Chromista

Chromista is a biological kingdom consisting of some single-celled and multicellular eukaryotic organisms, which share similar features in their photosynthetic organelles (plastids).[1] It includes all protists such as some algae, diatoms, oomycetes, and protozoans whose plastids contain chlorophyll c. It is probably a polyphyletic group whose members independently arose as separate evolutionary group from the common ancestor of all eukaryotes.[2] As it is assumed the last common ancestor already possessed chloroplasts of red algal origin, the non-photosynthetic forms evolved from ancestors able to perform photosynthesis. Their plastids are surrounded by four membranes, and are believed to have been acquired from some red algae.

Chromista
Scientific classification
Domain:
Eukaryota
(unranked):
Kingdom:
Chromista

Phyla

Chromista as a biological kingdom was created by British biologist Thomas Cavalier-Smith in 1981 to differentiate some protists from typical protozoans and plants.[3] According to Cavalier-Smith, the kingdom originally included only algae, but his later analysis indicated that many protozoans also belong to the new group. As of 2018, the kingdom is as diverse as kingdoms Plantae and Animalia, consisting of eight phyla. Notable members include marine algae, potato blight, dinoflagellates, Paramecium, brain parasite (Toxoplasma) and malarial parasite (Plasmodium).[4]

Biology

Structure of typical chromists compared with plant cell (left)

Members of Chromista are single-celled and multicellular eukaryotes having basically either or both features:[3]

  1. plastid(s) that contain chlorophyll c and lies within an extra (periplastid) membrane in the lumen of the rough endoplasmic reticulum (typically within the perinuclear cisterna);
  2. cilia with tripartite or bipartite rigid tubular hairs.

Even though the kingdom includes diverse organisms from algae to malarial parasites (Plasmodium), they are genetically related and are believed to have evolved from a common ancestor with all other eukaryotes but in an independent line of evolution. As a result of evolution, many have retained their plastids and cilia, while some have lost them.[5] Molecular evidences indicate that the plastids in chromists were derived from red algae through secondary symbiogenesis in a single event.[6] (In contrasts, plants acquired their plastids from cyanobacteria through primary symbiogenesis.[7]) These plastids are now enclosed in two extra cell membranes, making a four membrane envelope, as a result of which they acquired many other membrane proteins for transporting molecules in and out of the organelles. The diversity of chromists arose from degeneration, loss or replacement of the plastids in some lineages.[8] There was also additional symbiogenesis of green algae, the genes of which are retained in some members (such as heterokonts),[9] as well as bacterial chlorophyll (indicated by the presence of ribosomal protein L36 gene, rpl36) in haptophytes and cryptophytes.[10]

History and groups

The name Chromista was first introduced by Cavalier-Smith in 1981;[3] the earlier names Chromophyta, Chromobiota and Chromobionta correspond to roughly the same group.

It has been described as consisting of three different groups:[11]

Cavalier-Smith later (in 2009) stated his reason for making a new kingdom, saying:

I established Chromista as a kingdom distinct from Plantae and Protozoa because of the evidence that chromist chloroplasts were acquired secondarily by enslavement of a red alga, itself a member of kingdom Plantae, and their unique membrane topology.[5]

Since then Chromista has been defined in different ways at different times. In 2010, Cavalier-Smith himself indicated his desire to move Alveolata, Rhizaria and Heliozoa into Chromista.

Some examples of classification of the Chromista and related groups are shown below.[12][13]

Chromophycées (Chadefaud, 1950)

The Chromophycées (Chadefaud, 1950),[14] renamed Chromophycota (Chadefaud, 1960),[15] included the current Ochrophyta (autotrophic Stramenopiles), Haptophyta (included in Chrysophyceae until Christensen, 1962), Cryptophyta, Dinophyta, Euglenophyceae and Choanoflagellida (included in Chrysophyceae until Hibberd, 1975).

Chromophyta (Christensen 1962, 1989)

The Chromophyta (Christensen 1962, 1989), defined as algae with chlorophyll c, included the current Ochrophyta (autotrophic Stramenopiles), Haptophyta, Cryptophyta, Dinophyta and Choanoflagellida. The Euglenophyceae were transferred to the Chlorophyta.

Chromophyta (Bourrelly, 1968)

The Chromophyta (Bourrelly, 1968) included the current Ochrophyta (autotrophic Stramenopiles), Haptophyta and Choanoflagellida. The Cryptophyceae and the Dinophyceae were part of Pyrrhophyta (= Dinophyta).

Chromista (Cavalier-Smith, 1981)

The Chromista (Cavalier-Smith, 1981) included the current Stramenopiles, Haptophyta and Cryptophyta.

Chromalveolata (Adl et al., 2005)

The Chromalveolata (Cavalier-Smith, 1981) included Stramenopiles, Haptophyta, Cryptophyta and Alveolata.

Classification

Chromista classification

In his original classification in 1981, Cavalier-Smith included three phyla under Chromista, such as:[3]

  1. Heterokonta
  2. Haptophyta
  3. Cryptomonada

In 2010, Cavalier-Smith reorganised Chromista to include SAR supergroup such as (Stramenopiles, Alveolata and Rhizaria) and Hacrobia (Haptista, Cryptista).[5]

In 2017, he and his colleagues made a new higher-level grouping of all organisms as a revision of seven kingdoms model. In it, they classified Chromista into 2 subkingdoms and 10 phyla, namely:[16]

  1. Subkingdom Hacrobia
    1. Phulym N.N. (=nomen nominandum, name unknown)
    2. Phylum Cryptista
    3. Phylum Haptophyta
    4. Phylum Heliozoa
  2. Subkingdom Harosa (=Supergroup SAR)
    1. Phylum Ciliophora
    2. Phylum Miozoa
    3. Phylum Bigyra
    4. Phylum Pseudofungi (=Oomycota)
    5. Phylum Cercozoa
    6. Phylum Retaria

Cavalier-Smith made a new analysis of Chromista in 2018 in which he classified all chromists into 8 phyla, as follows:[4]

  1. Phylum Miozoa
  2. Phylum Ciliophora
  3. Phylum Gyrista
  4. Phylum Bigyra
  5. Phylum Cercozoa
  6. Phylum Retaria
  7. Phylum Cryptista
  8. Phylum Haptista

Phylogeny

Chromista
Hacrobia
Cryptista

Corbihelia Cavalier-Smith 2015

Cryptophyta Silva 1962

Haptista

Centroheliozoa Cushman & Jarvis 1929 sensu Durrschmidt & Patterson 1987

Haptophyta Hibberd 1976 emend. Edvardsen & Eikrem 2000

SAR supergroup
Rhizaria

Phytomyxea Engler & Prantl 1897 em. Cavalier-Smith 1993

Vampyrellidea Cavalier-Smith 2017

Filosa Leidy 1879 emend. Cavalier-Smith 2003 stat. nov.

Retaria Cavalier-Smith 1999 emend. Cavalier-Smith 2017

Halvaria
Alveolata

Ciliophora Doflein 1901 stat. n. Copeland 1956

Miozoa Cavalier-Smith 1987

Heterokonta

Platysulcea Cavalier-Smith 2017

Sagenista Cavalier-Smith 1995 stat. n. 2006

Bikosea Cavalier-Smith 2013

Placidozoa Cavalier-Smith 2013

Gyrista

Bigyromonadea Cavalier-Smith 1998

Peronosporomycota Dick 1995

Hyphochytriomycota Whittaker 1969

Pirsonea Cavalier-Smith 2017

Ochrophyta Cavalier-Smith 1986

Controversy

Molecular trees have had some difficulty resolving relationships between the different groups. All three may share a common ancestor with the alveolates (see chromalveolates), but there is evidence that suggests that the haptophytes and cryptomonads do not belong together with the heterokonts or the SAR clade, but may be associated with the Archaeplastida.[1][17] Cryptista specifically may be sister or part of Archaeplastida.[18]

See also

References

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  2. Cavalier-Smith T, Allsopp MT, Chao EE (November 1994). "Chimeric conundra: are nucleomorphs and chromists monophyletic or polyphyletic?". Proceedings of the National Academy of Sciences of the United States of America. 91 (24): 11368–72. Bibcode:1994PNAS...9111368C. doi:10.1073/pnas.91.24.11368. PMC 45232. PMID 7972066.
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  7. Ponce-Toledo, Rafael I.; Deschamps, Philippe; López-García, Purificación; Zivanovic, Yvan; Benzerara, Karim; Moreira, David (2017). "An Early-Branching Freshwater Cyanobacterium at the Origin of Plastids". Current Biology. 27 (3): 386–391. doi:10.1016/j.cub.2016.11.056. PMC 5650054. PMID 28132810.
  8. Keeling, Patrick J. (2010). "The endosymbiotic origin, diversification and fate of plastids". Philosophical Transactions of the Royal Society B: Biological Sciences. 365 (1541): 729–748. doi:10.1098/rstb.2009.0103. PMC 2817223. PMID 20124341.
  9. Morozov, A.A.; Galachyants, Yuri P. (2019). "Diatom genes originating from red and green algae: Implications for the secondary endosymbiosis models". Marine Genomics. 45: 72–78. doi:10.1016/j.margen.2019.02.003. PMID 30792089.
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