Keith R. Roberts

Light and electron microscopical observations on the type species of Gymnodinium, G. fuscum (Dinophyceae)

Abstract Gymnodinium fuscum (Ehrenberg) Stein is the type species of Gymnodinium, one of the largest genera of dinoflagellates. It has the typical fine structure of dinoflagellates, but also possesses several unusual features. The nuclear envelope is specialized by having vesicular chambers in which the nuclear pores are located; the pusular complex includes an internal collectjon chamber; and a nuclear fibrous connective links the nucleus with the longitudinal microtubular root of the flagellar apparatus. A delicate horseshoe-shaped apical groove is visible by scanning electron microscopy. The same morphological features have been observed in a few other gymnodinioids, indicating a close phylogenetic relationship, and have recently been used, together with large subunit (LSU) rDNA data, in redefining the genus Gymnodinium [Daugbjerg et al. (2000) Phycologia 39: 302–317]. However, one of the most unusual features of G. fuscum is the absence of a transverse striated flagellar root, a feature presently known only in the zoospores of the otherwise very different genus Noctiluca. This feature, together with the lack of striated collars around the flagellar canals, suggests a phylogenetically more isolated position for G. fuscum. Future investigations may reveal the need for further ‘splitting’ of the genus Gymnodinium.

THE FLAGELLAR APPARATUS OF CHILOMONAS PARAMECIUM (CRYPTOPHYCEAE) AND ITS COMPARISON WITH CERTAIN ZOOFLAGELLATES1

The major components of the internal flagellar apparatus of Chilomonas paramecium Ehr. are two large microtubular roots and a striated root paralleled by three microtubules. The two microtubular roots overlap at the basal bodies. One microtubular root follows a curved path in the anterior of the cell, and the other extends straight to the posterior passing through a groove in the nucleus. The striated root extends laterally from the basal bodies. Except that it is smaller, the posteriorly directed root bears a strong resemblance to the axostyle of oxymonads. The overall arrangement and structure of the flagellar roots is similar to the pelta, axostyle and costa of trichomonads and the pelta and axostyle of oxymonads, groups of mitochondrion‐less, largely parasitic or symbiotic protozoans. An affinity between cryptomonads and oxymonads or trichomonads would have many phylogenetic implications, some of which are discussed.

COMPARATIVE CYTOLOGY AND TAXONOMY OF THE ULVAPHYCEAE. I. THE ZOOSPORE OF ULOTHRIX ZONATA (CHLOROPHYTA)1

This fine structural study of the quadriflagellate zoospore of Ulothrix zonata (Weber & Mohr) Kützing, with special attention to the flagellar root system, demonstrates that it is very similar to the zoospore of Ulva lactuca L. in several aspects. Common features include the presence of a cruciate root system (4‐2‐4‐2 type), a non‐striated band that connects basal bodies, a so‐called terminal cap, and system I and system II striated root components. Only slight differences exist, i.e. in the shape of the terminal cap, and in the number and position of the system II root components. It is concluded that the taxonomic affinities of U. zonata lie with the Ulvaphyceae sensu Stewart and Mattox rather than with the Chlorophyceae. Additional support for this conclusion is the discovery of tiny, flat body scales on the zoospore of U. zonata. A summary of the distinctive characteristics of the Chlorophyceae, Charophyceae and Ulvaphyceae reflecting the current state of knowledge is given.

THE FLAGELLAR APPARATUS OF OXYRRHIS MARINA (PYRROPHYTA)1

The three‐dimensional structure of the flagellar apparatus in the dinoflagellate Oxyrrhis marina has been reinvestigated and found to consist of several previously unknown components and component combinations that appear strikingly similar to those of some gymnodinoid taxa.

STRUCTURE AND SIGNIFICANCE OF THE CRYPTOMONAD FLAGELLAR APPARATUS. I. CRYPTOMONAS OVATA (CRYPTOPHYTA)1

The absolute configuration of the flagellar apparatus in Cryptomonas ovata has been elucidated and found to be similar to that reported for Chilomonas paramecium. Variations apparent in the flagellar apparatus of Cryptomonas ovata include the presence of striations in the mitochondrion associated lamella, a rhizostyle which does not bear wing‐like extensions from the microtubules and does not lie close to the nucleus, and a striated fibrous anchoring structure associated with one basal body which has not hitherto been described. The flagellar apparatus also includes a four stranded microtubular root which traverses into the anterior dorsal lobe of the cell, a striated fibrous root which is associated with a five stranded microtubular root, and a two stranded Cr root. The homologous nature of these roots to those in the larger cryptomonads is discussed in relation to the apparent reduction in flagellar apparatus size and complexity among the smaller cryptomonads. A diagrammatic reconstruction of the flagellar apparatus of Cryptomonas ovata is also presented.

COMPARATIVE CYTOLOGY AND TAXONOMY OF THE ULVAPHYCEAE. III. THE FLAGELLAR APPARATUSES OF THE ANISOGAMETES OF DERBESIA TENUISSIMA (CHLOROPHYTA)1

The components of the flagellar apparatuses of the male and female gametes of Derbesia tenuissima (De Not.) Crouan are compared with those in other swarmers of green algae. Both the male and female gametes were found to have a cruciate microtubular root system, a non‐striated capping plate which connects basal bodies, two electron dense terminal caps which partially cover the proximal end of the basal bodies, and two small system II fibrous roots. Similarities exist between these components and those suggested to be typical of ulvalean swarmers. Based upon these similarities, it is proposed that the Caulerpales be classified in the Ulvaphyceae rather than in the Charophyceae or Chlorophyceae.

THE MICROTUBULAR CYTOSKELETON OF AMPHIDINIUM RHYNCHOCEPHALUM (DINOPHYCEAE)

The sub‐thecal microtubular cytoskeleton of Amphidinium rhynchocephalum Anissimowa was investigated using indirect immunofluorescence microscopy and transmission electron microscopy. The majority of sub‐thecal microtubules are longitudinally oriented and radiate from one of two sub‐thecal transverse microtubular bands that lie adjacent to the anterior and posterior edge of the cingulum.Both transverse bands consist of 3–5 microtubules and are loop shaped with one end adjacent to the cells right edge of the sulcus and the other end adjacent to the fibrous ventral ridge. The posterior transverse microtubular band (PTB) defines the posterior edge of the cingulum and gives rise to numerous posteriorly directed longitudinal microtubular bundles that consist of 1–3 microtubules per bundle. These bundles end at the posterior end of the cell. The PTB also gives rise to the cingular longitudinal microtubules that underlie the cingular groove and terminate at the anterior transverse microtubular band (ATB). The ATB defines the anterior edge of the cingulum and loops around the base of the epicone. This band gives rise to anteriorly directed longitudinal microtubular bundles that terminate in the small epicone of the cell. The longitudinal microtubular root of the flagellar apparatus is directed posteriorly and lies immediately beneath the theca but is distinct from the subthecal microtubule system. A narrow fibrous ridge is ventrally located to the cells left between the exit apertures of the transverse and longitudinal flagella. In this position, the ventral ridge lies between and also connects with the anterior and posterior transverse microtubular bands. The ventral ridge is also associated with three microtubules that are distinct from other cytoskeletal microtubules. Our results demonstrate that the majority of sub‐thecal microtubules originate from one of two microtubular bands associated with the cingulum. The possible role of the fibrous ventral ridge and its associated microtubules is also discussed.

STRUCTURE OF THE ANISOGAMETES OF THE GREEN SIPHON PSEUDOBRYOPSIS SP. (CHLOROPHYTA)1

The fine structure of the male and female gametes of Pseudobryopsis, particularly that of the flagellar apparatus, is compared with that of swarmers of other green algae. There is general similarity, with differences in detail, to the Ulvales and other green siphons that have been studied. The similarities include overlapping basal bodies, the capping plate type of connective between basal bodies, terminal caps, and system II fibrous roots (rhizoplasts). The capping plate of the female gamete differs from that in other green siphons and the Ulvales in form and in the presence of a faint striation. A diagram illustrating the actual arrangement of the components of the flagellar apparatus is given, along with a discussion of the fact that the mirror image of the true arrangement has been given in some reports on ulvaphycean algae.

Comparative cytology and taxonomy of theUlvaphyceae II. Ulvalean characteristics of the stephanokont flagellar apparatus ofDerbesia tenuissima

SummaryThe stephanokont flagellar apparatus of the zoospores ofDerbesia tenuissima (De Not.) Crouan is examined and compared to the flagellar apparatuses of other green algae. The flagella ofDerbesia are attached to two of three bands which lie at the junction of the body and papilla. Serial longitudinal and cross sections reveal that the basal bodies are attached to the bands along their sides and at their proximal ends. The bands are not striated in any plane. The lack of striation in the bands and the partial covering of the proximal end of the basal bodies by one of the bands closely resemble the type of flagellar connection system described as the “Bryopsis-type” byMelkonian (1980). Zoospores of ulvalean green algae also possess these features, suggesting that green siphons are phylogenetically related to theUlvales. It is proposed that green siphons be tentatively classified in theUlvaphyceae rather than in theChlorophyceae orCharophyceae.

COMPARATIVE ANALYSES OF THE DINOFLAGELLATE FLAGELLAR APPARATUS. III. FREEZE SUBSTITUTION OF AMPHIDINIUM RHYNCHOCEPHALUM1

The flagellar apparatus of the marine dinoflagellate Amphidinium rhynchocephalum Anissimowa was examined using the techniques of rapid freezing/freeze substitution and serial thin section three dimensional reconstruction. The flagellar apparatus is composed of two basal bodies that are offset from one another and lie at an angle of approximately 150° The transverse basal body is associated with two individual microtubules that extend from the proximal end of the basal body toward the flagellar opening. One of these microtubules is closely appressed to a striated fibrous root that also extends from the proximal base of the transverse basal body. The longitudinal basal body is associated with a nine member microtubular root that extends from the proximal end of the basal body toward the posterior of the cell. The longitudinal microtubular root and the transverse striated fiber are connected by a striated connective fiber. In addition to the microtubules associated with the transverse and longitudinal basal bodies, a group of microtubules originates adjacent to one of the transverse flagellar roots and extends into the cytoplasm. Vesicular channels extend from the flagellar openings to the region of the basal bodies where they expand to encompass the various connective structures of the flagellar apparatus. The possible function and evolutionary importance of these structures is discussed.