Modes of embryo development in angiosperms

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Abstract

The general ideas that determine the structure of the embryo are discussed. A comparative analysis of the classifications of the endospermogenesis and embryogenesis types has shown that the principles of their creation are similar. However, during the endosperm development, the process of separation of the types (cellular and helobial, the latter includes the nuclear endosperm) begins as early as after the second division, and during the formation of the embryo, only after the third division during the formation of the proembryo, i. e. the ontogenetic time of typing differs in both systems. Using these principles, we proposed an original classification of the embryo formation modes. It uses the concept of “megatypes of embryogenesis” and analyzes existing modes and systems of types of embryo development. The data on the Piperad-type have been revised, and a new Orchidad-type has been proposed for orchids, including 3 variations.

After the division of the zygote, patterns of clusters of future types of embryogenesis are outlined: transverse, inclined, irregular (features of the formation of the first and subsequent partitions during the formation of the early embryo), coenocytic (nuclear stage in the development of the early embryo). The main cluster is transverse, which is inherent in most flowering plants. It is accompanied by the formation of apical and basal cells, further divisions of which lead to the formation of two ways of development based on the T-shaped or linear tetrad of cells. In each way, the participation of derivatives of apical and basal cells is different, which leads to the emergence of autonomous types of embryogenesis — Asterad-type, Caryophyllad-type, Chenopodiad-type, Onagrad-type, Solanad-type. The oblique cluster of megatypes is characterized by inclined partitions during the formation of proembryo (Poad-type of embryogenesis), while the irregular cluster is characterized by a combination of longitudinal, oblique, and transverse partitions (Piperad-type of embryogenesis) or a variety of tetrad forms of proembryo and the lack of regularity in the division and fate of basal cell derivatives (Orchidad-type of embryogenesis). The coenocyte cluster is represented by Paeonad-type only.

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About the authors

I. I. Shamrov

Herzen State Pedagogical University of Russia; Komarov Botanical Institute of Russian Academy of Sciences

Author for correspondence.
Email: shamrov52@mail.ru
Russian Federation, St. Petersburg; St. Petersburg

G. M. Anisimova

Komarov Botanical Institute of Russian Academy of Sciences

Email: galina0353@mail.ru
Russian Federation, St. Petersburg

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2. Fig. 1. Embryogenesis stages in Ceratophyllum demersum: 1–17 — early embryogenesis stages and embryo structure before organogenesis; 18–23 — organogenesis: cotyledon formation and epicotyle reorganization into embryo plumule. са — apical cell, сb — basal cell, сi — lower cell, icс — central cylinder initials, iec — root bark initials, ieр — epicotyle initials, l, l— upper and lower quadrants, m — middle cell, pсo — cotyledon, pl — embryo plumule. Scale bar, µm: 1–18 – 20; 19–23 – 100.

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3. Fig. 2. Some stages of embryogenesis in Ceratophyllum demersum: 1 — stage of octants; 2 — globular embryo; 3 — embryo before organogenesis; 4 — initiation of cotyledon and epicotyle primordia. ieр — epicotyle initials, pсo — cotyledon. Scale bar, µm: 30.

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4. Fig. 3. Embryogenesis stages in Luzula pedemontana: 1–13 — early embryogenesis stages and embryo structure before organogenesis, starch in cells is shown as granules, dextrines as dots; 14 — mature embryo. са — apical cell, сb — basal cell, сi — lower cell, co — root cap, h — hypophysial cell, icс — central cylinder initials, iec — root bark initials, l, l' — upper and lower quadrant-cells, m — middle cell, n, n'— derivates of middle cell, pсo — cotyledon, phy — hypocotyl, pvt — shoot apical meristem, q — quadrant-cells, s — suspensor. Scale bar, µm: 50.

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5. Fig. 4. Embryogenesis stages in Gymnadenia conopsea: 1–11 — successive stages of embryo development: 1 — zygote, 2 — two-celled proembryo, 3 — T-shaped cell tetrad, 4 — quadrant stage, 5 — octant stage, 6–11 — embryoderm differentiation and formation of a long single-row suspensor; starch in cells is shown as granules, dextrines as dots. са — apical cell, сb — basal cell, сi — lower cell, h — hypophysial cell, hp — hypostase, l, l — upper and lower quadrants, m — middle cell, n e — nucellus epiderm, pl — placenta, ps — postament, s — suspensor. Scale bar, µm: 30.

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6. Fig. 5. Embryogenesis stages in Listera ovata: 1–5 — development of the embryo surrounded by seed tissue: 1 — two-celled proembryo, 2 — T-shaped cell triad, 3 — quadrant stage, 4 — octant stage, 5 — mature seed, the basal cell of the embryo does not divide and forms a unicellular suspensor. cb, q — tiers of proembryo, em — embryo, en — endosperm, hp — hypostase. Scale bar, µm: 20.

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7. Fig. 6. Possible modes of embryo development in angiosperms: 1 — embryogenesis megatypes: 2 — transverse, 3 — oblique, 4 — irregular, 5 — coenocytic, 6 — embryogenesis types, 7 —T-shaped cell tetrad, 8 — linear cell tetrad, 9, 10 — contribution of ca and cb cells in the construction of the embryo, 11, 12 — predominantly ca cell — Onagrad-type (11), Solanad-type (12), 13, 14 — both cells — Asterad-type (13), Chenopodiadtype (14), 15 — cb cell does not divide, Caryophyllad-type, 16 — predominantly oblique cell divisions — Poad-type, 17, 18 —irregular type: combination of longitudinal, oblique and transverse divisions (Piperad-type — 17), variety of tetrad forms of proembryo cells and lack of regularity in the division and fate of basal cell derivatives (Orchidad-type — 18), 19 — nuclear divisions in proembryo — Paeoniad-type. са — apical cell, сb — basal cell.

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2. Категории обрабатываемых данных: файлы «cookies» (куки-файлы). Файлы «cookie» – это небольшой текстовый файл, который веб-сервер может хранить в браузере Пользователя. Данные файлы веб-сервер загружает на устройство Пользователя при посещении им Сайта. При каждом следующем посещении Пользователем Сайта «cookie» файлы отправляются на Сайт Оператора. Данные файлы позволяют Сайту распознавать устройство Пользователя. Содержимое такого файла может как относиться, так и не относиться к персональным данным, в зависимости от того, содержит ли такой файл персональные данные или содержит обезличенные технические данные.

3. Цель обработки персональных данных: анализ пользовательской активности с помощью сервиса «Яндекс.Метрика».

4. Категории субъектов персональных данных: все Пользователи Сайта, которые дали согласие на обработку файлов «cookie».

5. Способы обработки: сбор, запись, систематизация, накопление, хранение, уточнение (обновление, изменение), извлечение, использование, передача (доступ, предоставление), блокирование, удаление, уничтожение персональных данных.

6. Срок обработки и хранения: до получения от Субъекта персональных данных требования о прекращении обработки/отзыва согласия.

7. Способ отзыва: заявление об отзыве в письменном виде путём его направления на адрес электронной почты Оператора: info@rcsi.science или путем письменного обращения по юридическому адресу: 119991, г. Москва, Ленинский просп., д.32А

8. Субъект персональных данных вправе запретить своему оборудованию прием этих данных или ограничить прием этих данных. При отказе от получения таких данных или при ограничении приема данных некоторые функции Сайта могут работать некорректно. Субъект персональных данных обязуется сам настроить свое оборудование таким способом, чтобы оно обеспечивало адекватный его желаниям режим работы и уровень защиты данных файлов «cookie», Оператор не предоставляет технологических и правовых консультаций на темы подобного характера.

9. Порядок уничтожения персональных данных при достижении цели их обработки или при наступлении иных законных оснований определяется Оператором в соответствии с законодательством Российской Федерации.

10. Я согласен/согласна квалифицировать в качестве своей простой электронной подписи под настоящим Согласием и под Политикой обработки персональных данных выполнение мною следующего действия на сайте: https://journals.rcsi.science/ нажатие мною на интерфейсе с текстом: «Сайт использует сервис «Яндекс.Метрика» (который использует файлы «cookie») на элемент с текстом «Принять и продолжить».