Glaucophyte

The glaucophytes, also known as glaucocystophytes or glaucocystids, are a small group of unicellular algae found in freshwater and moist terrestrial environments,<ref name="keeling">Template:Cite journal</ref><ref>Genomic Insights Into the Biology of Algae</ref> less common today than they were during the Proterozoic.<ref name="Cruzan 2018">Template:Cite book</ref> The stated number of species in the group varies from about 14 to 26.<ref name=FiguJackReye19/><ref>The monoplastidic bottleneck in algae and plant evolution | Journal of Cell Science</ref><ref>Template:AlgaeBase taxon</ref> Together with the red algae (Rhodophyta) and the green algae plus land plants (Viridiplantae or Chloroplastida), they form the Archaeplastida.

The glaucophytes are of interest to biologists studying the evolution of chloroplasts as they may be similar to the original algal type that led to the red algae and green plants, i.e. glaucophytes may be basal Archaeplastida.<ref name="keeling" /><ref>Template:Cite journal</ref><ref name=FiguJackReye19/>

Unlike red and green algae, glaucophytes only have asexual reproduction.<ref name="Walker 2012">Template:Cite book</ref>

Reproduction

Unlike red and green algae, glaucophytes reproduce exclusively through asexual means. They undergo open mitosis without centrioles, a trait shared with other basal eukaryotes. Reproductive modes include binary fission, zoospore formation, and autosporulation. For example, Cyanophora paradoxa divides longitudinally, producing two daughter cells, each inheriting a single cyanelle. Species of Glaucocystis reproduce via non-motile autospores. To date, there is no evidence of sexual reproduction in glaucophytes.<ref>Jackson, C. (2015). The Glaucophyta: The blue-green plants in a nutshell. Acta Societatis Botanicorum Poloniae 84(4): 439–443. https://doi.org/10.5586/asbp.2015.049</ref>

Characteristics

The plastids of glaucophytes are known as 'muroplasts',<ref name="Wise—Plastid diversity">Template:Cite book</ref> 'cyanoplasts', or 'cyanelles'. Unlike the plastids in other organisms, they have a peptidoglycan layer, believed to be a relic of the endosymbiotic origin of plastids from cyanobacteria.<ref name="keeling" /><ref name="Miyagishima">Template:Cite journal</ref>This peptidoglycan layer plays a functional role in plastid division and is considered molecular evidence of their cyanobacterial ancestry.<ref>Template:Cite journal</ref> Glaucophytes contain the photosynthetic pigment chlorophyll a.<ref name="keeling" /> Along with red algae<ref name="keeling" /> and cyanobacteria, they harvest light via phycobilisomes, structures consisting largely of phycobiliproteins. The green algae and land plants have lost that pigment.<ref name=Skuja>Skuja, A. (1948). Taxonomie des Phytoplanktons einiger Seen in Uppland, Schweden. Symbolae Botanicae Upsalienses 9(3): 1-399.Template:AlgaeBase taxon</ref> Like red algae, and in contrast to green algae and plants, glaucophytes store fixed carbon in the cytosol.<ref name="ball_2011">Template:Cite journal</ref>

This cytosolic carbon fixation, rather than fixation within plastids, is considered a retained ancestral trait. Glaucophyte phycobilisomes are composed primarily of phycocyanin and allophycocyanin, two key pigments also present in cyanobacteria. These pigments allow absorption of light at wavelengths that chlorophyll cannot, enhancing light harvesting in low-light aquatic environments.<ref>Template:Cite journal</ref>Studies of endosymbiotic gene transfer (EGT) suggest that several genes originally encoded in cyanobacterial ancestors have been relocated to the nuclear genome in glaucophytes, reflecting early stages of plastid-host genomic integration.<ref>Nowack, E. C. M., et al. (2008). Gene transfers from diverse bacteria compensate for reductive genome evolution in the chromatophore of Paulinella chromatophora. PNAS 105(5): 16782–16787. https://doi.org/10.1073/pnas.0809772105</ref>The evolution of glycogen and starch metabolism in eukaryotes gives molecular clues to understand the establishment of plastid endosymbiosis.Template:Cn

The most early-diverging genus is Cyanophora, which only has one or two plastids. When there are two, they are semi-connected.<ref name="de Vries Gould 2017 p. ">Template:Cite journal</ref>

Glaucophytes have mitochondria with flat cristae, and undergo open mitosis without centrioles. Motile forms have two unequal flagella, which may have fine hairs and are anchored by a multilayered system of microtubules, both of which are similar to forms found in some green algae.<ref name=Skuja/>

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Phylogeny

External

Together with red algae and Viridiplantae (green algae and land plants), glaucophytes form the Archaeplastida – a group of plastid-containing organisms that may share a unique common ancestor that established an endosymbiotic association with a cyanobacterium. The relationship among the three groups remains uncertain, although it is most likely that glaucophytes diverged first:<ref name=FiguJackReye19/>

The alternative, that glaucophytes and red algae form a clade, has been shown to be less plausible, but cannot be ruled out.<ref name=FiguJackReye19/>

Internal

The internal phylogeny of the glaucophytes and the number of genera and species varies considerably among taxonomic sources. A phylogeny of the Glaucophyta published in 2017 divided the group into three families, and includes five genera:<ref>Template:Cite book</ref> Template:Clade

Taxonomy

A 2019 list of the described glaucophyte species has the same three subdivisions, treated as orders, but includes a further five unplaced possible species, producing a total of between 14 and 19 possible species.<ref name=FiguJackReye19>Template:Citation</ref>

• Order Cyanophorales
  • Genus Cyanophora – 5–6 species

A recent study by Takahashi et al. (2023) used both morphological and molecular data to distinguish five distinct species within the genus Cyanophora, confirming species boundaries and supporting the utility of combined phylogenetic analysis.<ref>Takahashi, T., Hirano, Y., Ichinomiya, M., & Nozaki, H. (2023). Five *Cyanophora* (Cyanophorales, Glaucophyta) species delineated based on morphological and molecular data. Journal of Phycology 59(1): 29–42. https://doi.org/10.1111/jpy.13310</ref>

• Order Glaucocystales
  • Genus Glaucocystis – 7–8 species
• Order Gloeochaetales
  • Cyanoptyche – 1 species
  • Gloeochaete – 1 species
• Other possible species
  • ?Archaeopsis monococca Skuja
  • ?Chalarodora azurea Pascher
  • ?Glaucocystopsis africana Bourrelly
  • ?Peliaina cyanea Pascher
  • ?Strobilomonas cyaneus Schiller

Template:As of, AlgaeBase divided glaucophytes into only two groups, placing Cyanophora in Glaucocystales rather than Cyanophorales (however the entry was dated 2011).<ref name=AB_g44133>Template:AlgaeBase genus</ref> AlgaeBase included a total of 26 species in nine genera:<ref name=AB_t4301>Template:AlgaeBase taxon</ref>

• Glaucocystales
  • Chalarodora Pascher – 1 species
  • Corynoplastis Yokoyama, J.L.Scott, G.C.Zuccarello, M.Kajikawa, Y.Hara & J.A.West – 1 species
  • Cyanophora Korshikov – 6 species
  • Glaucocystis Itzigsohn – 13 species
  • Glaucocystopsis Bourrelly – 1 species
  • Peliaina Pascher – 1 species
  • Strobilomonas Schiller – 1 species
• Gloeochaetales
  • Cyanoptyche Pascher – 1 species
  • Gloeochaete Lagerheim – 1 species

None of the species of Glaucophyta is particularly common in nature.<ref name="keeling" />

The glaucophytes were considered before as part of family Oocystaceae, in the order Chlorococcales.<ref>Template:Cite web</ref>