In many cases, changes in the normal colouring[887] are caused by the
breaking up of the acids on contact with atmospheric or soil ammonia.
Alkaline salts are thus formed which may be oxidized by the oxygen in
the air to yellow, red, brown, violet-brown or even to entirely black
humus-like products which are insoluble in water. These latter substances
are frequently to be found at the base of shrubby lichens or on the under
surface of leafy forms that are closely appressed to the substratum.
_c._ COLOUR DUE TO AMORPHOUS SUBSTANCES. These are the various pigments
which are deposited in the cell-walls of the hyphae. The only instance,
so far as is known, of colours within the cell occurs in _Baeomyces
roseus_, in which species the apothecia owe their rose-colour to
oil-drops in the cells of the paraphyses, and in _Lecidea coarctata_
where the spores are rose-coloured when young. In a few instances the
colouring matter is excreted (_Arthonia gregaria_ and _Diploschistes
ocellatus_); but Bachmann[888], who has made an extended study of this
subject and has examined 120 widely diversified lichens, found that with
few exceptions the pigment was in the membranes.
Bachmann was unable to determine whether the pigments were laid down
by the protoplasm or were due to changes in the cell-wall. The middle
layer, he found, was generally more deeply coloured than the inner one,
though that was not universal. In other cases the outer sheath was the
darkest, especially in cortices one to two cells thick such as those
of _Parmelia olivacea_, _P. fuliginosa_ and _P. revoluta_, and in the
brown thick-walled spores of _Physcia stellaris_ and of _Rhizocarpon
geographicum_. Still another variation occurs in _Parmelia tristis_ in
which the dark cortical cells show an outer colourless membrane over the
inner dark wall.
The coloured pigments are mainly to be found in the superficial tissues,
but if the thallus is split by areolation, as in crustaceous lichens, the
internal hyphae may be coloured like those of the outer cortex wherever
they are exposed. The hyphae of the gonidial layer are persistently
colourless, but the lower surface and the rhizoids of many foliose
lichens are frequently very deeply stained, as are the hypothalli of
crustaceous species.
The fruiting bodies in many different families of lichens have dark
coloured discs owing to the abundance of dark-brown pigment in the
paraphyses. In these the walls, as determined by Bachmann, are composed
generally of an inner wall, a second outer wall, and the outermost sheath
which forms the middle lamella between adjacent cells. In some species
the second wall is pigmented, in others the middle lamella is the one
deeply coloured. The hymenium of many apothecia and the hyphae forming
the amphithecium are often deeply impregnated with colour. The wall
hyphae of the pycnidia are also coloured in some forms; more frequently
the cells round the opening pore are more or less brown.
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