Chromatin condensation begins as the multilayered structure (MLS) in Phaeoceros reaches its maximally structured differentiation. As nuclear elongation and chromatin condensation proceed, the S2–4 strata disappear, and the nuclear beak extends between the S1 and the nearly spherical anterior mitochondrion. In a mature sperm the mitochondrion is elongate and lies completely anterior to the blunt front end of the nucleus. The 12 S1 tubules extend over the anterior mitochondrion and nucleus, but their number becomes reduced to five at the level where the nucleus' midportion is constricted. The anterior ends of the S1 tubules lie embedded in a rather conical osmiophilic crest. Flagellar insertion is restricted to the extreme anterior tip of the S1 tubules. The locomotory apparatus in Phaeoceros is compared with that of other bryophytes.
Electron microscopic examination of thin sections showed that the blepharoplast of a young spermatid of Phaeoceros consists of two side‐by‐side centrioles and an accumulation of osmiophilic, granular matrix at their proximal ends. Lying between these nearly parallel organelles is a dark‐staining body that will later disappear at the onset of flagellogenesis. For a brief period the centrioles are oriented perpendicular to the nuclear surface so that the granular matrix at their proximal ends is confluent with the nuclear envelope; furthermore, the nucleoplasm immediately in front of the centrioles becomes densely staining. The multilayered structure (MLS) develops directly under the centrioles. It comprises a band of 12 microtubules (the S1 stratum) and three lower strata (S2–4) whose constitutent lamellae are oriented at an oblique angle to the S1 axis. While the S1 tubules grow rearward over the nucleus which forms a beak adjacent to the posterior end of the lamellar strata, the centrioles are transformed into basal bodies with the distal growth of the axonemes and the proximal growth of the central cartwheels and lowermost triplets. The proximal ends of the basal bodies and the S1 tubules overlying the lamellar strata are invested with osmiophilic matrix that extends down to the S2 layer and may temporarily occlude the lamellar plates. At the onset of nuclear elongation an anterior mitochondrion becomes situated close beneath the lamellar strata which extend laterally beyond the S1 tubules.
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