4. Produce and Disperse Pollen
Androecium Stamens
Stamens Represent Modified Leaves
All Angiospermous Anthers are Bilobed and Quadrilocular (i.e.,
formed of four micro- sporangia) at an Early Stage of Development
and This Condition is Seen in Most Mature Stamens.
The final anther anatomy and the progression of developmental
stagesaregenerallyconservedinangiosperms,althoughtherearemanyv
ariationsinanthergrowth parameters , number of days to anthesis,
and mechanisms of pollen shed (the termination ofanther
development)
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differentiation of organelles in pre-meiotic rice anthers. Cell Biology International, 29(9), 792–802.
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Scott, R. J. (2004). Stamen Structure and Function. The Plant Cell Online, 16(suppl_1), S46–S60.
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(2012). Maize multiple archesporial cells 1 (mac1), an ortholog of rice TDL1A, modulates cell
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Fig. 2. (A) Typical epidermal cells with thick external walls covered in cuticle, most cells also having a large central vacuole with the tonoplast
membrane visible (arrowheads). Plasmodesmata connect the adjacent cells (arrows). Nucleoli were observed within the nuclei. (B) At high
magnification the bi-lamellate nature of the evenly distributed cuticle on the epidermal cells (arrows) is apparent. (C,D) Portions of endothecium cells
exhibiting numerous starch containing chloroplasts and clusters of small mitochondria respectively. In (D), a region of cortical cytoplasm, lipid
bodies and microtubules are also visible (arrows). en, endothecium; chl, chloroplast; m, mitochondrion; p, plastid; v, vacuole; s, starch granule; lb,
lipid body; mt, microtubules; neo, nucleolus.
Fig. 4. (A) The chloroplasts in the endothecium contain starch granules surrounded by immature but well defined thylakoids and grana. Microbodies
were often associated with the chloroplasts. (B) Amyloplasts are also present in the endothecium. (C) A section of cell wall between neighbouring
endothecium cells showing a plasmodesma (arrows) linking the cells. (D) Middle layer cell during pre-meiotic development exhibiting a central
nucleus with mitochondria, plastids and vacuoles. (E) Middle layer cells are connected with one another by plasmodesmata. en, endothecium; ml,
middle layer; n, nucleus; v, vacuole; lb, lipid body; m, mitochondrion; am, amyloplast; p, plastid; th, thylakoid; gr, grana; mb, microbody.
Fig. 5. (A) Tapetal cells exhibiting large central nuclei, numerous mitochondria, small vacuoles, vesicles, lipid bodies and plastids. The nuclear
envelope displays dilatation with large spaces present between the double membranes (arrowheads). In the neighbouring sporogenous cell, electron
opaque bodies are visible near the cells wall (arrows). (B) Like the surrounding tapetal cells, the sporogenous cells are densely cytoplasmic with
numerous organelles. (C) High magnification image of sporogenous cells containing numerous organelles, many of which are similar in size and
shape and in a relatively undifferentiated state. Lipid bodies and numerous vesicles of different sizes including electron opaque bodies are also seen in
the cytoplasm close to the cell wall (arrows). (D,E) Plasmodesmata (arrows) exist between tapetal cells, and tapetal and middle layer cells (D) and
neighbouring sporogenous cells. (F) Detail of cell wall between tapetum and sporogenous cells indicating wall thickenings (arrows). t, tapetum; ml,
middle layer; m, mitochondrion; p, plastid; lb, lipid body; r, ribosomes; v, vacuole; n, nucleus; neo, nucleolus; ch, chromatin; sp, sporogenous cell; vs,
vesicle.
Fig. 8. Phenotypic characterization of the mac1; ocl4 double mutant. (A-D)Transverse section of anther lobes. (A)In the wild-type anther (500m), four cell types are present: the epidermis (EP), endothecium (EN), secondary parietal cells (SPC) and archesporial cells (AR). The EN cells are elongated and have started to accumulate starch granules. (B)In the ocl4 mutant, after L2-d cells have divided periclinally to form two layers, the outer layer divided periclinally again to form two EN-like cell layers. (C)In the mac1 mutant, excess AR cells are observed. An EN-like layer with starch granules forms from a few cell layers with unknown identity. (D)In the mac1; ocl4 double mutant, excess AR cells are present with unorganized somatic cells surrounding them. In addition, two EN-like layers form (blue dotted lines). The border of AR cells is evident in both wildtype (A) and the ocl4 mutant (B) (green dotted lines); however, it becomes disordered in the mac1 mutant (C) and mac1; ocl4 double mutant (D). (E-H)Confocal images of longitudinal optical sections of anthers. (E)A wild-type locule (500m anther) illustrates that EN and SPC have differentiated. (F)A 450m mac1 anther shows that excess AR cells enclosed by one layer of cells derived from L2 (L2-d) and EP. (G)A mac1; ocl4 double mutant anther (500m) shows an increased number of AR plus irregular patches of two somatic cell layers (arrowhead). (H)A mac1; ocl4 double mutant anther (1 mm) exhibits a burst of cell divisions as seen in mac1 mutant. Scale bars: 20m.