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Zea mays

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stack (2.14, 2.52) of smooth cisternae 1-2 I-lm in<br />

diameter. Their margins arc frequemly branched<br />

imo a tubular network (2.51) which proliferates a<br />

number of small vesicles. The cisternae are<br />

separated from each other by about 10 nm, but<br />

individual dictyosomes remain intact when<br />

isolated from the cell.<br />

In the longitudinal view of a dictyosome a<br />

polarity is sometimes evident (2.52). At its<br />

forming face the cisternae are thought to be<br />

reconstituted from vesicles budded off from<br />

adjacent membranes of the ER (2.14,2.52). The<br />

cisternal membranes progressively increase in<br />

thickness across the dictyosome and at irs maturing<br />

face the cisternae frequently become concave,<br />

witb vesicles budding off from their margins<br />

(2.14,2.51,2.52).<br />

The vesicles apparently migrate and fuse with<br />

the plasmalemma where the contents are voided<br />

into the apoplasr. Within the dict)'osome, proteins<br />

derived from the RER combine with various<br />

sugars; the secreted vesicles contain carbohydrates<br />

and glycoproteins concerned with cell wall synthesis<br />

(2.8), mucilage (2.19), or nectar secretion.<br />

Dicryosomes are particularly abundant in cells<br />

actively undergoing extensive wall thickening or<br />

forming new dividing walls (2.8, 2.53); in root cap<br />

cells, where cisternae often become inflated with<br />

mucilage and sloughed off whole (2.19); and in<br />

glandular cells. In actively secreting cells a vast<br />

pOtential excess of membranous components<br />

(derived from the dicryosomes) arrives at the plasmalemma,<br />

but some of this material is apparently<br />

returned to the cytoplasm in the 'coated' vesicles<br />

(2.14).<br />

Vacuole<br />

1n the meristematic cell a number of small<br />

vacuoles occur, each bounded by the membranous<br />

tonoplast (2.2, 2.4, 2.13). However, during cell<br />

growth these vacuoles massively enlarge and fuse<br />

so that up to 90% of the volume in a parenchyma<br />

cell is occupied by the vacuole (1.18), "while the<br />

cytoplasm and other organelles are mainly<br />

peripheral (2.24, 2.42). The vacuole contains<br />

various solutes (normally at aboUT O.S M concentration)<br />

and its consequent turgidity greatly contributes<br />

to the [Urgor of the whole protoplast.<br />

The vacuole also contains a number of hydrolytic<br />

enzymes and the tonoplast breaks dmvn during<br />

differentiation of sclercnchyma and tracheary<br />

cells (2.7 to 2.9). The enzymes which are liberated<br />

digest the proroplasts so that only the walls<br />

remain intact (lA, 1.11). Vacuoles may contain<br />

anthocyanins and other pigments and also become<br />

modified as protein bodies in a number of seeds<br />

(2.43,2.54).<br />

Microbodies<br />

These small, membrane-bounded bodies are of<br />

two types. Pcroxysomes occur in close proximity<br />

to chloroplasts (2.38, 2.39); they contain a variety<br />

of enzymes which oxidise the glycolic acid<br />

resulting from photorespiration and then return<br />

glyoxylic acid to the chloroplasts. Glyoxysomes<br />

occur in fatty seeds and contain enzymes catalysing<br />

fatty acid breakdown to release energy<br />

during germination, while the hydrogen peroxide<br />

produced is broken down by peroxidase.<br />

Ribosomes<br />

The cytoplasmic ribosomes are 17-20 nm wide<br />

and occur both floating free within the cytosol<br />

(cytoplasmic ground substance) as well as<br />

attached to the outer surfaces of the RER (2.48,<br />

2.50 and 2.51). Ribosomes contain RNA and<br />

protein and these are composed of two sub-units<br />

which are synthesised in the nucleolus (2.26) bur<br />

subsequentiy combine in the cytoplasm. Ribosomes<br />

arc especially dense in cells which are<br />

rapidly synthesising protein where they frequently<br />

occur in clusters termed polysomes (2.48). The<br />

cytoplasmic ribosomes (80S) are slightly larger<br />

and biochemically distinct from those located in<br />

the plastids and mitochondria (70S).<br />

Microtubules and microfilaments<br />

The microtubules are proteinaceous structures<br />

about 25 nm wide, with a hollow core 12 nm<br />

wide, and may be up to several micrometers long<br />

(2.55). In the non-dividing cell they are normally<br />

located adjacent to the plasmalemma (2.2, 2.55<br />

and 2.56), bur at nuclear division these become<br />

reassembled and aggregate into the fibres of the<br />

mitotic spindle (2.28, 2.56), When mitosis is<br />

completed, the microtubules apparently guide<br />

dictyosome vesicles to the equatorial region of the<br />

spindle where the vesicles fuse to form the cell<br />

plate (2.53. 2.57, 2.58). It has been suggested that<br />

the peripheral microtubules arc concerned with<br />

the orientation of the cellulose microfibrils which<br />

are being formed in the young wall on rhe outside<br />

of the plasmalemma (2.16, 2.17); however, the<br />

evidence for [his is equivocal.<br />

Another smaller type of proteinaceous element<br />

has sometimes been observed in [he plant cell.<br />

This is termed a microfilament and is abom 7 run<br />

in width. In the green alga Nite/la, the micro·<br />

filaments a.re concerned with cytoplasmic streaming;<br />

in pollen tubes of flowering plants they<br />

apparentl)' guide vesicles concerned with wall<br />

synthesis through the cytosol to the growing tip of<br />

the tube.<br />

29

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