FURTHER EVIDENCE FOR A MEMBRANE-BOUND ENDOSYMBIONT WITHIN THE DINOFLAGELLATE PERIDINIUM FOLIACEUM1

The fine structure of the binucleate, fucoxanthin-containing dinoflagellate Peridinium foliaceum (Stein) Biechler was re-examined for evidence of an endosymbiout. The eucaryotic nucleus, chloroplasts and associated ribosome-dense cytoplasm were separated by a single invaginating membrane from the rest of the dinoflagellate cell. The triple membrane-enclosed eyespot, mesocaryotic nucleus, trichocysts and accumulation bodies resided in the dinoflagellate cytoplasm. These observations suggest that P. foliaceum contains a membrane-bound endosymbiont, similar to that already described for the closely related species. P. balticum (Levander) Lemmermann.     

NUCLEAR BASIC PROTEINS FROM THE BINUCLEATE DINOFLAGELLATE PERIDINIUM FOLIACEUM (PYRROPHYTA)1

Histories of the endosymbiont nucleus of the binucleate dinoflagellate Peridinium foliaceum Stein were prepared from isolated nuclei and analyzed by peptide mapping, ammo acid composition, and two-dimensional gel electrophoresis. Using these criteria, we identified the presence of two HI-like histories and the core histones H3, H2A, H2B, and H4. These histones are similar but not identical to those of the endosymbiont nucleus of the bi-nucleate dinoflagellate Peridinium balticum Levander.     

THE CALCAREOUS RESTING CYST OF PENTAPHARSODINIUM TYRRHENICUM COMB. NOV. (DINOPHYCEAE)1

While investigating dinoflagellate cyst assemblages in surface sediments of the Gulfs of Naples and Salerno (Mediterranean Sea), we found a new calcareous resting cyst. This cyst has a round to oval body surrounded by a thick mineral layer, which gives it the shape of a Napoleon hat, with a flat, oval face bearing the archeopyle and a convex keel on the opposite side. The cyst shape is variable in both natural samples and clonal cultures. The organic membrane underlying the calcareous covering is resistant to acetolysis, thus demonstrating the presence of sporopolleninlike material. The cyst germinated into a motile stage having the same morphological features and thecal plate pattern as Peridinium tyrrhenicum Balech. We believe the validity of the genus Pentapharsodinium Indelicato & Loeblich should be accepted. Based on the comparative examination of the species we collected and of a similar species, Pentapharsodinium trachodium Indelicato & Loeblich, we propose Pentapharsodinium tyrrhenicum as a new combination for Peridinium tyrrhenicum. The genus Pentapharsodinium also includes P. dalei Indelicato & Loeblich (= Peridinium faeroense Dale), which produces spiny, organic-walled cysts. The presence of species forming calcareous cysts and species producing noncalcareous cysts in the same genus raises questions about maintaining the family Calciodinellaceae. This family should only include calcareous cyst-forming peridinioids, in order to maintain a unified system of classification of fossil and recent dinoflagellates.     

Fitoplanctonti Raccolti in Occasione Della Moria di Pesci Verificatasi Lungo la Costa Marchigiano-Romagnola Nel Maggio 1969

Abstract

Phytoplankton collected during the fishes death occured off the western middle Adriatic coasts in May 1969. — A taxonomic inventory of some phytoplankton samples taken during a « red bloom » occured off the Italian Adriatic coasts from Cattolica to Cervia is given and discussed. The most common species was Peridinium depressum Bailey. It is also noticeable the presence of many Ceratium species and of Oxytoxum sceptrum (Stein) Schroder, a dinoflagellate rare for the Adriatic sea. The study of hydrological data and the comparison of this list with that of the species collected six days before, 50 miles southward, let us suppose that, for an anomalous circulation of waters with mediterranean characteristic, a stock of tipic pelagic species were transported very close to our coasts. Particular favourable conditions allowed big development of Peridinium depressum and its metabolism was responsible of the Oxygen deficency that was noted and was the cause of the fishes' death.     

PIGMENTS OF THE DINOFLAGELLATE PERIDINIUM BALTICUM AND ITS PHOTOSYNTHETIC ENDOSYMBIONT1

An examination of the pigments of the binucleate dinoflagellate Peridinium balticum (Levander) Lemmerman revealed the presence of chlorophylls a, c₁ and c₂ and the carotenoids: fucoxanthin (most abundant), diadinoxanthin, diatoxanthin, an unidentified fucoxanthin-like xanthophyll, β-carotene, γ-carotene and astaxanthin. A comparison of the pigments of P. balticum and P. foliaceum (Stein) Biecheler, also a binucleate dinoflagellate, demonstrated similar compositions. However P. balticum lacked the β-carotene precursors (e.g. phytoene) which accumulated outside the chloroplast in P. foliaceum. This study indicates that P. balticum and P. foliaceum are closely related; each species is a heterotrophic dinoflagellate with a photosynthetic endosymbiont taxonomically affiliated with the Chrysophyta (Chrysophyceae or Bacillariophyceae).     

Four new dinoflagellates

Fragilidium fissile is a new species of this rare genus. It somewhat resemblesF. subglobosum. It differs from the latter in having a slot and a pore in the first apical plate 1′ (the nomenclature of dinoflagellate plate designation follows the Kofoid system). Both species are also distinguishable by noticeable differences in Po, 1″″ and 1‴.Peridinium tyrrhenicum is a small species differing from all the other known species ofPeridinium in its shape, apical channel and several plates, especially some of the sulcal components.Alexandrium foedum somewhat resemblesA. balechii, but it differs from the latter in that its width is greater than its length, and in the characters of all the main sulcal plates. The above listed species were found in a sample from the Tyrrhenian Sea. The fourth species,Alexandrium andersoni, is a small dinoflagellate obtained in coastal waters off Cape Cod. It differs from all the other species of the minutum group in the very typical shapes of both the 6″ and the S. s.Pentapharsodinium daleii Indelicato and Loeblich is transferred toPeridinium.     

OBSERVATIONS ON THE SYMBIOSIS OF PERIDINIUM BALTICUM AND ITS INTRACELLULAR ALGA. I. ULTRASTRUCTURE1,2

The ultrastructure of Peridinium balticum indicates that an eudosymbiont is present within the dinoflagellate. The structure and probable function of the organelles of both protists arc described and discussed. The origin of photosynthetic dinoflagellates from heterotrophic forms by means of a chrysophyte-like endosymbiont is postulated.     

Spatial and temporal variations of Dinophyceae in subtropical reservoirs in southern Brazil

Knowledge of dinoflagellate diversity in Brazilian reservoirs is limited, especially in subtropical environments. We investigated as to how nutrients and other environmental variables influenced the biomass of Dinophyceae species in three subtropical ecosystems. The reservoirs Samuara, Faxinal, and São Miguel were sampled fortnightly from 2002 to 2006, and eight dinoflagellate taxa were identified. High temperature was a determining factor for the occurrence of Peridinium africanum Lemmermann. Peridinium umbonatum Stein and P. willei Huitfeld-Kass required high concentrations of nutrients. P. willei was inversely related to temperature and directly related to nutrients. P. umbonatum Stein var. umbonatum Stein showed the largest range of tolerance toward resources. Durinskia baltica Carty &; Cox and Peridinium gatunense Nygaard could be opportunistic, since they did not show any spatial or temporal pattern.     

A TECHNIQUE FOR COUNTING CHROMOSOMES OF ARMORED DINOFLAGELLATES,AND CHROMOSOME NUMBERS OF SIX FRESHWATER DINOFLAGELLATE SPECIES

A technique is presented to stain, squash, and enumerate thecate dinoflagellate chromosomes using a cellulose incubation and propionocarmine stain. Chromosome numbers for six freshwater armored dinoflagellates (Peridinium cinctum (O.F.M.) Ehrenberg, P. inconspicuum Lemm., P. limbatum (Stokes) Lemm., P. volzii Lemm., P. willei Huit.-Kaas, and Peridiniopsis polonicum (Wolosz.) Bourrelly) range from 41 (P. inconspicuum) to 210 (P. cinctum). Evidence is presented for dinoflagellate aneuploidy in culture.     

Isolation and Chemical Composition of Dinoflagellate Nuclei*

Nuclei were isolated from Peridinium cinctum. Peridinium trochoideum, Gyrodinium cohnii (Cryptothecodinium cohnii) and Gymnodinium nelsoni. The nuclei of G. cohnii, P. trochoideum and G. nelsoni were found to contain ～ 6.9, 34 and 143 picograms of DNA respectively. The ratios to DNA of RNA, acid-soluble and acid-insoluble protein for G. cohnii were 0.32, 0.13 and 0.99, respectively. The corresponding values for P. trochoideum were 0.22, 0.08 and 1.22, while those for G. nelsoni were 0.21, 0.10 and 1.09. The chemical composition of dinoflagellate nuclei is compared and contrasted with that of typical eukaryotic nuclei. The culture of these difficult organisms also is discussed.     

Dinoflagellate resting cysts as factors in phytoplankton ecology of the North Sea

The occurrence and distribution of dinoflagellate resting cysts were investigated at 11 locations in the south-eastern part of the North Sea. Twenty-six known cyst species and 7 unknown cyst types, which may act as seed population for planktonic dinoflagellate blooms, have been recorded for the first time in the area. The most common cysts in recent sediments were those ofScrippsiella trochoidea, Zygabikodinium lenticulatum, Peridinium dalei, Scrippsiella lachrymosa, Protoceratium reticulatum, Protoperidinium denticulatum, andP. conicum. At all stations,S. trochoidea dominated the cyst assemblages with a maximal abundance of 1303 living cysts/cm³ in the uppermost half centimetre. Cysts of the potentially toxic dinoflagellatesAlexandrium cf.excavatum andA. cf.tamarense were scarce. In the upper 2-cm layer of sediment, dinoflagellate cysts were found in concentrations of 1.8 up to 682 living cysts/cm³. Empty cysts constituted 22–56% of total cyst abundance. The comparative distribution of the cysts showed a general increase in abundance from inshore sites to the offshore area, whereby sandy stations exhibited the lowest cyst abundance and diversity. The wide distribution of living and empty cysts ofScrippsiella lachrymosa suggests that its motile form, which has not been officially recorded in the area until now, is a common plankton organism in German coastal waters. The relatively high abundance of cysts in recent sediments demonstrates the potential importance of benthic resting stages for the initiation of dinoflagellate blooms in the study area.     

WINTER PHYTOPLANKTON BLOOMS UNDER ICE ASSOCIATED WITH ELEVATED OXYGEN LEVELS1

Many shallow lakes in north temperate zones experience reduced dissolved oxygen concentration under ice. However, some shallow lakes display supersaturated dissolved oxygen concentrations (>20 mg·L ^{? 1}) in late winter under conditions of maximum ice thickness. During the winters of 1996, 1997, and 1999, we collected phytoplankton samples from Arrowwood Lake near Pingree, North Dakota to determine whether a specific alga was involved in dissolved oxygen supersaturation in this lake. Although dissolved oxygen supersaturation was not observed during this period, we did observe an increase in dissolved oxygen concentration that was associated with a phytoplankton bloom during late February and early March in both 1996 and 1997. In 1996, the bloom was composed of the dinoflagellate, Peridinium aciculiferum (Lemm.) Lemm. and several species of cryptomonads. A similar bloom of P. aciculiferum was followed by a bloom of several species of euglenoids in 1997. In contrast, P. aciculiferum was only a minor component of the winter phytoplankton, dissolved oxygen concentrations remained low, and no bloom event was observed in 1999. Statistical analyses indicated a significant relationship (r_s = 0.57, P = 0.019) between dissolved oxygen levels and the density of the dinoflagellate, P. aciculiferum, but no significant relationship between dissolved oxygen levels and densities of other phytoplankton. These results suggest that the elevated levels of dissolved oxygen are associated with the dinoflagellate, P. aciculiferum. This bloom was most likely the result of an excystment event rather than a general growth response.     

OBSERVATIONS ON THE TIDEPOOL ECOLOGY AND BEHAVIOR OF PERIDINIUM GREGARIUM1,2

Dense populations of a new species of dinoflagellate, Peridinium gregarium, have been found in a limited number of small rock tidepools on the coast of southern California. The organism is present throughout the year, but during periods of low temperatures and heavy tides it remains on the bottom of the pools in small aggregations. At other seasons P. gregarium leaves the aggregations during the morning hours and collects around the upper surface of small rocks in masses held together by a mucous matrix. Oxygen bubbles accumulated in the matrix eventually float the mass to the surface of the tidepool. During the afternoon or evening, the individuals return to the bottom aggregations leaving the matrix behind. Field and laboratory studies have attempted to define the nature of this organism's tidepool habitat and factors involved in the regulation of its diurnal movements.     

Phototactic responses of four marine dinoflagellates with different types of eyespot and chloroplast

Great structural variety is seen in the eyespot of dinoflagellates, a structure involved in phototaxis. Although there are several works on the phototactic responses in some species of dinoflagellates, none of the dinoflagellates used in these studies possessed an eyespot and, therefore, we have no knowledge of the relationship between eyespot type and phototactic response. In this study, we determined wavelength dependency curves for phototaxis in four marine dinoflagellates that possess a different type of either eyespot or chloroplast. These include: (i) a dinoflagellate possessing a peridinin-containing ohioroplast with an eyespot (Scrippsiella hexapraecingula Horiguchi et Chihara); (ii) a dinoflagellate containing a diatom endosymbiont and with the type B eyespot sensu Dodge (1984; (Peridinium foli-aceum (Stein) Biecheler); (iii) a dinoflagellate with peri-dinin-containing chloroplasts, but lacking an eyespot (Atexandrium hiranoi Kita et Fukuyo); and (iv) a dinoflagellate with fucoxanthin, 19′-hexanoyloxyfucoxanthin and 19′-butanoyloxyfucoxanthin, but lacking an eyespot (Gymnodinium mikimotoi Miyabe et Kominami ex Oda), Regardless of the eyespot or the chloroplast type, all four dinoflagellates showed similar wavelength dependency curves for phototaxis, with sensitivity between 380 and 520 nm, the highest peak at approximately 440 or 460 nm and smaller peaks or shoulders at 400–420 nm and 480–500 nm. Substantial peaks have also been noted in the ultraviolet range (260–280 nm). The ultrastructural study of the eye-spot of Scrippsiella hexapraecingula revealed that the eyespot consists of two layers of lipid globules and probably acts as a quarter-wave stack antenna.     

FERTILIZATION AND ZYGOTE DEVELOPMENT IN THE BINUCLEATE DINOFLAGELLATE PERIDINIUM BALTICUM (PYRRHOPHYTA)

Peridinium balticum (Pyrrhophyta) exists as a symbiosis between a nonphotosynthetic dinoflagellate host and a chlorophyll c-containing alga. It is hypothesized that P. balticum is an evolutionary link between primitive nonphotosynthetic and advanced photosynthetic dinoflagellates. This study documents pre- and postfertilization events of sexual reproduction in this unusual dinoflagellate for the first time. Light microscopy and TEM observations showed that gametes resemble vegetative cells except in the organization of their chloroplasts. Fusion of gametes occurred in a specific orientation, i.e., apical to sulcal area. The presence of an intact membrane between fusing pairs prior to plasmogamy was suggestive of enzymatic digestion of plates during fertilization. Rupture of this membrane triggers plasmogamy and karyogamy of the host, followed by that of the algal symbiont. A discussion of the cellular processes involved in gamete formation, fertilization, and zygote development is presented. The results of this investigation demonstrate that a synchronous sexual reproduction cycle has evolved for the symbiont and its dinoflagellate host.     

OBSERVATIONS ON THE PECULIAR DIURNAL MIGRATION OF A RED TIDE DINOPHYCEAE IN TROPICAL SHALLOW WATERS1

The dinoflagellate Peridinium cf. quinquecorne Abé forms red tide-like blooms in eutrophic shallow waters in the Philippines. The organism moves into a distinct near-surface layer when intensive solar radiation occurs, but only during the incoming tide. Shortly before high tide, regardless of light levels, the dinoflagellates seem to disappear. Simple experiments show that once intensive radiation has been reduced Peridinium quinquecorne moves out of the water column and attaches itself to solid objects away from the light. The morphology of the organism, especially as related to attachment, was studied through SEM. Its high swimming velocity and the reaction to radiation and tidal changes are described. The possibility that, superimposed on its reaction to light, this dinoflagellate may follow intrinsic tide-dependent oscillations is discussed.     

Durinskia baltica (Dinophyceae), a newly recorded species and genus from China, and its systematics

A freshwater dinoflagellate was identified as Durinskia baltica (Levander) Carty & Cox by morphological characteristics,with the plate formula:Po,x,4',2a,6',5c,4s,5',2'.Durinskia was a newly recorded dinoflagellate genus for China with two anterior intercalary plates and six characteristic precingular plates.Partial sequences of the small and large subunit ribosomal DNA and internal transcribed spacer sequences for the dinoflagellate cells were obtained from field samples.Molecular phylogenetic results indicated Durinskia species could cluster into a monophyletic group,which were distinct from Peridinium species.According to morphological and molecular evidence,it was agreed that the genus Durinskia was separated from the genus Peridinium,which could be a polyphyletic group.In addition,D.baltica was an infrequent diatom-harboring dinoflagellate which was known to possess an endosymbiotic diatom or diatom-like alga.The phylogenetic analyses indicated that D.baltica had a close affinity with Peridiniopsis penardii and P.niei,common freshwater bloom-forming species in China.     

A SURVEY OF AUXOTROPHY IN FIVE FRESHWATER DINOFLAGELLATES (PYRRHOPHYTA)1

Peridiniopsis polonicum (Wolosz.) Bourrelly requires vitamin B₁₂, and Peridinium limbatum (Stokes) Lemm. requires thiamin for growth. Unlike marine Peridinium species, Peridinium willei Huit.-Kaas, P. volzii Lemm., and P. inconspicuum Lemm. do not display auxotrophy. Peridinium volzii is strongly inhibited by concentrations of biotin above 1 μg L^?1.     

THE EFFECTS OF TEMPERATURE,IRRADIANCE, AND NITROGEN ON THE ENCYSTMENT AND GROWTH OF THE FRESHWATER DINOFLAGELLATES PERIDINIUM CINCTUM AND P. WILLEI IN CULTURE (DINOPHYCEAE)1

Effects of temperature, irradiance, and nitrogen availability on the encystment and growth of the freshwater dinoflagellates Peridinium cinctum Ehrenberg and Peridinium willei Huitfeld-Kaas were studied in culture. Lack of nitrogen was the main trigger of encystment in both species. Irradiance had a secondary effect on the percentage of the population of each species that encysted. Temperature did not significantly affect encystment in either species. In both species, only a small percentage of the population underwent encystment. Low light had an inhibitory effect on the growth of P. willei growing in nitrogen-sufficient medium.     

THE DINOFLAGELLATE TRANSVERSE FLAGELLUM: THREE-DIMENSIONAL RECONSTRUCTIONS FROM SERIAL SECTIONS1

Serial sections through in situ transverse flagella of the dinoflagellate Peridinium cinctum f. irregulatum (Lindem.) Lefévre are presented. Three-dimensional reconstructions based upon tangential and radial series show a helically coiled axoneme lying external to and distinct from an accessory strand. Hitherto undescribed vesicles within the expanded flagellar sheath are suggested to provide a decoupling effect between axoneme and strand. The flagellar axis bears two types of hair but anchoring threads between cingulum and flagellum have not been found. Functional and taxonomic implications of these observations are briefly discussed.