vavilovВавиловский журнал генетики и селекцииVavilov Journal of Genetics and Breeding2500-3259Institute of Cytology and Genetics of Siberian Branch of the RAS10.18699/VJ15.109vavilov-489Research ArticleБиоинформатика растенийPlant BioinformaticsКлеточная стенка растений и механизмы устойчивости к патогенамPlant cell wall and the mechanisms of resistance to pathogensСмирноваО. Г.SmirnovaO. G.planta@bionet.nsc.ruКочетовА. В.KochetovA. V.Федеральное государственное бюджетное научное учреждение «Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук», Новосибирск, Россия Федеральное государственное автономное образовательное учреждение высшего образования «Новосибирский национальный исследовательский государственный университет», Новосибирск, РоссияРоссияInstitute of Cytology and Genetics SB RA S, Novosibirsk, Russia Novosibirsk State University, Novosibirsk, RussiaRussian Federation201504012016196715723Copyright © Смирнова О.Г., Кочетов А.В., 20162016Смирнова О.Г., Кочетов А.В.Smirnova O.G., Kochetov A.V.This work is licensed under a Creative Commons Attribution 4.0 License.https://vavilov.elpub.ru/jour/article/view/489

Огромное число грибов, бактерий и вирусов потенциально способны инфицировать ткани и вызывать заболевания растений. Устойчивость растений к патогенам основывается на сложной сети конститутивных и индуцированных защитных реакций, в контроле которых задействовано большое число генов. Клеточная стенка является первым препятствием, которое должны преодолеть патогенные микроорганизмы. Успешная защита на уровне клеточной стенки может остановить вторжение подавляющего большинства потенциальных фитопатогенов. Разные виды растений различаются по структуре клеточной стенки. Основу клеточной стенки составляет сеть из микрофибрилл целлюлозы, пересекаемых молекулами гемицеллюлозы. В растущих частях растения эта сеть встроена в матрикс из пектиновых полисахаридов. В уже сформировавшихся тканях клеточные стенки усилены лигнином. Кроме полисахаридов, клеточная стенка содержит значительное количество белков, выполняющих структурную и ферментативную функции. Информация о многочисленных белках клеточных стенок разных видов растений представлена в базе данных WallProtDB. Каждый из компонентов клеточной стенки вносит вклад в формирование устойчивости к патогенам. В местах контакта с потенциальными патогенами происходит дополнительное укрепление клеточной стенки и накопление антимикробных вторичных метаболитов. Патогены секретируют ферменты, способные расщеплять компоненты клеточной стенки. В ответ на атаку микробов растение продуцирует ингибиторы микробных гидролитических ферментов. Растение также способно оценивать количество компонентов клеточной стенки. Так, мутанты с дефицитом целлюлозы обычно имеют повышенный уровень лигнификации и усиление защитного ответа. Возникающие после действия микробных ферментов низкомолекулярные фрагменты клеточной стенки выполняют сигнальную функцию, усиливая защитную реакцию растения. Таким образом, клеточная стенка является динамической структурой, способной предотвращать проникновение большинства потенциальных патогенов и запускать разные варианты иммунного ответа. Реконструкция генных сетей, контролирующих структурно-функциональную организацию клеточной стенки в процессе роста и в условиях биотических и абиотических стрессов, необходима для понимания молекулярных механизмов развития и стрессоустойчивости. В обзоре рассматриваются механизмы специфической и неспецифической устойчивости растений к патогенам различной природы, связанные с клеточной стенкой. Обсуждаются структура клеточной стенки и роль различных компонентов в детекции инвазии фитопатогенов и индукции защитных механизмов.

A huge variety of phytopathogens (viruses, bacteria, fungi) are potentially able to infect plant tissues and cause diseases. Numerous plant genes control a complex network of defense mechanisms based on both constitutive and inducible processes. The cell wall is a primary barrier the pathogens have to penetrate to start the infection process. However,it is able to block invasion by most non-specific potential pathogens. The cell wall structure may differ in various plant species. It is based on the net of cellulose microfibrils linked by hemicellulose molecules. Pectin and lignin are the other important cell wall constituents. Dozens of proteins inside the cell wall are involved in structural and metabolic processes as well as in signal transduction and regulatory circuits (more information is available in W allProtDB database). Each of these components contributes to resistance to pathogens. At the points of contact with potential pathogens cell wall structural changes and accumulation of metabolites with antimicrobial, antifungal or antiviral activities occur. Some pathogens could produce hydrolytic enzymes able to degrade cellulose and pectin to counteract these non-specific plant resistance mechanisms. In turn, plants developed the inhibitors of pathogen-related enzymes and this “arms race” is an important part of plant evolution and host-pathogen interaction mechanisms. Plants also can evaluate the cell wall state to compensate for imbalances and deficiencies. For instance, mutants with cellulose deficiency may have a higher lignification rate and a stronger stress response. The cell wall is also a source of signal molecules triggering the initiation of response mechanisms. In total, the plan cell wall is a complex dynamic structure able to prevent infection by most potential (non-specific) pathogens and switch on the mechanisms of plant immune response. The reconstruction of gene networks controlling the cell wall structural and functional organization during the growth, and under normal and stressful conditions is vitally important for understanding the basic molecular mechanisms of development and stress resistance. The mechanisms of specific and non- specific plant resistance to various phytopathogens connected to the cell wall structure are reviewed. The roles of the cell wall constituents in pathogen detection and the induction of defense mechanism are discussed

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The authors declare that there are no conflicts of interest present.