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Introduction to Fungi, Third Edition

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STRUCTURE AND MORPHOGENESIS OF BASIDIOCARPS<br />

523<br />

Fig19.8 Modes of gill attachment <strong>to</strong> the stipe, and<br />

their terminology.<br />

of gill maturation begins at the cap margin and<br />

passes slowly upwards and inwards. After the<br />

basidia at the lower edge of the gill have discharged<br />

their spores, the gill tissue undergoes<br />

au<strong>to</strong>lysis (deliquescence) in<strong>to</strong> an inky black<br />

liquid which drips away from the cap. The gravitropic<br />

gill curvature characteristic of aequihymenial<br />

types is absent, but the stipe may still<br />

curve <strong>to</strong> bring the gills in<strong>to</strong> an approximately<br />

vertical position.<br />

An important criterion for identification of<br />

gill-bearing agarics in the field is the way in<br />

which the lamellae are attached <strong>to</strong> the stipe<br />

(Fig. 19.8). Gills are said <strong>to</strong> be free if their blade<br />

does not <strong>to</strong>uch the stipe. Adnate gills show<br />

attachment <strong>to</strong> the stipe with their entire base,<br />

whereas adnexed gills are attached only partially<br />

<strong>to</strong> the stipe. In sinuate gills, the gill margin<br />

shows an S-shaped curve near the point of<br />

junction with the stipe. Decurrent gills run<br />

down the surface of the stipe.<br />

19.2.4 The hymenophoral trama<br />

The hymenophore structures of some<br />

Homobasidiomycetes are shown in Fig. 19.9. In<br />

most hymenophores, there is a central group of<br />

hyphae running from the underside of the cap <strong>to</strong><br />

the tip of the gill, pore or spine, and these<br />

hyphae are collectively called the hymenophoral<br />

trama. Various distinctive types have been<br />

recognized by Reijnders and Stalpers (1992), but<br />

these do not correlate well with the phylogeny of<br />

the Homobasidiomycetes as shown in Fig. 19.2.<br />

1. In the trame<strong>to</strong>id type (e.g. in Lentinus,<br />

Fistulina and Schizophyllum), development begins<br />

with a bundle of parallel hyphae from which<br />

branches grow out in various directions and may<br />

become interwoven.<br />

2. In the cantharelloid type, the developing<br />

hymenophore is at first smooth and covered by a<br />

palisade of hyphae which will form<br />

the hymenium. Later, locally increased<br />

activity in the subhymenium may result<br />

in irregular ridges (Serpula), more regular veins<br />

(Cantharellus, Craterellus) or gills (Hygrophoropsis).<br />

3. The bole<strong>to</strong>id type is shown by Boletus and<br />

relatives (Fig. 19.9a). The first stage is made up of<br />

divergent hyphae which form a narrow central<br />

layer (mediostratum), followed by swelling and<br />

differentiation of the cells <strong>to</strong> form a lateral<br />

stratum. The diverging hyphae of the lateral<br />

strata curve sharply outwards <strong>to</strong> form the<br />

hymenium, and the term divergent trama is<br />

therefore used <strong>to</strong> describe this type of hymenophore.<br />

A subhymenium made up of narrow<br />

interwoven hyphae is also present.<br />

4. The agaricoid type is by far the most<br />

common and includes the coprinoid, russuloid,<br />

agaricoid, pluteoid and amani<strong>to</strong>id subtypes. In<br />

cross-section, gills usually appear differentiated<br />

in<strong>to</strong> a central trama, a subhymenium and the<br />

hymenium (Fig. 19.9b). The hyphae in the trama<br />

often run parallel <strong>to</strong> each other, but in the<br />

pluteoid subtype (e.g. Volvariella and Pluteus), the<br />

wide hyphae making up the trama are arranged<br />

in a V-shaped pattern (Fig. 19.9d), and this is<br />

called the inverted trama. In the russuloid<br />

subtype (Russula and Lactarius), the mature<br />

trama is said <strong>to</strong> be intermixed because in<br />

addition <strong>to</strong> the narrow generative hyphae it<br />

contains swollen cells (sphaerocysts) of very<br />

different width (Fig. 19.9c). The amani<strong>to</strong>id type<br />

shows a central mediostratum made up of<br />

narrow generative hyphae from which wider<br />

subhymenial elements diverge (Fig. 19.9e). This<br />

type of trama is described as bilateral.

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