</surname><given-names>K. N.</given-names></name></name-alternatives><xref ref-type="aff" rid="aff-1"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru">ФГБОУ  ВО  «Воронежский  государственный  медицинский  университет  им.  Н.Н.  Бурденко» Минздрава России, Воронеж<country>Россия</country></aff><aff xml:lang="en">N.N. Burdenko Voronezh State Medical University, Voronezh<country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-2"><aff xml:lang="ru">ФГБОУ ВО «Ингушский государственный университет», Магас<country>Индия</country></aff><aff xml:lang="en">Ingush State University, Magas<country>India</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2021</year></pub-date><pub-date pub-type="epub"><day>15</day><month>07</month><year>2021</year></pub-date><volume>10</volume><issue>2</issue><fpage>73</fpage><lpage>79</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Шарапов И.Ю., Кварацхелиия А.Г., Болгучева М.Б., Коротких К.Н., 2021</copyright-statement><copyright-year>2021</copyright-year><copyright-holder xml:lang="ru">Шарапов И.Ю., Кварацхелиия А.Г., Болгучева М.Б., Коротких К.Н.</copyright-holder><copyright-holder xml:lang="en">Sharapov I.Y., Kvaratskheliiya A.G., Bolgucheva M.B., Korotkikh K.N.</copyright-holder><license 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://anatomy.elpub.ru/jour/article/view/1315">https://anatomy.elpub.ru/jour/article/view/1315</self-uri><abstract><p>Желудочно-кишечный тракт постоянно подвергается воздействию различных физических и химических  факторов.  В  кишечнике  контакт  бактерий  с  эпителиоцитами  в  значительной  степени  зависит  от слизи, которая в основном состоит из высоко  гликозилированного муцина-2, секретируемого бокаловидными клетками эпителия слизистой. Бокаловидные клетки располагаются по всей длине тонкой и толстой кишки и отвечают за выработку и поддержание защитного слоя слизи путем синтеза и секрециивысоко-молекулярных  гликопротеинов,  известных  как  муцины.  В  статье  представлены  данные  об  эмбриогенезе тонкой  кишки  в целом  и,  бокаловидных  клеток,  в частности;  представлен  литературный обзор, раскрывающий роль бокаловидных клеток в морфофункциональной организации кишечного тракта и функциональной  активности  их  секрета.  Ввиду  особенностей  этих  сильно  поляризованных  экзокринных  клеток обсуждаются клеточные механизмы, с помощью которых бокаловидные клетки секретируют свои продукты.</p></abstract><trans-abstract xml:lang="en"><p>The gastrointestinal tract is constantly exposed to various physical and chemical factors. In the intestine, the contact of bacteria and the epithelium largely depends on mucus, which mainly consists of highly glycosylated mucin-2 secreted by goblet cells in the epithelium. Goblet cells are located along the entire length of the small and large intestine and are responsible for the production and maintenance of a protective layer of mucus through the synthesis and secretion of high-molecular glycoproteins known as mucins. The article presents data on the embryogenesis of the small intestine in general and goblet cells, in particular, a literary review of the role of goblet cells in the morphology of the intestinal tract, the functional aсtivity of their secretion is carried out. Due to the unique nature of this highly polarized exocrine cell, the cellular mechanisms by which goblet cells  secrete their products are discussed.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>тонкая кишка</kwd><kwd>бокаловидные клетки</kwd><kwd>муцин</kwd><kwd>иммунная  система</kwd><kwd>факторы  среды</kwd></kwd-group><kwd-group xml:lang="en"><kwd>small intestine</kwd><kwd>goblet cells</kwd><kwd>mucin</kwd><kwd>immune system</kwd><kwd>environmental factors</kwd></kwd-group></article-meta></front><back><ref-list><title>References</title><ref id="cit1"><label>1</label><citation-alternatives><mixed-citation xml:lang="ru">Афанасьев Ю.И., Юрина Н.А. Гистология. М.: Медицина; 1983</mixed-citation><mixed-citation xml:lang="en">Afanas'ev  YuI,  Yurina  NA. Gistologiya. M.: Meditsina; 1983] (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Ачасова К.М., Литвинова Е.А., Кожевникова Е.Н. Роль компонентов муцина-2 в изменении состава нормальной микрофлоры кишечника и мукозального иммунитета мышей. Russian Scientist. 2017;1(2):8–9</mixed-citation><mixed-citation xml:lang="en">Litvinova  EA, Achasova  KM,  Kozhevnikova  EN.  The  role  of muсin-2  components  in  changes  in  the lora and mucosal immunity of mice. Russian Scientist. 2017;1(2):8–9] (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Бахтин А.А. Особености гистохимических показателей в тонкой кишке при воздействии неблагоприятных экологических факторов. Современные проблемы науки и образования. 2012;1:18</mixed-citation><mixed-citation xml:lang="en">Bakhtin  AA.  Peculiarity  of histochemicall  indicators  in  a  small  intestine under influence of bad ecological factors. Modern problems of science and education. 2012;1:18] (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Бахтин А.А., Наумова Л.И. Морфологическая характеристика стенки тонкой кишки при воздействии сероводородсодержащего газа. Астраханский медицинский журнал. 2012;7(4):37–40</mixed-citation><mixed-citation xml:lang="en">Bakhtin  AA,  Naumova  LI.  The  morfological characteristic  of  the  wall  of  small  intestine  after influence  of  hydrogen  sulphide  gas.  Astrakhan Medical Journal. 2012;7(4):37–40] (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Воробьев А.А., Несвижский Ю.В., Липницкий Е.М. и др. Микробное сообщество пристеночного муцина различных отделов желудочно-кишечного тракта человека. Вестник РАМН. 2004;(4):23–8</mixed-citation><mixed-citation xml:lang="en">Vorobyov  AA,  Nesvizhsky  YuV, Lipnitsky EM, et al. The parietal-mucin microbial population  in  different  sections  of  the  human gastrointestinal  tract.  Annals  Of  The  Russian Academy  Of Medical  Sciences.  2004;(4):23–8](in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Давлатова И.С., Наумова Л.И., Шишкина Т.А.Реактивность кишечного эпителия на фоне действия гипоксии разного генеза. Актуальные вопросы современной медицины: Материалы IV международной научно-практической конференции прикаспийских государств. Астра-хань. 2019; 274–6 [Davlatova IS, Naumova LI,</mixed-citation><mixed-citation xml:lang="en">Shishkina  TA.  The  reactivity  of  intestinal epithelium against the background of the action of hypoxia  of  different  genesis.  Aktual'nye  voprosy sovremennoi  meditsiny  :  Materialy  IV mezhdunarodnoi  nauchno-prakticheskoi konferentsii  prikaspiiskikh  gosudarstv. Astrakhan'. 2019; 274–6](in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Могильная Г.М., Пейливаньян Э.Г. Синтез муцинов бокаловидными гландулоцитами двенадцатиперстной кишки на различных этапах онтогенеза. Астраханский медицинский журнал. 2013;8(1):158–60</mixed-citation><mixed-citation xml:lang="en">Mogilnaya  GM, Peylivanyan EG. The synthesis of mucin by goblet glandulocytes  of  duodenum  at  different  stages  of ontogenesis.  Astrakhan  Medical  Journal. 2013;8(1):158–60] (in Russian).</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Симаненков В.И., Маев И.В., Ткачева О.Н., Алексеенко С.А., Андреев Д.Н., и др. Синдром повышенной эпителиальной проницаемости в клинической практике. Мультидисциплинарный национальный консенсус. Кардиоваскулярная терапия и профилактика. 2021;20(1):2758 [</mixed-citation><mixed-citation xml:lang="en">Simanenkov  VI,  Maev  IV, Tkacheva  ON,  Alekseenko  SA, Andreev  DN,  et  al. Syndrome  of  increased  epithelial  permeability  in clinical  practice.  Multidisciplinary  national Consensus.  Cardiovascular  Therapy  and Prevention. 2021 Feb 19;20(1):2758]. (in Russian) doi: 10.15829/1728-8800-2021-2758</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Al Alam D, Danopoulos S, Schall K, Sala FG, Almohazey D, Fernandez GE, et al. Fibroblast growth factor 10 alters the balance between goblet and Paneth cells in the adult mouse small intestine. American Journal of PhysiologyGastrointestinal and Liver Physiology. 2015 Apr 15;308(8):G678–90. doi: 10.1152/ajpgi.00158.2014</mixed-citation><mixed-citation xml:lang="en">Al  Alam D,  Danopoulos S,  Schall K,  Sala FG,  Almohazey D,  Fernandez  GE,  et  al. Fibroblast growth factor 10 alters the balance between goblet and  Paneth  cells  in  the  adult  mouse  small  intestine.  American  Journal  of  PhysiologyGastrointestinal  and  Liver  Physiology.  2015  Apr 15;308(8):G678–90.  doi: 10.1152/ajpgi.00158.2014</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Becciolini A, Fabbrica D, Cremonini D, Balzi M.Quantitative Changes in the Goblet Cells of the Rat Small Intestine after Irradiation. Acta Radiologica: Oncology. 1985 Jan;24(3):291–9. doi: 10.3109/02841868509134403</mixed-citation><mixed-citation xml:lang="en">Becciolini A,  Fabbrica D,  Cremonini D,  Balzi M.Quantitative Changes in the Goblet Cells of the Rat Small Intestine after Irradiation. Acta Radiologica: Oncology.  1985  Jan;24(3):291–9.  doi: 10.3109/02841868509134403</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Birchenough GMH, Johansson ME, Gustafsson JK, Bergström JH, Hansson GC. New developments in goblet cell mucus secretion and function. Mucosal Immunology. 2015 Apr 15;8(4):712–9. doi: 10.1038/mi.2015.32</mixed-citation><mixed-citation xml:lang="en">Birchenough GMH,  Johansson ME, Gustafsson JK,  Bergström JH,  Hansson GC. New developments  in  goblet  cell  mucus  secretion  and function.  Mucosal  Immunology.  2015  Apr 15;8(4):712–9. doi: 10.1038/mi.2015.32</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Boshuizen JA, Reimerink JHJ, Korteland-van Male AM, van Ham VJJ, Bouma J, Gerwig GJ, et al.Homeostasis and function of goblet cells during rotavirus infection in mice. Virology. 2005 Jul;337(2):210–21. doi: 10.1016/j.virol.2005.03.039</mixed-citation><mixed-citation xml:lang="en">Boshuizen JA,  Reimerink JHJ,  Korteland-van Male AM, van Ham VJJ, Bouma J, Gerwig  GJ, et al.Homeostasis and function of goblet cells during rotavirus  infection  in  mice.  Virology.  2005 Jul;337(2):210–21.  doi: 10.1016/j.virol.2005.03.039</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Calvert R, Bordeleau G, Grondin G, Vezina A, Ferrari J.On the presence of intermediate cells in the small intestine. The Anatomical Record. 1988 Mar;220(3):291–5. doi: 10.1002/ar.1092200310</mixed-citation><mixed-citation xml:lang="en">Calvert R,  Bordeleau G,  Grondin G,  Vezina  A, Ferrari  J.On the presence of intermediate cells in the small intestine. The Anatomical Record. 1988 Mar;220(3):291–5. doi: 10.1002/ar.1092200310</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Chairatana P, Nolan EM. Defensins, lectins, mucins, and secretory immunoglobulin A: microbe-binding biomolecules that contribute to mucosal immunity in the human gut. Crit Rev Biochem Mol Biol. 2017;52(1):45-56. doi:10.1080/10409238.2016. 1243654</mixed-citation><mixed-citation xml:lang="en">Chairatana P,  Nolan  EM. Defensins,  lectins, mucins,  and  secretory  immunoglobulin  A:  microbe-binding  biomolecules  that  contribute  to mucosal immunity in the human gut. Crit Rev Biochem  Mol  Biol.  2017;52(1):45-56. doi:10.1080/10409238.2016. 1243654</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Collins P. Development of midgut. 39th edn. Gray’s anatomy. Standring S, Ellis H, Healy JC, et al. London, Churchill Livingstone, 2005:1256-9.</mixed-citation><mixed-citation xml:lang="en">Collins P. Development  of  midgut.  39th  edn. Gray’s anatomy. Standring S, Ellis H, Healy JC, et al. London, Churchill Livingstone, 2005:1256-9.</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Dharmani P, Srivastava V, Kissoon-Singh V, et al.Role of intestinal mucins in innate host defense mechanisms against pathogens. J Innate Immun 2009;1(2):123–35.</mixed-citation><mixed-citation xml:lang="en">Dharmani P,  Srivastava V,  Kissoon-Singh V,  et al.Role of intestinal mucins in innate host defense mechanisms  against  pathogens.  J  Innate  Immun 2009;1(2):123–35.</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Eroschenko VP. diFiore’s atlas of histology with functional correlations. 11th edn. Lippincott Williams and Wilkins, 2005:341-2.</mixed-citation><mixed-citation xml:lang="en">Eroschenko VP. diFiore’s  atlas  of  histology  with functional  correlations.  11th  edn.  Lippincott  Williams and Wilkins, 2005:341-2.</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Forder REA, Howarth GS, Tivey DR, Hughes RJ.Bacterial Modulation of Small Intestinal Goblet Cells and Mucin Composition During Early Posthatch Development of Poultry. Poultry Science. 2007 Nov;86(11):2396–403. doi: 10.3382/ps.2007-00222</mixed-citation><mixed-citation xml:lang="en">Forder REA,  Howarth GS,  Tivey DR,  Hughes RJ.Bacterial  Modulation  of  Small  Intestinal  Goblet Cells  and  Mucin  Composition  During  Early  Posthatch  Development  of  Poultry.  Poultry  Science. 2007  Nov;86(11):2396–403.  doi: 10.3382/ps.2007-00222</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Gomes JR, Ayub LC, dos Reis CA, Machado MJ, da Silva J, Omar NF, et al.Goblet cells and intestinal Alkaline phosphatase expression (IAP) du ring the development of the rat small intestine.Acta Histochemica. 2017 Jan;119(1):71–7. doi: 10.1016/j.acthis.2016.11.010</mixed-citation><mixed-citation xml:lang="en">Gomes JR,  Ayub LC,  dos  Reis CA,  Machado MJ, da Silva J, Omar NF, et al.Goblet cells and intestinal  Alkaline  phosphatase  expression  (IAP)  du ring  the  development  of  the  rat  small  intestine.Acta  Histochemica.  2017  Jan;119(1):71–7.  doi: 10.1016/j.acthis.2016.11.010</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Johansson ME, Phillipson M, Petersson J, Velcich A, Holm L, Hansson GC.The inner of the two Muc2 mucin-dependent mucus layers in colon is devoid ofbacteria. Proc Natl Acad Sci USA. 2008;105:15064–15069.</mixed-citation><mixed-citation xml:lang="en">Johansson ME,  Phillipson M,  Petersson J,  Velcich A, Holm L, Hansson  GC.The inner of the two Muc2  mucin-dependent  mucus  layers  in  colon  is devoid ofbacteria. Proc Natl Acad Sci USA.  2008;105:15064–15069.</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Karam SM.Lineage commitment and maturation of epithelial cells in the gut. Front Biosci 1999;4:D286–98</mixed-citation><mixed-citation xml:lang="en">Karam SM.Lineage commitment and maturation of  epithelial  cells  in  the  gut.  Front  Biosci 1999;4:D286–98</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Kishida K, Pearce SC, Yu S, Gao N, Ferraris RP.Nutrient sensing by absorptive and secretory progenies of small intestinal stem cells. American Journal of Physiology-Gastrointestinal and Liver Physiology. 2017 Jun 1;312(6):G592–605. doi: 10.1152/ajpgi.00416.2016</mixed-citation><mixed-citation xml:lang="en">Kishida K,  Pearce SC,  Yu S,  Gao  N,  Ferraris RP.Nutrient  sensing  by  absorptive  and  secretory progenies of small intestinal stem cells. American Journal  of  Physiology-Gastrointestinal  and  Liver Physiology.  2017  Jun  1;312(6):G592–605.  doi: 10.1152/ajpgi.00416.2016</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Lang T, Klasson S, Larsson E, Johansson MEV, Hansson GC, Samuelsson T.Searching the Evolutionary Origin of Epithelial Mucus Protein Components—Mucins and FCGBP. Molecular Biology and Evolution. 2016 Apr 4;33(8):1921–36</mixed-citation><mixed-citation xml:lang="en">Lang T,  Klasson S,  Larsson E,  Johansson MEV, Hansson GC, Samuelsson T.Searching the Evolutionary  Origin  of  Epithelial  Mucus  Protein  Components—Mucins  and  FCGBP.  Molecular  Biology and Evolution. 2016 Apr 4;33(8):1921–36</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Larsen WJ. Development of the gastrointestinal tract. Human embryology. 2nd edn. New York. Churchill Livingstone; 1997:245–8.</mixed-citation><mixed-citation xml:lang="en">Larsen WJ. Development  of  the  gastrointestinal tract.  Human  embryology.  2nd  edn.  New  York. Churchill Livingstone; 1997:245–8.</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Lei W, Ren W, Ohmoto M, Urban JF, Matsumoto I, Margolskee RF, et al.Activation of intestinal tuft cell-expressed Sucnr1 triggers type 2 immunity in the mouse small intestine. Proceedings of the National Academy of Sciences. 2018 May 7;115(21):5552–7. doi: 10.1073/pnas.1720758115</mixed-citation><mixed-citation xml:lang="en">Lei W,  Ren W,  Ohmoto M,  Urban JF,  Matsumoto I,  Margolskee RF,  et  al.Activation  of  intestinal  tuft  cell-expressed  Sucnr1  triggers  type  2 immunity  in  the  mouse  small  intestine.  Proceedings  of  the National  Academy  of  Sciences.  2018 May  7;115(21):5552–7.  doi: 10.1073/pnas.1720758115</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Levin DE, Barthel ER, Speer AL, Sala FG, Hou X, Torashima Y, et al. Human tissue-engineered small intestine forms from postnatal progenitor cells. Journal of Pediatric Surgery. 2013 Jan;48(1):129–37. doi: 10.1016/j.jpedsurg.2012.10.029</mixed-citation><mixed-citation xml:lang="en">Levin DE,  Barthel ER,  Speer AL,  Sala FG,  Hou X, Torashima Y,  et  al. Human  tissue-engineered small  intestine  forms  from  postnatal  progenitor cells.  Journal  of  Pediatric  Surgery.  2013 Jan;48(1):129–37.  doi: 10.1016/j.jpedsurg.2012.10.029</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Lievin-Le Moal V, Servin AL. The front line of enteric host defense against unwelcome intrusion of harmful microorganisms: mucins, antimicrobial peptides, and microbiota. Clin Microbiol Rev2006;19(2):315–37.</mixed-citation><mixed-citation xml:lang="en">Lievin-Le  Moal V,  Servin  AL. The  front  line  of enteric host defense against unwelcome intrusion of harmful microorganisms: mucins, antimicrobial peptides,  and  microbiota.  Clin  Microbiol  Rev2006;19(2):315–37.</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Liu SP, Chang CY, Huang WH, Fu YS, Chao D, Huang HT.Dimethylthiourea pretreatment inhibits endotoxin-induced compound exocytosis in goblet cells and plasma leakage of rat small intestine. Journal of Electron Microscopy. 2009 Oct 12;59(2):127–39. doi: 10.1093/jmicro/dfp049</mixed-citation><mixed-citation xml:lang="en">Liu SP,  Chang CY,  Huang WH,  Fu YS,  Chao D, Huang HT.Dimethylthiourea pretreatment inhibits  endotoxin-induced  compound  exocytosis  in goblet cells and plasma leakage of rat small intestine.  Journal  of  Electron  Microscopy.  2009  Oct 12;59(2):127–39. doi: 10.1093/jmicro/dfp049</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Merzel J, Leblond CP.Origin and renewal of goblet cells in the epithelium of the mouse small intestine. American Journal of Anatomy. 1969 Mar;124(3):281–305. doi: 10.1002/aja.1001240303</mixed-citation><mixed-citation xml:lang="en">Merzel J, Leblond CP.Origin and renewal of goblet  cells  in  the  epithelium  of  the  mouse  small  intestine.  American  Journal  of  Anatomy.  1969 Mar;124(3):281–305.  doi: 10.1002/aja.1001240303</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Middendorp S, Schneeberger K, Wiegerinck CL, Mokry M, Akkerman RDL, van Wijngaarden S, et al.Adult Stem Cells in the Small Intestine Are Intrinsically Programmed with Their LocationSpecific Function. STEM CELLS. 2014 Apr 17;32(5):1083–91. doi: 10.1002/stem.1655</mixed-citation><mixed-citation xml:lang="en">Middendorp S,  Schneeberger K,  Wiegerinck CL, Mokry M, Akkerman RDL, van Wijngaarden S, et al.Adult Stem Cells in the Small Intestine Are Intrinsically  Programmed  with  Their  LocationSpecific  Function.  STEM  CELLS.  2014  Apr 17;32(5):1083–91. doi: 10.1002/stem.1655</mixed-citation></citation-alternatives></ref><ref id="cit31"><label>31</label><citation-alternatives><mixed-citation xml:lang="ru">Pelaseyed T, Bergström JH, Gustafsson JK, Ermund A, Birchenough GMH, Schütte A, et al.The mucus and mucins of the goblet cells and enterocytes provide the first defense line of the gastrointestinal tract and interact with the immune system. Immunological reviews. 2014;260(1):8–20. doi: 10.1111/imr.12182</mixed-citation><mixed-citation xml:lang="en">Pelaseyed T,  Bergström JH,  Gustafsson JK,  Ermund A, Birchenough GMH, Schütte  A, et al.The mucus and mucins of the goblet cells and enterocytes provide the first defense line of the gastrointestinal  tract  and  interact  with  the  immune  system.  Immunological  reviews.  2014;260(1):8–20. doi: 10.1111/imr.12182</mixed-citation></citation-alternatives></ref><ref id="cit32"><label>32</label><citation-alternatives><mixed-citation xml:lang="ru">Pfoze K, Rajshree H.Time of appearance of goblet cells in human small intestine. Journal of Evolution of Medical and Dental Sciences. 2018 Apr 23;7(17):2099–103. doi: 10.14260/jemds/2018/470</mixed-citation><mixed-citation xml:lang="en">Pfoze K, Rajshree H.Time of appearance of goblet cells  in  human  small  intestine.  Journal  of  Evolution  of  Medical  and  Dental  Sciences.  2018  Apr 23;7(17):2099–103.  doi: 10.14260/jemds/2018/470</mixed-citation></citation-alternatives></ref><ref id="cit33"><label>33</label><citation-alternatives><mixed-citation xml:lang="ru">Rojanapo W, Lamb AJ, Olson JA.The Prevalence, Metabolism and Migration of Goblet Cells in Rat Intestine following the Induction of Rapid, Synchronous Vitamin A Deficiency. The Journal of Nutrition. 1980 Jan 1;110(1):178–88. doi: 10.1093/jn/110.1.178</mixed-citation><mixed-citation xml:lang="en">Rojanapo W, Lamb AJ, Olson JA.The Prevalence, Metabolism  and  Migration  of  Goblet  Cells  in  Rat Intestine  following  the  Induction  of  Rapid,  Synchronous  Vitamin  A  Deficiency.  The  Journal  of Nutrition.  1980  Jan  1;110(1):178–88.  doi: 10.1093/jn/110.1.178</mixed-citation></citation-alternatives></ref><ref id="cit34"><label>34</label><citation-alternatives><mixed-citation xml:lang="ru">Rubio CA.Paneth cells and goblet cells express the neuroendocrine peptide synaptophysin. I. Normal duodenal mucosa. In Vivo. 2012 JanFeb;26(1):135-8.</mixed-citation><mixed-citation xml:lang="en">Rubio CA.Paneth cells and goblet cells express the neuroendocrine peptide synaptophysin. I. Normal duodenal  mucosa.  In  Vivo.  2012  JanFeb;26(1):135-8.</mixed-citation></citation-alternatives></ref><ref id="cit35"><label>35</label><citation-alternatives><mixed-citation xml:lang="ru">Salim SY, Söderholm JD.Importance of disrupted intestinal barrier in inflammatory bowel diseases. Inflamm Bowel Dis. 2011;17(1):362-81.</mixed-citation><mixed-citation xml:lang="en">Salim SY, Söderholm JD.Importance of disrupted intestinal barrier in inflammatory bowel diseases. Inflamm Bowel Dis. 2011;17(1):362-81.</mixed-citation></citation-alternatives></ref><ref id="cit36"><label>36</label><citation-alternatives><mixed-citation xml:lang="ru">Specian RD, Oliver MG. Functional biology of intestinal goblet cells. The American journal of physiology. 1991;260(2 Pt 1):C183-93. doi: 10.1152/ajpcell.1991.260.2.C183</mixed-citation><mixed-citation xml:lang="en">Specian RD,  Oliver MG. Functional  biology  of intestinal  goblet  cells.  The  American  journal  of physiology.  1991;260(2  Pt  1):C183-93.  doi: 10.1152/ajpcell.1991.260.2.C183</mixed-citation></citation-alternatives></ref><ref id="cit37"><label>37</label><citation-alternatives><mixed-citation xml:lang="ru">van Es JH, van Gijn ME, Riccio O, van den Born M, Vooijs M, Begthel H, et al. Notch/γ-secretase inhibition turns proliferative cells in intestinal crypts and adenomas into goblet cells. Nature. 2005 Jun;435(7044):959–63. doi: 10.1038/nature03659</mixed-citation><mixed-citation xml:lang="en">van  Es JH,  van  Gijn ME,  Riccio O,  van  den Born M,  Vooijs M,  Begthel H,  et  al. Notch/γ-secretase  inhibition turns proliferative cells in intestinal crypts and adenomas into goblet cells. Nature.  2005  Jun;435(7044):959–63.  doi: 10.1038/nature03659</mixed-citation></citation-alternatives></ref><ref id="cit38"><label>38</label><citation-alternatives><mixed-citation xml:lang="ru">Verburg M, Renes IB, Meijer HP, Taminiau JAJM, Büller HA, Einerhand AWC, et al. Selective sparing of goblet cells and Paneth cells in the intestine of methotrexate-treated rats. American Journal of Physiology-Gastrointestinal and Liver Physiology. 2000 Nov 1;279(5):G1037–47. doi: 10.1152/ajpgi.2000.279.5.G1037</mixed-citation><mixed-citation xml:lang="en">Verburg M,  Renes IB,  Meijer  HP, Taminiau JAJM,  Büller HA,  Einerhand  AWC,  et al. Selective  sparing  of  goblet  cells  and  Paneth cells in the intestine of methotrexate-treated rats. American  Journal  of  Physiology-Gastrointestinal and Liver Physiology. 2000 Nov 1;279(5):G1037–47. doi: 10.1152/ajpgi.2000.279.5.G1037</mixed-citation></citation-alternatives></ref><ref id="cit39"><label>39</label><citation-alternatives><mixed-citation xml:lang="ru">von Moltke J, Ji M, Liang H-E, Locksley RM.Tuft-cell-derived IL-25 regulates an intestinal ILC2–epithelial response circuit. Nature. 2015 Dec 14;529(7585):221–5. doi: 10.1038/nature16161</mixed-citation><mixed-citation xml:lang="en">von  Moltke J,  Ji M,  Liang H-E,  Locksley  RM.Tuft-cell-derived  IL-25  regulates  an  intestinal ILC2–epithelial response circuit. Nature. 2015 Dec 14;529(7585):221–5. doi: 10.1038/nature16161</mixed-citation></citation-alternatives></ref><ref id="cit40"><label>40</label><citation-alternatives><mixed-citation xml:lang="ru">Wang H, Liu J, Zhao W, Zhang Z, Li S, Li S, et al.Effect of Fluoride on Small Intestine Morphology and Serum Cytokine Contents in Rats. Biological Trace Element Research. 2018 Sep 13;189(2):511–8. doi: 10.1007/s12011-018-1503-y</mixed-citation><mixed-citation xml:lang="en">Wang H, Liu J, Zhao W, Zhang Z, Li S, Li S, et al.Effect  of  Fluoride  on  Small Intestine  Morphology and  Serum  Cytokine Contents  in  Rats.  Biological Trace Element Research. 2018 Sep 13;189(2):511–8. doi: 10.1007/s12011-018-1503-y</mixed-citation></citation-alternatives></ref><ref id="cit41"><label>41</label><citation-alternatives><mixed-citation xml:lang="ru">Wang Y, Song W, Wang J, Wang T, Xiong X, Qi Z, et al. Single-cell transcriptome analysis reveals differential nutrient absorption functions in human intestine. Journal of Experimental Medicine. 2019 Nov 21;217(2) doi: 10.1084/jem.20191130</mixed-citation><mixed-citation xml:lang="en">Wang Y,  Song W,  Wang J,  Wang T,  Xiong X, Qi Z,  et  al. Single-cell  transcriptome  analysis  reveals  differential  nutrient  absorption  functions  in human  intestine.  Journal  of  Experimental  Medicine.  2019  Nov  21;217(2)  doi: 10.1084/jem.20191130</mixed-citation></citation-alternatives></ref><ref id="cit42"><label>42</label><citation-alternatives><mixed-citation xml:lang="ru">Wigley C. Microstructure of the small intestine. Gray’s anatomy. 39th edn. Standring S, Ellis H, Healy JC, et al. London, Churchill Livingstone, 2005:1157-69.</mixed-citation><mixed-citation xml:lang="en">Wigley C. Microstructure  of  the  small  intestine. Gray’s  anatomy.  39th  edn.  Standring  S,  Ellis  H, Healy  JC,  et  al.  London,  Churchill  Livingstone, 2005:1157-69.</mixed-citation></citation-alternatives></ref><ref id="cit43"><label>43</label><citation-alternatives><mixed-citation xml:lang="ru">Ye DZ, Kaestner KH.Foxa1 and Foxa2 control the differentiation of goblet and enteroendocrine Land D-cells in mice. Gastroenterology. 2009 Dec 1;137(6):2052–62. doi: 10.1053/j.gastro.2009.08.059</mixed-citation><mixed-citation xml:lang="en">Ye DZ, Kaestner KH.Foxa1 and Foxa2 control the differentiation  of  goblet  and  enteroendocrine  Land  D-cells  in  mice.  Gastroenterology.  2009  Dec 1;137(6):2052–62.  doi: 10.1053/j.gastro.2009.08.059</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>
