_a_. HISTORICAL. The presence of spores, as such, in the lichen fruit was
first established by Hedwig[660] in _Anaptychia_ (_Physcia_) _ciliaris_.
He rightly judged the minute bodies to be the “semina” of the plant. In
that species they are fairly large, measuring about 50µ, long and 24µ
thick, and as they are very dark in colour when mature, they stand out
conspicuously from the surrounding colourless tissue of the hymenium.
Acharius[661] also took note of these “semina” and happily replaced the
term by that of “spores.” They may be produced, he states, in a compact
nucleus (_Sphaerophoron_), in a naked disc (_Calicium_), or they may be
embedded in the disc (_Opegrapha_ and _Lecidea_). Sprengel[662] opined
that the spores—which he figures—were true seeds, though he allows
that there had been no record of their development into new plants.
Luyken[663] made a further contribution to the subject by dividing
lichens into gymnocarpous and angiocarpous forms, according as the
spores, enclosed in cells or vesicles (thecae), were borne in an open
disc or in a closed perithecium.
In his _Systema_ of lichen genera Eschweiler[664], some years later,
described and figured the spores as “thecae” enclosed in cylindrical
asci. Fée[665] in contemporary works gave special prominence to the
colour and form of the spores in all the lichens dealt with.
_b_. DEVELOPMENT OF THE ASCUS. The first attempt to trace the origin
and development of lichen asci and spores was made by Mohl[666]. He
describes the mother-cell (the ascus) as filled at first with a clouded
granular substance changing later into a definite number—usually eight—of
simple or septate spores. Dangeard[667] included the lichens _Borrera_
(_Physcia_) _ciliaris_ and _Endocarpon_ (_Dermatocarpon_) _miniatam_
among the plants that he studied for ascus and spore development. He
found that in lichens, as in fungi, the ascus arose usually from the
penultimate cell of a crooked hypha (Fig. 104) and that it contained
at first two nuclei derived from adjoining cells. These nuclei are
similar in size to those of the vegetative hyphae, and in each there is
a large nucleolus with chromatin material massed on one side. Fusion
takes place, as in fungi, between the two nuclei, and the secondary or
definitive nucleus thus formed divides successively to form the eight
spore-nuclei. Baur[668] and Nienburg[669] have confirmed Dangeard’s
results as regards lichens, and René Maire[670] has also contributed
important cytological details on the development of the spores. In
_Anaptychia_ (_Physcia_) _ciliaris_ he found that the fused nucleus
became larger and that a synapsis stage supervened during which the long
slender chromatin filaments became paired, and at the same time shorter
and thicker. The nuclear membrane disappeared as the chromatin filaments
were united in masses joined together by linin threads which also
disappeared later. At the most advanced stage observed by Maire there
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