Apothecium Development

Germination of ascospores produce septate mycelium that is monokaryotic or homokaryotic. This mycelium will produce ascogonium and antheridium (Fig. 1), which may be designated as the female and male gametangia, respectively. Each gametangium is multinucelate and transfer of the nuclei from the antheridium to the ascogonium is made possible by the formation of a trichogyne, a tubular structure originating from the ascogonium that fuses with the antheridium. Once migration has occurred, each nucleus from the antherdium pairs off with one from the ascogonium (Fig. 2).
 
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Figure 1: Nuclei from the antheridium are transported into the ascogonium through the trichogyne , which originates from the ascogonium. Figure 2: Migration of nuclei from ascogonium to antheridium, via the trichogyne, is completed and nuclei from antheridum are paired paired off with those from the ascogonium. 

Once the nuclei have paired off, ascogenous hyphae form from the ascogonium and the paired nuclei undergo congugate mitotic division. Thus, the ascogenous hyphae are dikaryotic. This binucleate stage is prolonged and does not undergo karyogamy as in other life cycles that you have observed. At the same time the stalk cell is producing interwoven monokaryotic mycelium that will make up most of the apothecium and the paraphyses.


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Figure 3. Formation of ascogenous hyphae, which is a proloned binucleate stage and will be the cells that eventually give rise to the asci and ascospores. Stalk cell produce sterile cells of the ascocarp and paraphyses.



The ascogoenous hyphae grow and branch, repeatedly until the tips (Fig. 4a) of the ascogenous cells reach the site where the asci and ascospores will be produced. The tips of these ascogenous cells will elongate (Fig. 4b) and become recurved to form "hooks" or croziers (Fig. 4c). This is followed by the simultaneous division of the two nuclei in this cell (Fig. 4c) and division of the crozier into an apical cell, the recurved part of the hyphal tip, which is uninucleate and the penultimate cell, the cell immediately behind the apical cell (Fig. 4d). It is here that the karyogamy and meiosis occurs, and where the asci and ascospores will develop. Below the penultimate cell is the uninucleate stalk cell (Fig. 4d).
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Figure 4a-d: Crozier formation. a. Hyphal tip of ascogenous cell; b. Elongation of hyphal tip; c. Hyphal tip recurves forming crozier; d. Simultaneous division of nuclei and division of crozier into uninucleate apical and binucleate penultimate cells. Stalk cell below penultimate cell is also uninucleate.
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