Epidermal patterning and stomatal development in Gnetales

Background and Aims The gymnosperm order Gnetales, which has contentious phylogenetic affinities, includes three extant genera (Ephedra, Gnetum, Welwitschia) that are morphologically highly divergent and have contrasting ecological preferences: Gnetum occupies mesic tropical habitats, whereas Ephedra and Welwitschia occur in arid environments. Leaves are highly reduced in Ephedra, petiolate with a broad lamina in Gnetum and persistent and strap-like in Welwitschia. We investigate stomatal development and prepatterning stages in Gnetales, to evaluate the substantial differences among the three genera and compare them with other seed plants. Methods Photosynthetic organs of representative species were examined using light microscopy, scanning electron microscopy and transmission electron microscopy. Key Results Stomata of all three genera possess lateral subsidiary cells (LSCs). LSCs of Ephedra are perigene cells derived from cell files adjacent to the stomatal meristemoids. In contrast, LSCs of Gnetum and Welwitschia are mesogene cells derived from the stomatal meristemoids; each meristemoid undergoes two mitoses to form a ‘developmental triad’, of which the central cell is the guard mother cell and the lateral pair are LSCs. Epidermal prepatterning in Gnetum undergoes a ‘quartet’ phase, in contrast with the linear development of Welwitschia. Quartet prepatterning in Gnetum resembles that of some angiosperms but they differ in later development. Conclusions Several factors underpin the profound and heritable differences observed among the three genera of Gnetales. Stomatal development in Ephedra differs significantly from that of Gnetum and Welwitschia, more closely resembling that of other extant gymnosperms. Differences in epidermal prepatterning broadly reflect differences in growth habit between the three genera.

SOL1 and SOL2 Regulate Fate Transition and Cell Divisions in the Arabidopsis Stomatal Lineage

In the stomatal lineage, cells make fate transitions from asymmetrically dividing and self-renewing meristemoids, to commitment to the guard mother cell identity, and finally though a single division to create mature, post-mitotic stomatal guard cells. Flexibility in the stomatal lineage allows plants to alter leaf size and stomatal density in response to environmental conditions; however, transitions must be clean and unidirectional in order to produce functional and correctly patterned stomata. Among direct transcriptional targets of the stomatal initiating factor, SPEECHLESS, we found a pair of genes, SOL1 and SOL2, required for effective transitions in the lineage. Here we show that these two genes, which are homologues of the LIN54 DNA-binding components of the mammalian DREAM complex, are expressed in a cell cycle dependent manner and regulate cell fate and division properties in the self-renewing early lineage. In the terminal division of the stomatal lineage, however, these two proteins appear to act in opposition to their closest paralogue, TSO1, revealing complexity in the gene family may enable customization of cell divisions in coordination with development.

Stimulation of the final cell cycle in the stomatal lineage by the cyclin CYCD7;1 under regulation of the MYB transcription factor FOUR-LIPS

(180 words)Stomatal guard cells are formed through a sequence of asymmetric and symmetric divisions in the epidermis of the sporophyte of most land plants. We show that several D-type cyclins are consecutively activated in the stomatal linage in the epidermis of Arabidopsis thaliana. Whereas CYCD2;1 and CYCD3;2 are activated in the meristemoids early in the lineage, CYCD7;1 is activated before the final division. CYCD7;1 expression peaks in the guard mother cell, where its transcription is modulated by the FOUR-LIPS/MYB88 transcription factor. FOUR-LIPS/MYB88 interacts with the CYCD7;1 promoter and represses CYCD7;1 transcription. CYCD7;1 stimulates the final symmetric division in the stomatal lineage, since guard cell formation is delayed in the cycd7;1 mutant epidermis and guard mother cell (GMC) divisions in four-lips mutant guard mother cells are limited by loss of function of CYCD7;1. Hence, the precise activation of a specific D-type cyclin, CYCD7;1, is required for correct timing of the last symmetric division that creates the stomatal guards cells, and CYCD7;1 expression is regulated by the FLP/MYB pathway that ensures cell cycle arrest in the stomatal guard cells.Summary StatementThe formation of paired guard cells in the epidermis of the Arabidopsis thaliana shoot, requires the activity of the D-type cyclin CYCD7;1 for the normal timing of the final division.

The guard mother cell dormancy of soybean cotyledon

The cotyledons of soybean seeds serve both as a food source and a photosynthetic organs for the early development of soybean seedlings. The development of the stomata of cotyledons should be important and related to the food source utilization and the mechanism of photosynthesis. During the study of the development of the cotyledon stomata of soybean seeds, a phenomenon was observed on the upper epidermal layer of the cotyledon, we call it. “the dormancy of the guard mother cell”, which is not reported in the literature available to us.Soybean seeds were soaked, planted and sampled at 24, 48, and 72 hours after planting. The upper epidermal layers were stripped with a sharp razor blade, and processed for observation with Hitachi S-2300 scanning electron microscope.

Development of the Stomatal Complexes During Ontogeny in Eucalyptus and Angophora (Myrtaceae)

The mode of stomatal development is studied in cotyledons, seedling and adult leaves of species of eucalypts and three species of Angophora. In the cotyledons of all species examined the early stomatal initials are unilabrate or dolabrate. The stomatal initials in seedling leaves of species of the Corymbosae and Clavigerae are anisocytic. In the 4th seedling leaf in species of a group we have previously called Monocalyptus the stomatal initials are also anisocytic. All other eucalypts retain the early cotyledonary mode of origin of stomata throughout life. These two modes of origin, whether anisocytic or by unilabrate and dolabrate initials, are set in all eucalypts from the 4th seedling leaf onward. Secondary characteristics of the adult stomata, e.g. number of subsidiary cells, are more complex than those of the seedling leaves; rarely, the relatively simple pattern of the seedling leaves may persist in the adult leaves of a given species. In species in which the initials in adult leaves are unilabrate or dolabrate, groups of stomata may share one or more subsidiary cells or be juxtaposed without an intervening subsidiary cell. The sister cell(s) of the guard mother cell may precociously develop a thicker cuticle than ordinary epidermal cells, and this may be apparent at maturity. The abaxial stomata of the cotyledons (but not of seedling or adult leaves) are regularly aligned parallel to the main venation. The existence of three main types of origin of stomata characteristic of three large non-interbreeding groups of eucalypts is of interest in the taxonomy of the genus.

The subcellular organization of stomatal initials in sugarcane leaves: the guard and subsidiary mother cells

The subcellular organization of guard and subsidiary mother cells in sugarcane leaves was examined by electron microscopy. Guard and subsidiary mother cells assume a characteristic shape before mitosis and contain variable numbers of mitochondria, proplastids, dictyosomes, and cisternae of rough endoplasmic reticulum. In guard mother cells, the nucleus occupies a central position, whereas in subsidiary mother cells, the nucleus is located toward one end of the cell, near the guard mother cell. Microtubules are found in both guard and subsidiary mother cells and are either closely grouped to form defined preprophase bands or randomly dispersed between the nucleus and the preprophase bands. Many of the dispersed microtubules occur in close association with the nucleus in both guard and subsidiary mother cells. Possible functions for these preprophase microtubules are discussed in relation to their organization in the preprophase band, their orientation, and their distribution within guard and subsidiary mother cells.

Pre Prophase Microtubules and Stomatal Differentiation in Commelina Cy Anea

The relationship between preprophase microtubules and asymmetrical cell division in the formation of the stomatal complex of C. cyanea was investigated. Polarization of nuclei and other cell organelles adjacent to the guard mother cell occurred in most cases without a preprophase band of microtubules being present; the grouping of preprophase microtubules appeared immediately prior to cell division, and its situation, even during abnormal stomatal development, predicted the plane of future division. The results show that preprophase microtubules cannot be the cytoplasmic agents involved in orienting and positioning the nucleus prior to division. Clear evidence was obtained indicating that preprophase microtubules move intact into the spindle. Some aspects of abnormal stomatal development are discussed, and the results are related to some other work on stomatal differentiation.