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Abstract Background Volvocales in green algae have evolved by multicellularity of Chlamydomonas-like unicellular ancestor. Those with various cell numbers exist, such as unicellular Chlamydomonas, four-celled Tetrabaena, and Volvox species with different cell numbers (~1,000, ~5,000, and ~10,000). Each cell of these organisms shares two cilia and an eyespot, which are used for swimming and photosensing. They are all freshwater microalgae but inhabit different fluid environments: unicellular species live in low Reynolds-number (Re) environments where viscous forces dominate, whereas multicellular species live in relatively higher Re where inertial forces become non-negligible. Despite significant changes in the physical environment, during the evolution of multicellularity, they maintained photobehaviors (i.e., photoshock and phototactic responses), which allows them to survive under changing light conditions. Results In this study, we utilized high-speed imaging to observe flash-induced changes in the ciliary beating manner of 27 Volvocales strains. We classified flash-induced ciliary responses in Volvocales into four patterns: “1: temporal waveform conversion”, “2: no obvious response”, “3: pause in ciliary beating”, and “4: temporal changes in ciliary beating directions”. We found that which species exhibit which pattern depends on Re, which is associated with the individual size of each species rather than phylogenetic relationships. Conclusions These results suggest that only organisms that acquired different patterns of ciliary responses survived the evolutionary transition to multicellularity with a greater number of cells while maintaining photobehaviors. This study highlights the significance of the Re as a selection pressure in evolution and offers insights for designing propulsion systems in biomimetic micromachines.

Volvocales forms a species‐rich clade with wide morphological variety and is regarded as an ideal model for tracing the evolutionary transitions in multicellularity. The phylogenetic relationships among the colonial volvocine algae and its relatives are important for investigating the origin of multicellularity in the clade Reinhardtinia. Therefore, a robust phylogenetic framework of the unicellular and colonial volvocine algae with broad taxon and gene sampling is essential for illuminating the evolution of multicellularity. Recent chloroplast phylogenomic studies have uncovered five major orders in the Chlorophyceae, but the family‐level relationships within Sphaeropleales and Volvocales remain elusive due to the uncertain positions of some incertae sedis taxa. In this study, we contributed six newly sequenced chloroplast genomes in the Volvocales and analyzed a dataset with 91 chlorophycean taxa and 58 protein‐coding genes. Conflicting phylogenetic signals were detected among chloroplast genes that resulted in discordant tree topologies among different analyses. We compared the phylogenetic trees inferred from original nucleotide, RY‐coding, codon‐degenerate, and amino acid datasets, and improved the robustness of phylogenetic inference in the Chlorophyceae by reducing base compositional bias. Our analyses indicate that the unicellular Chlamydomonas and Vitreochlamys are close to or nested within the colonial taxa, and all the incertae sedis taxa are nested within the monophyletic Sphaeropleales s.l. We propose that the colonial taxa in the Reinhardtinia are paraphyletic and multicellularity evolved once in the volvocine green algae and might be lost in Chlamydomonas and Vitreochlamys. [ABSTRACT FROM AUTHOR]

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Graphical abstract Highlights • Thirty-two unicellular lineages related to colonial volvocaleans were recognized. • Phylogenetic resolution among the unicellular and colonial volvocaleans was improved. • Clade including Chlamydomonas pila is the probable sister of colonial volvocaleans. • Colony-associated characteristics were established within a short evolutionary time. Abstract In the green algal order Volvocales (Chlorophyceae), flagellate colonial forms have evolved at least four times. One of these colonial lineages, Tetrabaenaceae-Goniaceae-Volvocaceae (TGV), which belongs to the clade Reinhardtinia , is closely related to several unicellular chlamydomonads in the genera Chlamydomonas and Vitreochlamys. However, the unicellular sister of TGV has not been specified. Here, the largest ever 18S r RNA phylogenetic tree of Reinhardtinia was constructed including several newly isolated chlamydomonads, and a clade (core- Reinhardtinia) including 32 unicellular lineages and three colonial families were recognized. Interrelationships within core- Reinhardtinia were barely resolved in the tree, and therefore combined 18S- atp B- psa A- psa B- psb C- rbc L gene phylogenetic analyses were performed with selected representatives of 29 of the 32 unicellular lineages and three colonial families. The 29 unicellular lineages were clustered into five metaclades and an unassigned lineage; the metaclade that includes Chlamydomonas pila was resolved, with moderate support, as the sister clade to TGV. To examine possible biases from specific gene(s), long-branch taxa, and the heterogeneous base composition, phylogenetic analyses using several smaller data sets were also performed. Light microscopy of C. pila and its relatives indicated that any early steps towards colony evolution appeared after divergence of TGV from the C. pila lineage. [ABSTRACT FROM AUTHOR]

SUMMARY In 1960, Butcher isolated a strain of unicellular green alga, 'Chlamydomonas subehrenbergii' SAG 18.89, from a brackish water locality in England, facing the North Sea. Although C. subehrenbergii Butcher was previously described by the same author in 1959, the strain SAG 18.89 is in fact morphologically different, possessing shorter flagella, a different‐shaped papilla, and different‐sized vegetative cells. Light and fluorescence microscopy, molecular phylogenetic analyses, and ITS2 comparisons revealed that this strain represents an undescribed species of the genus Microglena (Volvocales, Chlorophyceae), described here as M. redcarensis sp. nov. The presence of contractile vacuoles, which are osmoregulatory organelles, suggests potential inhabitation in freshwater. Contractile vacuole activity and growth of M. redcarensis were therefore examined in media of different salinity. [ABSTRACT FROM AUTHOR]

Recent literary data on inversion (turning inside out) in the embryos of flagellated algae of the genus Volvox are critically analyzed. In this process, active changes in the shape of embryonic cells and the displacement of intercellular cytoplasmic bridges play an important role. After inversion, the flagella appear on the outer side of the young colony and provide its motility. Within the genus Volvox, two main modes of embryo inversion have been recently established during the asexual developmental cycle—inversion of type A and inversion of type B—represented by the two species most thoroughly studied, respectively, Volvox carterif. nagariensis and V. globator. However, the published opinion that the inversion of V. aureus embryos is of the type B seems to be doubtful. Comparative and evolutionary aspects of embryonic inversion in Volvox are discussed with the use of data on other genera of colonial volvocine algae. [ABSTRACT FROM AUTHOR]

Background: One of evolution's most important achievements is the development and radiation of multicellular organisms with different types of cells. Complex multicellularity has evolved several times in eukaryotes; yet, in most lineages, an investigation of its molecular background is considerably challenging since the transition occurred too far in the past and, in addition, these lineages evolved a large number of cell types. However, for volvocine green algae, such as Volvox carteri, multicellularity is a relatively recent innovation. Furthermore, V. carteri shows a complete division of labor between only two cell types - small, flagellated somatic cells and large, immotile reproductive cells. Thus, V. carteri provides a unique opportunity to study multicellularity and cellular differentiation at the molecular level. Results: This study provides a whole transcriptome RNA-Seq analysis of separated cell types of the multicellular green alga V. carteri f. nagariensis to reveal cell type-specific components and functions. To this end, 246 million quality filtered reads were mapped to the genome and valid expression data were obtained for 93% of the 14,247 gene loci. In the subsequent search for protein domains with assigned molecular function, we identified 9435 previously classified domains in 44% of all gene loci. Furthermore, in 43% of all gene loci we identified 15,254 domains that are involved in biological processes. All identified domains were investigated regarding cell type-specific expression. Moreover, we provide further insight into the expression pattern of previously described gene families (e.g., pherophorin, extracellular matrix metalloprotease, and VARL families). Our results demonstrate an extensive compartmentalization of the transcriptome between cell types: More than half of all genes show a clear difference in expression between somatic and reproductive cells. Conclusions: This study constitutes the first transcriptome-wide RNA-Seq analysis of separated cell types of V. carteri focusing on gene expression. The high degree of differential expression indicates a strong differentiation of cell types despite the fact that V. carteri diverged relatively recently from its unicellular relatives. Our expression dataset and the bioinformatic analyses provide the opportunity to further investigate and understand the mechanisms of cell type-specific expression and its transcriptional regulation. [ABSTRACT FROM AUTHOR]

Vampyrellid amoebae (Vampyrellida, Rhizaria) are widespread in freshwater, marine and terrestrial ecosystems and consume a wide range of eukaryotes, e.g. algae, fungi and micrometazoa. Environmental sequences indicate that only a small fraction of their genetic diversity is phenotypically characterised, emphasising the need to further explore unknown vampyrellids and their interactions with prey organisms. This study tests the prey range specificity of three vampyrellid amoebae with 49 strains of three common groups of freshwater algae (Zygnematophyceae, Euglenophyceae and Volvocales), and documents specific interactions by time-lapse microscopy. Two of the amoebae, here introduced as the novel genera Arachnomyxa and Planctomyxa based on morphology and SSU rRNA gene comparisons, display a complementary prey range and consume motile algae, namely Volvocales and Euglenophyceae, respectively. This reveals the existence of specialised 'plankton feeders' in the vampyrellid family Leptophryidae, contrasting with the strikingly broad prey range of Leptophrys vorax. The distinct autecological characteristics found in this group of morphologically rather indistinct amoebae contribute to our knowledge about the vastly understudied vampyrellid amoebae. Furthermore, time-lapse observations suggest that euglenoid movements exerted by the sluggish species of the 'Euglena deses group' as a reaction to vampyrellid contact may serve as an effective defence against microbial predators. [ABSTRACT FROM AUTHOR]

Pleodorina sphaerica Iyengar was considered to be a phylogenetic link between Volvox and the type species Pleodorina californica Shaw because it has small somatic cells distributed from the anterior to posterior poles in 64- or 128-celled vegetative colonies. However, cultural studies and molecular and ultrastructural data are lacking in P. sphaerica, and this species has not been recorded since 1951. Here, we performed light and electron microscopy and molecular phylogeny of P. sphaerica based on newly established culture strains originating from Thailand. Morphological features of the present Thai species agreed well with those of the previous studies of the Indian material of P. sphaerica and with those of the current concept of the advanced members of the Volvocaceae. The present P. sphaerica strains exhibited homothallic sexuality; male and facultative female colonies developed within a single clonal culture. Chloroplast multigene phylogeny demonstrated that P. sphaerica was sister to two other species of Pleodorina ( P. californica and Pleodorina japonica Nozaki) without posterior somatic cells, and these three species of Pleodorina formed a robust clade, which was positioned distally in the large monophyletic group including nine taxa of Volvox sect. Merrillosphaera and Volvox (sect. Janetosphaera) aureus Ehrenberg. Based on the present phylogenetic results, evolutionary losses of posterior somatic cells might have occurred in the ancestor of P. californica and P. japonica. Thus, P. sphaerica might represent an ancestral morphology of Pleodorina, rather than of Volvox. [ABSTRACT FROM AUTHOR]

The Earth is a cold place. Most of it exists at or below the freezing point of water. Although seemingly inhospitable, such extreme environments can harbour a variety of organisms, including psychrophiles, which can withstand intense cold and by definition cannot survive at more moderate temperatures. Eukaryotic algae often dominate and form the base of the food web in cold environments. Consequently, they are ideal systems for investigating the evolution, physiology, and biochemistry of photosynthesis under frigid conditions, which has implications for the origins of life, exobiology, and climate change. Here, we explore the evolution and diversification of photosynthetic eukaryotes in permanently cold climates. We highlight the known diversity of psychrophilic algae and the unique qualities that allow them to thrive in severe ecosystems where life exists at the edge. We focus on psychrophilic green algae from the Chlamydomonadales, discussing recent discoveries and directions for future research, and argue that they are among the best available models for studying psychrophily and life at the edge in photosynthetic eukaryotes. [ABSTRACT FROM AUTHOR]

New strains of a wall-less unicellular volvocalean flagellate were isolated from a freshwater environment in Japan. Observations of the alga, described here as Hapalochloris nozakii Nakada, gen. et sp. nov., were made using light, fluorescence, and electron microscopy. Each vegetative cell had two flagella, four contractile vacuoles, and a spirally furrowed cup-shaped chloroplast with an axial pyrenoid, and mitochondria located in the furrows. Based on the morphology, H. nozakii was distinguished from other known wall-less volvocalean flagellates. Under electron microscopy, fibrous material, instead of a cell wall and dense cortical microtubules, was observed outside and inside the cell membrane, respectively. Based on the phylogenetic analyses of 18S rRNA gene sequences, H. nozakii was found to be closely related to Asterococcus, Oogamochlamys, Rhysamphichloris, and ' Dunaliella' lateralis and was separated from other known wall-less flagellate volvocaleans, indicating independent secondary loss of the cell wall in H. nozakii. In the combined 18S rRNA and chloroplast gene tree, H. nozakii was sister to Lobochlamys. [ABSTRACT FROM AUTHOR]

Phytomonas spp. (phytomonads) are a diverse and globally distributed group of unicellular eukaryotes that parasitize a wide range of plants and are transmitted by insect hosts. Here we report the discovery and characterisation of a new species of Phytomonas , named Phytomonas oxycareni n. sp., which was obtained from the salivary glands of the invasive species of true bug Oxycarenus lavaterae (Heteroptera). The new Phytomonas species exhibits a long slender promastigote morphology and can be found both within the lumen of the insect host’s salivary glands as well as within the cells of the salivary gland itself. Sampling multiple individuals from the same population post-winter hibernation on two consecutive years revealed that infection was persistent over time. Finally, phylogenetic analyses of small subunit ribosomal RNA genes revealed that this species is sister to other species within the genus Phytomonas , providing new insight into the evolutionary history of the clade. [ABSTRACT FROM AUTHOR]

Background: Volvocine algae, which range from the unicellular Chlamydomonas to the multicellular Volvox with a germ-soma division of labor, are a model for the evolution of multicellularity. Within this group, the spheroidal colony might have evolved in two independent lineages: Volvocaceae and the goniacean Astrephomene. Astrephomene produces spheroidal colonies with posterior somatic cells. The feature that distinguishes Astrephomene from the volvocacean algae is lack of inversion during embryogenesis; the volvocacean embryo undergoes inversion after successive divisions to orient flagella toward the outside. The mechanisms of inversion at the molecular and cellular levels in volvocacean algae have been assessed in detail, particularly in Volvox carteri. However, embryogenesis in Astrephomene has not been subjected to such investigations. Results: This study relied on light microscopy time-lapse imaging using an actively growing culture of a newly established strain to conduct a developmental analysis of Astrephomene as well as to perform a comparison with the similar spheroidal volvocacean Eudorina. During the successive divisions involved in Astrephomene embryogenesis, gradual rotation of daughter protoplasts resulted in movement of their apical portions toward the embryonic posterior, forming a convex-to-spheroidal cell sheet with the apical ends of protoplasts on the outside. Differentiation of the posterior somatic cells from the embryo periphery was traced based on cell lineages during embryogenesis. In contrast, in Eudorina, the rotation of daughter protoplasts did not occur during successive cell divisions; however, inversion occurred after such divisions, and a spheroidal embryo was formed. Indirect immunofluorescence microscopy of basal bodies and nuclei verified this difference between Astrephomene and Eudorina in the movement of embryonic protoplasts. Conclusions: These results suggest different tactics for spheroidal colony formation between the two lineages: rotation of daughter protoplasts during successive cell divisions in Astrephomene, and inversion after cell divisions in Eudorina. This study will facilitate further research into the molecular and genetic mechanisms of the parallel evolution of the spheroidal colony in volvocine algae. [ABSTRACT FROM AUTHOR]

Background: The multicellular volvocine alga Pleodorina is intermediate in organismal complexity between its unicellular relative, Chlamydomonas, and its multicellular relative, Volvox, which shows complete division of labor between different cell types. The volvocine green microalgae form a group of genera closely related to the genus Volvox within the order Volvocales (Chlorophyta). Embryos of multicellular volvocine algae consist of a cellular monolayer that, depending on the species, is either bowl-shaped or comprises a sphere. During embryogenesis, multicellular volvocine embryos turn their cellular monolayer right-side out to expose their flagella. This process is called 'inversion' and serves as simple model for epithelial folding in metazoa. While the development of spherical Volvox embryos has been the subject of detailed studies, the inversion process of bowl-shaped embryos is less well understood. Therefore, it has been unclear how the inversion of a sphere might have evolved from less complicated processes. Results: In this study we characterized the inversion of initially bowl-shaped embryos of the 64- to 128-celled volvocine species Pleodorina californica. We focused on the movement patterns of the cell sheet, cell shape changes and changes in the localization of cytoplasmic bridges (CBs) connecting the cells. The development of living embryos was recorded using time-lapse light microscopy. Moreover, fixed and sectioned embryos throughout inversion and at successive stages of development were analyzed by light and transmission electron microscopy. We generated three-dimensional models of the identified cell shapes including the localization of CBs. Conclusions: In contrast to descriptions concerning volvocine embryos with lower cell numbers, the embryonic cells of P. californica undergo non-simultaneous and non-uniform cell shape changes. In P. californica, cell wedging in combination with a relocation of the CBs to the basal cell tips explains the curling of the cell sheet during inversion. In volvocine genera with lower organismal complexity, the cell shape changes and relocation of CBs are less pronounced in comparison to P. californica, while they are more pronounced in all members of the genus Volvox. This finding supports an increasing significance of the temporal and spatial regulation of cell shape changes and CB relocations with both increasing cell number and organismal complexity during evolution of differentiated multicellularity. [ABSTRACT FROM AUTHOR]

On the basis of prior phylogenetic analyses placing Gloeomonas as a lineage within the very diverse genus Chloromonas, it was hypothesized that the morphologically distinct Gloeomonas, bearing widely spaced basal bodies, evolved from ancestral organisms like Chloromonas. In addition, the phylogenetically related Chloromonas ( Cr.) rubrifilum, was expected to possess traits intermediate to the two forms. To test these hypotheses, we performed detailed transmission electron microscope analysis on nine species of these genera. The species were divided into two categories on the basis of ultrastructural features: group 1, consisting of four diverse species of Chloromonas and Ixipapillifera, had a V-shaped basal body arrangement, and group 2, of Cr. rubrifilum and four Gloeomonas species, forming a single clade possessing widely spaced basal bodies. Members of group 1 commonly had a simple, flat plate-like distal connecting fiber (dcf), three or sometimes two microtubules in the sinister root, continuous dexter and sinister striated microtubule-associated fibers (SMAFs), and a coarsely striated proximal fiber (CSPF) at the posterior-most regions of the basal bodies and probasal bodies, but lacked proximal and median proximal connecting fibers. Group 2 inherited these features, but had novel traits including the elongation and modification of the dcf, SMAFs, and CSPF that occurred with separation of basal bodies during growth, a dcf-associated layered structure, as well as protruding flagellar collars. The ultrastructural traits of Gloeomonas are interpreted as being evolutionarily modified from an ancestral Chloromonas morphology. Specific ultrastructural features were determined to be useful in characterizing these genera. Cr. rubrifilum of group 2 was not intermediate morphologically, but possessed the traits of Gloeomonas; thus G. rubrifilum comb. nov. was proposed. [ABSTRACT FROM AUTHOR]

The type of Haematococcus Flot. (Volvocales, Chlorophyceae) accumulates astaxanthin in its dormant stage. Currently, only one species is recognized in the genus; however, it is referred to by two names: H. lacustris (Gir.‐Chantr.) Rostaf. and H. pluvialis Flot. This confusion arose partly because of misunderstanding over the nomenclature and partly because of inadequate descriptions of Volvox lacustris Gir.‐Chantr. and H. pluvialis. Here, we designate an epitype based on a single cryopreserved culture that supports the newly designated lectotype of V. lacustris. Our treatment objectively synonymizes the two names and designates the correct name for the type of Haematococcus as H. lacustris, not H. pluvialis. In addition, we provide light microscopy descriptions and ITS rDNA information regarding the strain related to the epitype. [ABSTRACT FROM AUTHOR]

Chlamydomonas ( Cd.) is one of the largest but most polyphyletic genera of freshwater unicellular green algae. It consists of 400-600 morphological species and requires taxonomic revision. Toward reclassification, each morphologically defined classical subgenus (or subgroup) should be examined using culture strains. Chlamydomonas subg. Amphichloris is characterized by a central nucleus between two axial pyrenoids, however, the phylogenetic structure of this subgenus has yet to be examined using molecular data. Here, we examined 12 strains including six newly isolated strains, morphologically identified as Chlamydomonas subg. Amphichloris, using 18S rRNA gene phylogeny, light microscopy, and mitochondria fluorescent microscopy. Molecular phylogenetic analyses revealed three independent lineages of the subgenus, separated from the type species of Chlamydomonas, Cd. reinhardtii. These three lineages were further distinguished from each other by light and fluorescent microscopy-in particular by the morphology of the papillae, chloroplast surface, stigmata, and mitochondria-and are here assigned to three genera: Dangeardinia emend., Ixipapillifera gen. nov., and Rhysamphichloris gen. nov. Based on the molecular and morphological data, two to three species were recognized in each genus, including one new species, I. pauromitos. In addition, Cd. deasonii, which was previously assigned to subgroup ' Pleiochloris,' was included in the genus Ixipapillifera as I. deasonii comb. nov. [ABSTRACT FROM AUTHOR]

Background: The variability in gene organization and architecture of green algal mitochondrial genomes is only recently being studied on a finer taxonomic scale. Sequenced mt genomes from the chlorophycean orders Volvocales and Sphaeropleales exhibit considerable variation in size, content, and structure, even among closely related genera. However, sampling of mt genomes on a within-family scale is still poor and the sparsity of information precludes a thorough understanding of genome evolution in the green algae. Methods: Genomic DNA of representative taxa were sequenced on an Illumina HiSeq2500 to produce 2×100 bp paired reads, and mitochondrial genomes were assembled and annotated using Geneious v.6.1.5. Phylogenetic analysis of 13 protein-coding mitochondrial genes spanning the Sphaeropleales was performed. Results: This study presents one of the first within-family comparisons of mt genome diversity, and is the first to report complete mt genomes for the family Hydrodictyaceae (order Sphaeropleales). Four complete mt genomes representing three taxa and four phylogenetic groups, Stauridium tetras, Pseudopediastrum boryanum, and Pediastrum duplex, range in size from 37,723 to 53,560 bp. The size variability is primarily due to intergenic region expansion, and intron content is generally low compared with other mt genomes of Sphaeropleales. Conclusions: Certain gene rearrangements appear to follow a phylogenetic pattern, and with a more thorough taxon sampling genome-level sequence may be useful in resolving systematic conundrums that plague this morphologically diverse family. [ABSTRACT FROM AUTHOR]

The unicellular green alga Chlamydomonas reinhardtii has two actin genes, one encoding a conventional actin and the other an unconventional actin called novel actin-like protein (NAP). These actins apparently differ in their ability to polymerize, but their specific functions in the cell are unknown. To understand their evolutionary relationship, we investigated the molecular phylogeny of the actin/NAP family in Volvocales, using fully identified sequences ( Chlamydomonas moewusii, Gonium pectorale, and Volvox carteri) and newly determined partial sequences ( Eudorina elegans, and Volvulina steinii). Although the origin of the NAP clade remains ambiguous, the inferred phylogenetic trees strongly support the monophyly of NAP genes and show that only the genomes of volvocine species contain NAP genes. The nonsynonymous substitution rate of the NAP gene is, in consistence with its long branch length in the phylogeny, relatively high compared with that of the actin gene. NAP is thus apparently unique to volvocine species and most likely performs a cellular function with fewer constraints. [ABSTRACT FROM AUTHOR]

The role of melatonin (MLT) in micro- and macroalgae is the subject of continuous discussion. In eukaryotic unicellular organisms, the physiological role of MLT seems to differ from those of higher plants and mammals. Experimental data on the role of MLT in algae suggest that it acts mainly as an antioxidant. In the present study, the physiological role of MLT in phototrophic organisms was studied by measuring its effects on periodically controlled processes (cell division and HSP70B expression) as well as mechanisms involved in oxidative stress protection (carotenoids, FeSOD and MnSOD expression). The presence of MLT in Chlamydomonas reinhardtii was assessed and quantified using analytical thin-layer chromatography followed by fluorescence measurements. These data reveal that photoperiod had a significant influence on the processes and mechanisms studied in C. reinhardtii. Moreover, MLT had the greatest impact on C. reinhardtii under conditions promoting photo-oxidative stress. Addition of MLT to the growth medium down-regulated carotenoid content and the expression of FeSOD and MnSOD. These results support the idea that MLT has a physiological role as a direct and indirect antioxidant agent. [ABSTRACT FROM AUTHOR]

Volvox capensis was recorded previously only from South Africa. Here we established culture strains of this species from a sample collected in Montana, USA. Morphological details of asexual and sexual spheroids and molecular phylogeny of these strains were studied. The present alga was identified as V. capensis on the basis of morphological characteristics of asexual spheroids and zygotes. However, differences between the Montana and South African materials were recognized in number of sperm packets in a sexual, monoecious spheroid as well as in mode of gametic union between sperm and eggs. Possible polyspermy was observed in eggs of V. capensis by 4'-6-diamidino-2-phenylidole staining. Genetic difference between these two entities was small based on sequences of internal transcribed spacer 2 region of nuclear ribosomal DNA. [ABSTRACT FROM AUTHOR]

Variation is essential to ecological and evolutionary dynamics, but genetic variation of quantitative traits may be concentrated in a limited number of dimensions, constraining ecoevolutionary dynamics. We describe high-dimension variation in natural accessions of the model alga, Chlamydomonas reinhardtii, and test the hypothesis that extensive fitness variation across 30 environments is constrained to a small number of axes., We used high-throughput phenotyping to investigate morphological, fitness, and genotype × environment (G × E) variation in 18 natural C. reinhardtii accessions in 30 environments., The organismal phenotypes of cell cycle, cell size, and phototactic behavior exhibited substantial genetic variation between lines, and we found up to 74-fold fitness variation across accessions and environments. Approximately 47% of the extensive G × E variation is accounted for by the first two principal components (PCs) of the G-matrix corresponding to covariation in metals response, nitrogen availability, or salt and nutrient response., The natural variation of C. reinhardtii accessions supports the hypothesis that, despite abundant genetic variation across single environments, the species' adaptive response should be constrained along few major axes of selection. These results highlight the utility of natural accessions for integrating ecoevolutionary and genetic research. [ABSTRACT FROM AUTHOR]

Acetic acid at pH 5.0 can induce programmed cell death ( PCD) in Chlamydomonas reinhardtii cells, and abundant volatile organic compounds ( VOCs) were released during the process. In this study, the caspase-3-like activity was determined during the PCD, and it was increased significantly after 1 h. During the PCD, the dynamic release of VOCs from the cells was analyzed, and the emissions of total VOCs were raised markedly and reached the highest level at 2 h. Among the seven types of VOCs, such as alkanes, alkenes, terpenoids, alcohols, aldehydes, ketones and esters, three oxygenated compounds (aldehydes, ketones and esters) showed the most significant increase. O2-· and H2O2 were rapidly accumulated to high levels in the cells at the beginning of the PCD, but their content was reduced during the process. The activities of antioxidant enzymes were reduced gradually and even disappeared completely, demonstrating that the reduction of reactive oxygen species ( ROS) may not be scavenged by the antioxidant enzyme system. ROS have an intense oxidation and scavenging ability to volatile compounds, and the oxidation results in the production of oxygenated compounds. Therefore, the abundant production of oxygenated compounds indicated that ROS may play an important role in the dynamic release of VOCs from C. reinhardtii cells during PCD. [ABSTRACT FROM AUTHOR]

This article describes the morphological characteristics of the populations of green alga, Tetraspora gelatinosa, growing in the stressful Arctic conditions (77°00'22' N, 015°32'54.33' E). We present the first detailed morphological characteristics of this spe-cies from such a high latitude. Populations from both stagnant and flowing waters were studied. Depending on the type of habitat, their mucilaginous colonies (thalli) have differ-ent shapes, but the structure, size and the placement of the vegetative cells, akinetes and ameboid forms, as well as the pseudocilia morphology of both populations, were very simi-lar. Literature data on the distribution of T. gelatinosa indicate that it is a cosmopolitan spe-cies. Our data are compared with some characteristic features of this species growing in dif-ferent geographical and climatic zones. No significant differences were found in the morphology of the colonies compared, nor in the location and the inner structure of cells. How-ever, there were slight differences in cell size between the populations from warm and cold zones. [ABSTRACT FROM AUTHOR]

Although zygote morphology is important in traditional taxonomic systems of snow-inhabiting species of Chloromonas (Volvocales, Chlorophyceae), induction of sexual reproduction or hypnospore formation is difficult in culture. Thus, alternative taxonomic methodologies are required when sexual reproduction does not occur in cultures. Here, we performed a taxonomic study of snow-inhabiting Chloromonas species based on culture strains with elongate or ellipsoidal vegetative cells. Our comparative light and electron microscopy demonstrated that the strains were clearly distinguished into six species based on differences in vegetative cell shape and chloroplast morphology, the number of zoospores within the parental cell and the presence or absence of cell aggregates in old cultures. The six species are Chloromonas chenangoensis, C. fukushimae sp. nov., C. hohamii. C. pichinchae, C. tenuis sp. nov. and C. tughillensis. Although four of the six species formed a small clade based on phylogenetic analysis of multiple genes, their separation was supported by comparison of secondary structures of the internal transcribed spacer 2 of nuclear ribosomal DNA and genetic differences of nuclear- and chloroplast-encoded genes. The present study demonstrated that polyphasic analyses of culture strains of snow Chloromonas species could be used for the identification of natural species [ABSTRACT FROM AUTHOR]

Multicellularity arose several times in evolution of eukaryotes. The volvocine algae have full range of colonial organization from unicellular to colonies, and thus these algae are well-known models for examining the evolution and mechanisms of multicellularity. Gonium pectorale is a multicellular species of Volvocales and is thought to be one of the first small colonial organisms among the volvocine algae. In these algae, a cytoplasmic bridge is one of the key traits that arose during the evolution of multicellularity. Here, we observed the inversion process and the cytoplasmic bridges in G. pectorale using time-lapse, fluorescence, and electron microscopy. The cytoplasmic bridges were located in the middle region of the cell in 2-, 4-, 8-, and 16-celled stages and in inversion stages. However, there were no cytoplasmic bridges in the mature adult stage. Cytoplasmic bridges and cortical microtubules in G. pectorale suggest that a mechanism of kinesin-microtubule machinery similar to that in other volvocine algae is responsible for inversion in this species. [ABSTRACT FROM AUTHOR]

Chloromonas kasaiae sp. nov. (Volvocales, Chlorophyceae) was described based on a new culture strain originating from Japan. Based on light microscopy, vegetative cells of this new species were very similar to those of C. reticulata and C. serbinowii. However, C. kasaiae differed from these two species in the ultrastructure of eyespots (number of globule layers). Our phylogenetic analyses, based on a combined data set from nuclear-encoded 18S ribosomal (r) RNA, the ATP synthase beta subunit and P700 chlorophyll α apoprotein A2 genes, robustly resolved that C. kasaiae was separate from C. reticulata. Although these analyses did not resolve the phylogenetic relationship between C. kasaiae and C. serbinowii, separation of the two was supported based on a comparison of the secondary structure of nuclear rDNA internal transcribed spacer 2. [ABSTRACT FROM AUTHOR]

Background: Isogamous organisms lack obvious cytological differences in the gametes of the two complementary mating types. Consequently, it is difficult to ascertain which of the two mating types are homologous when comparing related but sexual isolated strains or species. The colonial volvocalean algal genus Gonium consists of such isogamous organisms with heterothallic mating types designated arbitrarily as plus or minus in addition to homothallic strains. Homologous molecular markers among lineages may provide an “objective” framework to assign heterothallic mating types. Methodology/Principal Findings: Using degenerate primers designed based on previously reported MID orthologs, the “master regulator” of mating types/sexes in the colonial Volvocales, MID homologs were identified and their presence/absence was examined in nine strains of four species of Gonium. Only one of the two complementary mating types in each of the four heterothallic species has a MID homolog. In addition to heterothallic strains, a homothallic strain of G. multicoccum has MID. Molecular evolutionary analysis suggests that MID of this homothallic strain retains functional constraint comparable to that of the heterothallic strains. Conclusion/Significance: We coordinated mating genotypes based on presence or absence of a MID homolog, respectively, in heterothallic species. This scheme should be applicable to heterothallic species of other isogamous colonial Volvocales including Pandorina and Yamagishiella. Homothallism emerged polyphyletically in the colonial Volvocales, although its mechanism remains unknown. Our identification of a MID homolog for a homothallic strain of G. multicoccum suggests a MID-dependent mechanism is involved in the sexual developmental program of this homothallic species. [ABSTRACT FROM AUTHOR]

The transition from unicellular, to colonial, to larger multicellular organisms has benefits, costs, and requirements. Here we present a model inspired by the volvocine green algae that explains the dynamics involved in the unicellular-multicellular transition using life-history theory and allometry. We model the two fitness components (fecundity and viability) and compare the fitness of hypothetical colonies of different sizes with varying degrees of cellular differentiation to understand the general principles that underlie the evolution of multicellularity. We argue that germ-soma separation may have evolved to counteract the increasing costs and requirements of larger multicellular colonies. The model shows that the cost of investing in soma decreases with size. For lineages such as the Volvocales, as reproduction costs increase with size for undifferentiated colonies, soma specialization benefits the colony indirectly by decreasing such costs and directly by helping reproductive cells acquire resources for their metabolic needs. Germ specialization is favored once soma evolves and takes care of vegetative functions. To illustrate the model, we use some allometric relationships measured in Volvocales. Our analysis shows that the cost of reproducing an increasingly larger group has likely played an important role in the transition to multicellularity and cellular differentiation. The transition from unicellular, to colonial, to larger multicellular organisms has benefits, costs, and requirements. Here we present a model inspired by the volvocine green algae that explains the dynamics involved in the unicellular-multicellular transition using life-history theory and allometry. We model the two fitness components (fecundity and viability) and compare the fitness of hypothetical colonies of different sizes with varying degrees of cellular differentiation to understand the general principles that underlie the evolution of multicellularity. We argue that germ-soma separation may have evolved to counteract the increasing costs and requirements of larger multicellular colonies. The model shows that the cost of investing in soma decreases with size. For lineages such as the Volvocales, as reproduction costs increase with size for undifferentiated colonies, soma specialization benefits the colony indirectly by decreasing such costs and directly by helping reproductive cells acquire resources for their metabolic needs. Germ specialization is favored once soma evolves and takes care of vegetative functions. To illustrate the model, we use some allometric relationships measured in Volvocales. Our analysis shows that the cost of reproducing an increasingly larger group has likely played an important role in the transition to multicellularity and cellular differentiation. [ABSTRACT FROM AUTHOR]