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CHAPTER 3<br />

Plant histology<br />

Disttibution of cells and tissues<br />

The vascular plant is composed of dermal, ground<br />

and vascular tissue systems (3.1 to 3.4). The<br />

phloem and xylem constitute the vascular system;<br />

both arc complex tissues in which the conducting<br />

elements (2.10, 2.68, 2.69) arc associated with<br />

other cell types (1.4, 1.11,3.5). The ground system<br />

surrounds the vascular tissues (3.1 to 3.4) and<br />

comprises parenchyma, collenchyma and selecenchyma<br />

(3.6 to 3.10). The dermal system is initially<br />

represented by the epidermis (3.2, 3.3) containing<br />

various cell types (3.3, 3.lt, 3.12). Its structure is<br />

considered in Chapter 5.<br />

In plants which undergo woody thickening, the<br />

epidermis typically becomes replaced by the cork<br />

and associated tissues (the periderm, 3.13) whose<br />

structure is discussed in Chapter 6. Secretory cells<br />

do not normally develop as distinct tissues (except,<br />

for example, in nectaries) but rather occur<br />

cither as surface structures (3_11, 3.12) or within<br />

other tissues (3.4).<br />

Parenchyma<br />

These cells arc generally thin-walled and have<br />

large vacuoles (2.24, 3.3. 3.6). Thcy form a<br />

continuous tissue in the cortex and pith (1.27, 3.4)<br />

and in the leaf mesophyll (3.3), while vertical<br />

strands and horizontal rays of parenchyma occur<br />

in the vascular system (1.3, 3.14). The leaves and<br />

stem of seedlings and small herbs are largely held<br />

erect by the collective turgor of the parenchyma<br />

cells (3.3) and the shoot wilts if insufficient water<br />

is available from the root system to replace<br />

transpiration losses (1 .12).<br />

Parenchyma cells are often polyhedral or<br />

elongate (3.3, 3.6) but stellate and irregular forms<br />

occur (3.3, 3.15). The mesophyll cell of the leaf is<br />

specialised for photosynthesis and contains numcrous<br />

chloroplasts (3.3) while starch is frequently<br />

stored in the parenchyma of the root and<br />

stem (t.14). In plants growing in marshy conditions<br />

and in aquatics, the parenchyma of the<br />

shoot and root sometimes develops extensive<br />

intetcellular spaces and is termed aerenchyma<br />

(3.15).<br />

Mature parenchyma ceJls of the stem and root<br />

may resume division (3.16) and undergo partial<br />

dedifferentiation to form the lareral meristems of<br />

the phellogen and vascular cambium (3.5) from<br />

which the cork and secondary vascular tissues<br />

originate (3.13, 3.17). In some situations the<br />

parenchyma cells revert to an undifferentiated<br />

state (as found in the apical meristems, 2.2) and<br />

give rise to new plant organs. This phenomenon<br />

results in the formation of lateral and adventitious<br />

roots (3.18) and in the development of primordia<br />

(3.19) which give rise to adventitious shoots on<br />

various partS of the plant (3.20, 3.21).<br />

The development of adventitious organs is<br />

sometimes related ro injury (3.18); the plant's<br />

regenerarive capacity is exploited in horticulture<br />

by the rooting of shoot cuttings (3.J8) and regeneration<br />

from various excised plant organs (3.J9 to<br />

3.21). Parenchyma cells should therefore be<br />

regarded as potentially totipotent and single cells<br />

derived from carrot and tobacco have given rise to<br />

completely new plants when cultured in vitro on a<br />

suitable nutrient medium with exogenous growth<br />

substances.<br />

The parenchyma cell wall is often thin, with irs<br />

growth normally terminating at the end of cell<br />

vacuolation (2.24, 3.3, 3.15)_ However, in seed<br />

srorage tissue the walls may be greatly thichned<br />

(Correa, Phoenix, Phaseo/lts. 2.54) by noncellulosic<br />

reserve carbohydrates, while parenchyma<br />

cells often develop thick lignified secondary<br />

walls (1.3, 2_65) in the xylem and pith. In transfer<br />

cells labyrinthine ingrowths of the wall (2.63)<br />

vastly increase the plasmalemmal surface, while<br />

simpler vertical ingrowths of the wall also occur in<br />

the plicate mcsophyH of several conifers (3.22).<br />

Collenchyma<br />

These living cells arc thick-walled but relatively<br />

pliable (3.7) and are located in the outer ground<br />

tissue (3.23). Collenchyma is of great value in the<br />

mechanical support of the young shoot but is rare<br />

in the root. The proroplasts of such cells are<br />

generally vacuolate (3.7) while their highly<br />

hydrated primary walls are unevenly thickened<br />

(3.7, 3.24) and comain large amounts of peCtins<br />

and hemicelluloscs. Collenchyma cells are usually<br />

elongate (3.7) and in transverse section often<br />

appear angular with thickening localised in the<br />

cell corners (3.7, 3.24) but the thickening may be<br />

confined to the tangential walls (3.7). Intercellular<br />

spaces sometimes occur in collenchyma and in<br />

older cells the walls may become lignified.<br />

57

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