_b._ FORMATION OF CARPOGONIA. Stahl[607] had indicated that not only
in gymnocarpous but also in angiocarpous lichens, it would be found
that carpogonia were formed as in _Collema_. Baur[608] justified this
surmise, and demonstrated the presence of ascogonia in groups of three
to eight, with trichogynes that reached the surface in _Endocarpon_
(_Dermatocarpon_) _miniatum_ (Fig. 99). It is one of the few foliaceous
Pyrenolichens, and the leathery thallus is attached to the substratum
by a central point, thus allowing in the thallus not only peripheral
but also intercalary growth, the latter specially active round the
point of basal attachment; carpogonia may be found in any region where
the tissue is newly formed, and at any season. The upper cortex is
composed of short-celled thick-walled hyphae, with branching vertical
to the surface, and so closely packed that there is an appearance of
plectenchyma; the medullary hyphae are also thick-walled but with
longer cells. The carpogonia of this species arise as a branch from the
vegetative hyphae and are without special covering hyphae, so frequent
a feature in other lichens. The trichogynes bore their way through
the compact cortex and rise well above the surface. After they have
disappeared—presumably after fertilization—the vegetative hyphae round
and between the ascogonia become active and travel upwards slightly
converging to a central point. The asci begin to grow out from the
ascogenous hyphae of the base before the vertical filaments have quite
pierced the cortex.
_Pyrenula nitida_ has also been studied by Baur[609]. It is a very common
species on smooth bark, with a thin crustaceous thallus immersed among
the outer periderm cells. Unlike most other lichens, it forms carpogonia
in spring only, from February to April. A primordial coil of hyphae lies
at the base of the gonidial layer, and, before there is any appearance of
carpogonia, a thick strand of hyphae is seen to be directed upwards, so
that a definite form and direction is given to the perithecium at a very
early stage. The ascogonial cells which are differentiated are extremely
small, and, like those of all other species examined, are uninucleate.
There are five to ten carpogonia in each primordium; the trichogynes grow
up through the hyphal strand and emerge 5-10 µ above the surface. After
their disappearance, a weft of ascogenous tissue is formed at the base,
and, at the same time, the surrounding vegetative tissue takes part in
the building up of a plectenchymatous wall of minute dark-coloured cells.
Further development is rapid and occupies probably only a few weeks.
In many of the pyrenocarpous lichens—_Verrucariae_ and others—the walls
of the paraphyses dissolve in mucilage as the spores become mature, a
character associated with spore ejection and dispersal. In some genera
and species, as in _Pyrenula_, they remain intact.
D. APOGAMOUS REPRODUCTION
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