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76                                    Algae: Anatomy, Biochemistry, and Biotechnology

                     The ultrastructure of the flagellar root system has been described only in one species of Bacillar-
                  iophyceae, Biddulphia laevis. In the spindle-shaped spermatoids of this centric diatom a system of
                  microtubules radiates from the only basal body present into the cell, to form a cone on the anterior
                  part of the nucleus, and at least some of these microtubules extend throughout the length of the
                  cell. They appear to maintain the elongated shape of the nucleus, and thus probably of the cell.
                     An elaborate root system connects the basal bodies to the anterior surface of the nucleus in the
                  Raphidophyceae. In Chattonella two or three roots have been described, one multi-layered associ-
                  ated with the pleuronematic flagellum, and one with the smooth flagellum. The latter is a band of
                  9–10 microtubules which extends from the anterior part of the cell, passing between the basal
                  bodies, and joining with other microtubules before extending along the nucleus in a depression
                  of its surface. A large number of distinctly cross-banded fibrous roots are also present, extending
                  from the flagellar bases to ensheathe the anterior cone of the nucleus.
                     In the Phaeophyceae the structure of the flagellar root system appears remarkably uniform. The
                  main characteristic of the system is the root supporting the flat, elastic proboscis in spermatozoids
                  of brown algae. In Fucus this root is very broad, contains usually 7 microtubules (but there can be
                  up to 15 microtubules), which travel from the base of the anterior flagellum along the plasmalemma
                  to the anterior end of the cell, where the proboscis is located. Here, they bend back and run along the
                  cell membrane on the opposite side. A bypassing microtubular root, of about five microtubules,
                  originates in the anterior part of the cell, runs along the proboscis root, bypasses the basal
                  bodies, without contact, and continues towards the posterior part of the cell. Other two minor micro-
                  tubular roots, consisting of only one microtubule, are present, one extending anteriorly, the other
                  posteriorly. The basal bodies are interlinked by three cross-banded connectives, the deltoid, the
                  strap-shaped, and the button-shaped bands (Figure 2.50).

                  Haptophyta
                  In the Haptophyta there are different types of flagellar roots. Members of the order Pavlovales, such
                  as Pavlova and Diacronema, possess a fibrous root, non-striated, which extends from the base of the
                  anterior flagellum, and passes into the cell along the inner face of the nucleus, becoming progress-
                  ively wider. In some species, another fibrous root originates at the base of the haptonema. Two
                  microtubular roots extend from the base of the posterior flagellum: a seven-stranded root which
                  runs under the periplast, and a two-stranded root arising almost at right angles to the seven-stranded
                  root, running inside the cell opposite to the haptonema. Fibrous connecting bands are present
                  between the basal bodies (Figure 2.51). In the algae of the order Isochrysidales, such as Pleurochry-
                  sis, the structure of the flagellar root system is more complex. Three main microtubular roots are
                  associated with the two basal bodies, two broad roots, no. 1 and no. 2, arising near the left flagellum,
                  and a smaller root, no. 3, arising near the right flagellum. Root no. 1 extends from the basal body up
                  towards the cell apex and then curves backwards to run inside the cell. A fibrous root is associated
                  with root no. 1. Closely packed microtubules organized in a bundle branch off perpendicularly from
                  both root no. 1 and root no. 2. The basal bodies are connected to each other by distal, intermediate,
                  and proximal connecting bands. Accessory connecting bands link the haptonema to the basal bodies
                  and the left basal body to the broad microtubular root no. 1 (Figure 2.52).

                  Cryptophyta
                  The flagellar roots of these algae include two characteristic components, the rhizostyle and the com-
                  pound rootlet. The rhizostyle is a posteriorly directed microtubular structure. It originates alongside
                  the basal body of the dorsal flagellum, extends deep into the cell, parallel to the gullet, behind the
                  layer of trichocysts, and ends in the posterior part of the cell. On the way, it runs through a groove in
                  the nuclear surface. In many cryptomonads, such as Chilomonas, each of the rhizostyle microtubule
                  bears a wing-like lamellar projection. The compound rootlet consists of a cross-banded fibrous band
                  and microtubular roots. The fibrous band originates from the basal bodies of the dorsal flagellum,
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