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Narcissus and Daffodil

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The biology of <strong>Narcissus</strong> 13<br />

The flower stalk is a leafless single internode (or scape), which contains abundant<br />

starch grains <strong>and</strong> raphides of calcium oxalate (Chan, 1952). The scape is topped<br />

by a single flower or a cyme, the receptacle of which bends to bring the flower into<br />

a horizontal position for anthesis <strong>and</strong> straightens after fertilisation to hold the<br />

capsule upright, a tropic response to gravity accentuated by light. Flowers also<br />

turn towards the light when growing in clumps (Rees, 1988). After flowers have<br />

been picked, the lower part of the stem that is left behind is capable of considerable<br />

growth from its basal meristem.<br />

The flower bud is enclosed in a protective sheath or spathe, which is green during<br />

bud development <strong>and</strong> dries <strong>and</strong> splits before anthesis. Flower development in<br />

narcissus was described <strong>and</strong> reviewed by Huisman <strong>and</strong> Hartsema (1933) <strong>and</strong> Chan<br />

(1952). The flower consists of two whorls of three perianth segments (or tepals)<br />

which arise from a hypanthial tube, two whorls of three anthers, <strong>and</strong> a tricarpellate<br />

inferior ovary with two rows of anatropous ovules in each of three loculi. The<br />

corona (paracorolla, trumpet or cup), characteristic of the genus, is conspicuous<br />

between the perianth segments <strong>and</strong> the anthers (Webb, 1980); it is regarded as<br />

either a prolongation of the receptacle or as a distinct structure (Huisman <strong>and</strong><br />

Hartsema, 1933; Guédès, 1966). The floral parts are formed in a spiral sequence<br />

from the perianth inwards to the gynoecium, although for horticultural purposes<br />

the completion of flower initiation (important as a key stage in beginning cold<br />

storage or hot-water treatment) is regarded as when the corona initial can be<br />

clearly seen on dissection, following the initiation of the gynoecium (Huisman <strong>and</strong><br />

Hartsema, 1933; Cremer et al., 1974). There are many varieties with double flowers,<br />

the structure of which varies in complexity (Reynolds <strong>and</strong> Tampion, 1983). Gross<br />

flower abnormalities (such as two flowers from the same apex) are rare (Chan,<br />

1952), although physiological disorders may occur in flower development (Rees,<br />

1972). The floral biology of N. pseudonarcissus has been described in detail by Caldwell<br />

<strong>and</strong> Wallace (1955). The corona <strong>and</strong> perianth tube lead to nectaries between<br />

the staminal filaments, the flowers being pollinated by bumble bees (Bombus spp.).<br />

Pollination is often poor due to a lack of insects in the relatively cool conditions,<br />

<strong>and</strong> little pollination occurs in the absence of insects. Pollination of N. longispathus<br />

in Spain was studied by Herrera (1995); the temperature inside the flower was up<br />

to 8 °C above ambient temperatures, raising the temperature of pollinating bees<br />

sufficient for flight. Di- <strong>and</strong> polymorphism with respect to style length has been<br />

reported for N. tazetta <strong>and</strong> N. tri<strong>and</strong>rus, respectively (Arroyo <strong>and</strong> Dafni, 1995; Barrett<br />

et al., 1997). The fruits are ovoid green pods which grow to full size in about two<br />

weeks <strong>and</strong> dehisce a few weeks later. The seeds are black <strong>and</strong> round <strong>and</strong> are<br />

distributed over a restricted area; seeds of commercial narcissus cultivars were<br />

described by Chan (1952).<br />

Roots<br />

The roots of narcissus are adventitious, unbranched, with a prominent root cap,<br />

<strong>and</strong> are generally considered to have no root hairs, although a few poorly<br />

developed root hairs can occur in some circumstances (Chan, 1952; Rees, 1972;<br />

Kawa <strong>and</strong> De Hertogh, 1992). However, Chilvers <strong>and</strong> Daft (1981) found root hairs<br />

in all cultivars examined, while Wilson <strong>and</strong> Peterson (1982) described the roots of<br />

N. lobularis as initially glabrous <strong>and</strong> later with hairs. Price (1977) examined the

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