Становление щетиночного узора у Drosophila melanogaster: предструктура и комплекс генов achaete-scute

Внешние механорецепторы дрозофилы, локализованные на голове и теле имаго, представлены щетинками разного размера – макро- и микрохетами. Макрохеты образуют устойчивую структурную композицию, так называемый щетиночный узор, специфичный для каждого вида дрозофилы, в котором каждая из макрохет занимает строго определенное положение. Формирование щетиночного узора начинается с формирования его прообраза в имагинальном диске. Специфичность позиций будущих механорецепторов определяется локальной экспрессией двух пронейральных генов – achaete (ac) и scute (sc), входящих в комплекс AS-C, в ответ на действие неких факторов, за которыми закрепилось название «факторы предструктуры», гетерогенно распределенных в эктодерме имагинальных дисков. Топография их совокупного распределения и создает прообраз (предструктуру) щетиночного узора. Таким образом, полноценный щетиночный узор является результатом взаимодействия двух систем: предструктуры и системы ответа на предструктуру – генов achaete и scute. К настоящему времени накоплено значительное число разрозненных экспериментальных данных, касающихся различных аспектов формирования щетиночного узора, однако формализованное представление полного спектра молекулярно-генетических взаимодействий факторов предструктуры как между собой, так и с генами комплекса AS-C, в литературе отсутствует. В обзоре систематизированы данные о закономерностях этих взаимодействий. Показано, что экспрессия пронейральных генов achaete-scute детерминируется иерархически организованной двухуровневой системой управления, содержащей как прямые, так и непрямые регуляторы их активности. Предложена обобщенная схема системы, включающая функциональные взаимодействия ее компонентов.

Drosophila melanogaster, макрохеты, щетиночный узор, пронейральный кластер, предструктура, генный комплекс achaete-scute

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