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PubMed2003

Systematics of primary osmotrophic euglenids: a molecular approach to the phylogeny of Distigma and Astasia (Euglenozoa).

Nuclear-encoded SSU rRNA genes from nine strains of Distigma and three strains of Astasia were sequenced and analysed phylogenetically with maximum-likelihood and maximum-parsimony methods. It could be demonstrated that the genus Distigma is paraphyletic, consisting of two distinct clades: one comprises four strains of the type species, Distigma proteus, and the other includes four strains of Distigma curvatum, Distigma gracile, Distigma sennii and Distigma elegans. These findings are well corroborated by morphological characteristics. The investigated species of Astasia are closely related to members of the Rhabdomonadida, thus rendering the genus Astasia polyphyletic, with Astasia longa branching within the phototrophs. All of the species investigated cluster in a well-supported group of primary osmotrophic euglenids that are not derived from photosynthetic ancestors. The recovered clades are characterized by their sequence diversity. After different evolutionary rates among lineages had been determined, a modified slow-fast approach was used to differentiate phylogenetic signal from noise. Finally, a revised systematic scheme based on phylogenetic relationships is suggested to render euglenid taxonomy more transparent: primary osmotrophic euglenids are classified as Aphagea, and members of the D. curvatum group are transferred into the new subgenus Parvonema.

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PubMedDate not supplied

A model for the morphogenesis of strip reduction patterns in phototrophic euglenids: evidence for heterochrony in pellicle evolution.

We propose a general developmental model that explains the evolutionary origin, diversification, and inheritance of pellicle strip patterns in phototrophic euglenids. Dividing cells of Euglena gracilis, E. viridis, and Phacus similis were observed with scanning electron microscopy in order to study the morphogenesis of posterior whorls of strip reduction. We found evidence that constant whorl numbers are maintained through cell division because of organized strip growth before and during cytokinesis. Alternating nascent strips form a new whorl of strip reduction at each of the anterior and posterior ends of daughter cells. Strips that terminated to form posterior whorls in the mother cell change in length during the development of daughter cells. In the mother cells of E. gracilis, the strips forming whorls I and II grow to become whorls II and III, respectively, in the daughter cells; the strips forming whorl III in the mother cell lengthen and meet with other strips already present at the posterior tip of daughter cells. This process of whorl morphogenesis during asexual reproduction is consistent with known variation in pellicle strip patterns and suggests that heterochrony played a major role in the ultrastructural evolution of phototrophic euglenids.

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PubMedDate not supplied

A molecular study of euglenoid phylogeny using small subunit rDNA.

The euglenoids are an ancient and extremely diverse lineage of eukaryotic flagellates with unclear relationships among taxa. Synapomorphies for the euglenoids include a surface pellicle and a closed mitosis with a series of separate sub-spindles. The taxonomy currently in use is inconsistent with the available data and needs revision. Most euglenoid phylogenies are largely intuitive reconstructions based on a limited number of morphological characters. Therefore, we have added molecular characters from the Small Subunit (SSU) rDNA to generate an overall phylogenetic framework for the euglenoids. SSU rDNA sequences from photosynthetic, osmotrophic, and phagotrophic euglenoids were aligned based on secondary structure. Phylogenetic analysis using the conserved areas of the sequence was performed using parsimony, maximum likelihood, and distance methods. Trees derived using different criteria are in agreement. The euglenoids form a distinct monophyletic clade with phagotrophic members diverging prior to the phototrophic and osmotrophic members. Among photosynthetic members, the biflagellate form diverged prior to the uniflagellate form. Additionally, the genus Euglena appears to be paraphyletic, with osmotrophic taxa, such as Astasia and Khawkinea, diverging independently within the clade containing the photosynthetic genus Euglena.

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PubMedDate not supplied

Comparative morphology of the euglenid pellicle. I. Patterns of strips and pores.

In anticipation that improved knowledge of euglenid morphology will provide robust apomorphy-based definitions for clades, transmission and scanning electron microscopy were used to reveal novel morphological patterns associated with the euglenid pellicle. In some taxa, the number of pellicle strips around the cell periphery reduces as discrete whorls at the anterior and posterior ends of the cell. The number of whorls at either end varies between selected euglenid taxa but is invariant within a taxon. The pattern of strip reduction associated with these whorls is shown to have at least three evolutionarily linked states: exponential, pseudoexponential, and linear. Two general equations describe these states near the posterior end of euglenid cells. Exponential patterns of strip reduction near the anterior end are described by a third equation. In addition, several euglenid taxa were found to possess conspicuous pellicle pores. These pores are arranged in discrete rows that follow the articulation zones between adjacent strips. The number of strips between rows of pores varies between taxa and displays a series of consecutive character states that differ by a power of two. The patterns of pores may not only have phylogenetical and taxonomical value but may provide morphological markers for following strip maturation during cytoskeletal reproduction.

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PubMedDate not supplied

Comparative morphology of the euglenid pellicle. II. Diversity of strip substructure.

The morphological diversity associated with the strip substructure of the euglenid pellicle was examined, and after identifying characters and states, we outlined hypotheses about their evolution. We have attempted to standardize terms necessary for analytical comparisons of strips by providing a glossary and comparing published synonyms. Most of the substructural diversity found in euglenids is demonstrated with 13 representative taxa. Strips are generally composed of two subcomponents: frames and projections. Frames support the basic shape of strips and many can be described as either S-shaped, plateau-shaped, M-shaped, or A-shaped. Projections branch laterally from the frames, are usually periodic, and can be described as thread-like structures, an indented plate, tooth-like structures, and plate-like structures. The ancestral state included strips that were few in number, flat, and fused. The strips became S-shaped and disjoined in the lineage leading to most euglenid taxa. These strips became secondarily flattened and fused in one lineage. In some lineages of phototrophs, the strips became increasingly robust. Two strips of different morphology formed the repeating pellicular unit or doublet in four taxa. These doublets evolved convergently at least three times and may provide insights into developmental patterns of the cytoskeleton.

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PubMedDate not supplied

Food vacuole contents in the ciliate, Balantidium jocularum (Balantididae), a symbiont in the intestine of the surgeonfish, Naso tonganus (Acanthuridae).

During the past 16 years, the ciliate Balantidium jocularum has been collected from the intestines of many specimens of its fish host, Naso tonganus, all collected from the Great Barrier Reef near Lizard Island, Australia. Ciliates for this study of food consumption were isolated in 1988, 1989, 2003, and 2005. Nineteen specimens of B. jocularum were examined in the transmission electron microscope to determine the contents of both food vacuoles and a putative discharging cytoproct vacuole. Food vacuoles contained rod-shaped bacteria, tightly coiled spirilliform bacteria, and one or more euglenid flagellates. In several balantidia of somewhat different form than the type species of B. jocularum, the large bacterium, Epulopiscium fishelsoni, was observed in light microscope protargol preparations. Some putative phagolysosomes retained spirilliform bacteria that were apparently intact, and others contained partially digested flagellates. Food in a single discharging cytoproct vacuole consisted of normal appearing spirilliform bacteria, some other bacteria, and no flagellates. The results argue for non-selective ingestion of food and selective digestion; hence, somewhat inefficient food processing.

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PubMedDate not supplied

Dinoflagellate, Euglenid, or Cercomonad? The ultrastructure and molecular phylogenetic position of Protaspis grandis n. sp.

Protaspis is an enigmatic genus of marine phagotrophic biflagellates that have been tentatively classified with several different groups of eukaryotes, including dinoflagellates, euglenids, and cercomonads. This uncertainty led us to investigate the phylogenetic position of Protaspis grandis n. sp. with ultrastructural and small subunit (SSU) rDNA sequence data. Our results demonstrated that the cells were dorsoventrally flattened, shaped like elongated ovals with parallel lateral sides, 32.5-55.0 mum long and 20.0-35.0 mum wide. Moreover, two heterodynamic flagella emerged through funnels that were positioned subapically, each within a depression and separated by a distinctive protrusion. A complex multilayered wall surrounded the cell. Like dinoflagellates and euglenids, the nucleus contained permanently condensed chromosomes and a large nucleolus throughout the cell cycle. Pseudopodia containing numerous mitochondria with tubular cristae emerged from a ventral furrow through a longitudinal slit that was positioned posterior to the protrusion and flagellar apparatus. Batteries of extrusomes were present within the cytoplasm and had ejection sites through pores in the cell wall. The SSU rDNA phylogeny demonstrated a very close relationship between the benthic P. grandis n. sp. and the planktonic Cryothecomonas longipes. These ultrastructural and molecular phylogenetic data for Protaspis indicated that the current taxonomy of Protaspis and Crythecomonas is in need of re-evaluation. The composition and identity of Protaspis is reviewed and suggestions for future taxonomic changes are presented. Problems within the genus Cryothecomonas are highlighted as well, and the missing data needed to resolve ambiguities between the two genera are clarified.

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PubMed2005

Photoreceptor for curling behavior in Peranema trichophorum and evolution of eukaryotic rhodopsins.

When it is gliding, the unicellular euglenoid Peranema trichophorum uses activation of the photoreceptor rhodopsin to control the probability of its curling behavior. From the curled state, the cell takes off in a new direction. In a similar manner, archaea such as Halobacterium use light activation of bacterio- and sensory rhodopsins to control the probability of reversal of the rotation direction of flagella. Each reversal causes the cell to change its direction. In neither case does the cell track light, as known for the rhodopsin-dependent eukaryotic phototaxis of fungi, green algae, cryptomonads, dinoflagellates, and animal larvae. Rhodopsin was identified in Peranema by its native action spectrum (peak at 2.43 eV or 510 nm) and by the shifted spectrum (peak at 3.73 eV or 332 nm) upon replacement of the native chromophore with the retinal analog n-hexenal. The in vivo physiological activity of n-hexenal incorporated to become a chromophore also demonstrates that charge redistribution of a short asymmetric chromophore is sufficient for receptor activation and that the following isomerization step is probably not required when the rest of the native chromophore is missing. This property seems universal among the Euglenozoa, Plant, and Fungus kingdom rhodopsins. The rhodopsins of animals have yet to be studied in this respect. The photoresponse appears to be mediated by Ca2+ influx.

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PubMed2005

Unique mitochondrial genome structure in diplonemids, the sister group of kinetoplastids.

Kinetoplastid flagellates are characterized by uniquely massed mitochondrial DNAs (mtDNAs), the kinetoplasts. Kinetoplastids of the trypanosomatid group possess two types of mtDNA molecules: maxicircles bearing protein and mitoribosomal genes and minicircles specifying guide RNAs, which mediate uridine insertion/deletion RNA editing. These circles are interlocked with one another to form dense networks. Whether these peculiar mtDNA features are restricted to kinetoplastids or prevail throughout Euglenozoa (euglenids, diplonemids, and kinetoplastids) is unknown. Here, we describe the mitochondrial genome and the mitochondrial ultrastructure of Diplonema papillatum, a member of the diplonemid flagellates, the sister group of kinetoplastids. Fluorescence and electron microscopy show a single mitochondrion per cell with an ultrastructure atypical for Euglenozoa. In addition, DNA is evenly distributed throughout the organelle rather than compacted. Molecular and electron microscopy studies distinguish numerous 6- and 7-kbp-sized mitochondrial chromosomes of monomeric circular topology and relaxed conformation in vivo. Remarkably, the cox1 gene (and probably other mitochondrial genes) is fragmented, with separate gene pieces encoded on different chromosomes. Generation of the contiguous cox1 mRNA requires trans-splicing, the precise mechanism of which remains to be determined. Taken together, the mitochondrial gene/genome structure of Diplonema is not only different from that of kinetoplastids but unique among eukaryotes as a whole.

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PubMedDate not supplied

The Glu-modification of alpha-tubulin in the feeding apparatus of the primitive flagellate Entosiphon sulcatum is only apparent after detergent treatment.

Using specific monoclonal antibodies, we investigated the distribution of post-translational modified Tyr- and Glu-tubulins during interphase of the primitive flagellate Entosiphon sulcatum. Immunofluorescence studies of simultaneously permeabilized and fixed cells revealed that microtubular structures comprising Ca(2+)-labile subpellicular and flagellar MTs and Ca(2+)-stable MTs in the siphon complex (feeding organelle) reacted surprisingly unorthodox with antibodies against Tyr- and Glu-tubulin: Unexpectedly, the siphon complex consisting of Ca(2+)-stable MTs appeared exclusively Tyr-positive, whereas the Ca(2+)-labile subpellicular and flagellar MTs reacted with the Glu- as well as with the Tyr-antibody. That the siphon MTs were indeed Ca(2+)-stable and all other MTs had become solubilized, was verified by EM-observation. This surprising result contrasting considerably with the permanent nature of the siphon complex, was reconsidered after preceding lysis and extraction procedures. Depending on the type of detergent used and on extraction times applied, the MTs of the siphon complex now always showed also Glu-positivity, indicating the presence of detyrosinated alpha-tubulin as a biochemical marker of stabilized MTs. Since saponin, irrespective of subsequent extraction times, always produced a Glu-positive reaction and ultrastructural analysis never gave compelling evidence for a drastic MAP-removal, we conclude that the Glu-epitope became freely accessible due to conformational changes in the tubulin polymeres.

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PubMed2005

Algal community used for assessment of water quality of Haranbaree dam and Mosam river of Maharashtra.

An algal community was used to assess the quality of water of Haranbaree dam and Mosam river of Maharashtra. Pollution tolerant genera and species of 4 groups of algae from each of 3 stations of Haranbaree dam and Mosam river were recorded. By using Palmer's index of pollution for rating of water samples the total score of each station of study area was greater than 20 indicating the confirmed high organic pollution. 34 pollution tolerant genera were recorded at all stations of dam and river. Out of 34 pollution tolerant genera, 27 genera and 33 genera were observed at the dam and river sites respectively. In present study 22 pollution tolerant species are observed at dam and river sites.

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PubMedDate not supplied

[Organization of ribosomal DNA from the phytoflagellates Astasia longa and Euglena gracilis: comparison of the structure of 19S and 28S rRNA genes].

Restriction maps of Astasia longa and Euglena gracilis var. bacillaris were built and localization of 5.8S, 19S and 28S rRNA genes was established on them by blot-hybridization method. In the A. longa and E. gracilis plasmid rDNA three intergeneous regions were found, two of which were intergeneous transcribed spacers flanking 5.8S RNA gene, and the third region seems to be an untranscribed spacer. Localization of 9 primers from E. gracilis was established on A. longa 19S rRNA gene by PCR; it was similar to E. gracilis. Using amplified A. longa 19S rRNA gene (2300 bp) we have determined the sequence of its 3'-region, which showed 87% homology with the same region of E. gracilis. Using plasmid pA1 S3-H1 containing S3-H1 fragment of A. longa rDNA the sequence of 28S rRNA gene 3'-region was determined. This sequence includes regions homologous to corresponding regions of E. gracilis Z: 6 nucleotides of 12/13 internal spacer, complete 13/14 internal spacer (86 bp), genes for 13th and 14th 28S rRNA components (56 and 86 bp) with approximately 90% of homology with E. gracilis Z, and fragment of untranscribed spacer (136 bp) with approximately 70% homology. It was shown that 28S rRNA genes of A. longa and E. gracilis have similar structure. Our data allow to conclude that these phytoflagellats are closely related.

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