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Zea mays

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These vascular tissues occur nearer to the apex in<br />

slow growing or dormant roots than in actively<br />

elongaring roots. The firsr mature protOphloem<br />

elements appear at the circumference of the<br />

procambial cylindet and differentiate closer to the<br />

root apex than the first mature protOxylem (4.8).<br />

By contrast the prospective met'axylem becomes<br />

demarcated, by its prominent vacuolation growth<br />

(4.5), nearer the apex than the protophloem. Some<br />

distance basal ro the prorophloem, mature protoxylem<br />

elements develop between the prorophloem<br />

files. Maturation of the metaphloem and<br />

metaxylem proceeds centripetally (4.30) on the<br />

radii already demarcated by the protophloem and<br />

protoxylem (4.8), and the vascular system of the<br />

mature primary rOOt is formed (4.31). The outer<br />

layer of the procambium gives rise 10 the<br />

parenchymatous pcricyle (4.8,4.31).<br />

In dicotyledonous roots the tissue between the<br />

xylem and phloem often forms a cambium (4.31),<br />

which later spreads laterally over the protoxylem<br />

poles ro form a COntinuous meristem (1.25). The<br />

vascular cylinder is invested by the parenchymatous<br />

cortex (1.29, 4.30} which frequently<br />

contains conspicuous intercellular air spaces. The<br />

pericycle is bounded by the single layered<br />

endodermis (1.29, 4.31, 4.32) whose radial and<br />

transverse walls are impregnared with lignosuberin<br />

to form Casparian bands (4.33). These<br />

laterally continuous bands are impermeable, so<br />

that all water and solute movement across Ihis<br />

layer is confined to the symplast (4.33). In the<br />

roots of many species, particularly monocotyledons,<br />

the ligno-suberisation of the endodermis<br />

86<br />

later extends to all walls and additional cellulose<br />

thickening may be deposited (4.32). Such cells,<br />

however, still allow symplastic transport from rhe<br />

cortex ro the stele via their plasmodesmata.<br />

The endodermis is an important selective<br />

barrier but it allows the active transport into the<br />

vascular cylinder of certain beneficial ions<br />

(potassium, phosphare) absorbed from the soil.<br />

Calcium, however, apparently moves apoplastially<br />

and cannor corer the vascular cylinder through the<br />

endodermis, but instead passes into the vascular<br />

cylinder in the very immarure root where<br />

Casparian bands have not developed. The active<br />

transport of ions into the vascular cylinder<br />

accounts for the phenomenon of roOt pressute<br />

which sometimes plays an accessory role to<br />

transpiration in the movement of water to Ihe<br />

shoot.<br />

to some roots the ourer cortex diffetentiates as<br />

a one- ro several-layered exodermis (1.29) but the<br />

Casparian bands in the tadial walls are usually<br />

masked by deposition of subetised lamellae<br />

adjacent ro the protoplast (4.34). A short distance<br />

behind the apex a zone of absorptive root hairs<br />

develops from the epidetmis (4.27); but warer<br />

absorption also occurs over the test of the<br />

epidermal surface and in some species roar hairs<br />

are absent. In the aerial roots of some epiphytic<br />

orchids and aroids the multilayered epidermis<br />

develops into a dead velamen (4.34), whose walls<br />

are thickened by bands of cellulose. This is<br />

probably an adaption to absorb watet from the<br />

humid atmosphere of tropical forests.<br />

4.1 lu.s of the plwnule of Pht'lseo/us /lUfgaris (bean) seed.<br />

The hemispherical, densely-staining and small-eelled shoot<br />

apex (1) bears a pair of leaf primordia (2) at its margins<br />

which represent the first foliage leaves. A prominent strand<br />

of procambium (3) demarcates tbe longitudinal rows of<br />

vacuolated pith (4) from the cortex. (LM x 110.)<br />

1 Shoot apex<br />

2 Leaf primordium<br />

3 Procambium<br />

4 Pith

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