Hedlund followed the process of association between the two organisms
in the lichens _Micarea_ (_Biatorina_) _prasina_ and _M. denigrata_
(_Biatorina synothea_), crustaceous species which inhabit trunks of trees
or palings. In these the alga, one of the Chlorophyceae, has assumed the
character of a _Gloeocapsa_ but on cultivation it was found to belong
to the genus _Gloeocystis_. The cells are globose and rather small;
they increase by the division of the contents into two or at most four
portions which become rounded off and covered with a membrane before
they become free from the mother-cell. The lichen hypha, on contact with
any one of the green cells, bores through the outer membrane and swells
within to a haustorium, as in the gonidia of _Synalissa_.
[Illustration: Fig. 11. Gonidia from _Ramalina reticulata_ Nyl. A,
gonidium pierced and cell contents shrinking × 560; B, older stage, the
contents of gonidium exhausted × 900 (after Peirce).]
[Illustration: Fig. 12. _Pertusaria globulifera_ Nyl. Fungus and gonidia
from gonidial zone × 500 (after Darbishire).]
Penetrating haustoria were demonstrated by Peirce[213] in his study of
the gonidia of _Ramalina reticulata_. In the first stage the tip of a
hypha had pierced the outer wall of the alga, causing the protoplasm to
contract away from the point of contact (Fig. 11). More advanced stages
showed the extension of the haustorium into the centre of the cell, and,
finally, the complete disappearance of the contents. In many cases it was
found that penetration equally with clasping of the alga by the filament
sets up an irritation which induces cell-division, and the alga, as in
_Synalissa_, thus becomes free from the fungus. Hue[214] has recorded
instances of penetration in an Antarctic species, _Physcia puncticulata_.
It is easy, he says, to see the tips of the hyphae pierce the sheath of
the gonidium and penetrate to the nucleus.
Lindau[215] has described the association between fungus and alga in
_Pertusaria_ and other crustaceous forms as one of contact only (Fig.
12). He found that the cell-membrane of the two adhering organisms was
unbroken. Occasionally the algal cell showed a slight indentation, but
was otherwise unchanged. The hyphal branch was somewhat swollen at the
tip where it touched the alga, and the wall was slightly thinner. The
attachment between the two cells was so close, however, that pressure on
the cover-glass failed to separate them.
Generally the hypha simply surrounds the gonidium with clasping branches.
Many algae also lie free in the gonidial zone, and Peirce[216] claims
that these are larger, more deeply coloured and in every way healthier
looking than those in the grasp of the fungus. He ignores, however, the
case of the soredial algae which though very closely invested by the
fungus are yet entirely healthy, since on their future increase depends
in many cases the reproduction of new individual lichens.
Public-domain text, read in full here on John Shaqi.
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