Chromista is a
biological kingdom
In biology, a kingdom is the second highest taxonomic rank, just below domain. Kingdoms are divided into smaller groups called phyla.
Traditionally, some textbooks from the United States and Canada used a system of six kingdoms (Animalia, Plant ...
consisting of single-celled and multicellular
eukaryotic
Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
species that share similar features in their
photosynthetic
Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in c ...
organelles (
plastid
The plastid (Greek: πλαστός; plastós: formed, molded – plural plastids) is a membrane-bound organelle found in the Cell (biology), cells of plants, algae, and some other eukaryotic organisms. They are considered to be intracellular endosy ...
s).
It includes all
protists
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the excl ...
whose plastids contain
chlorophyll ''c'', such as some
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
,
diatoms
A diatom (New Latin, Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group com ...
,
oomycetes
Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the result ...
, and
protozoans
Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
. It is probably a
polyphyletic
A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...
group whose members independently arose as a separate evolutionary group from the common ancestor of all eukaryotes.
As it is assumed the
last common ancestor
In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
already possessed
chloroplasts
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, ...
of
red algal
Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
origin, the non-photosynthetic forms evolved from ancestors able to perform photosynthesis. Their plastids are surrounded by four
membranes
A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. Bi ...
, and are believed to have been acquired from some red algae.
Chromista as a biological kingdom was created by British biologist
Thomas Cavalier-Smith
Thomas (Tom) Cavalier-Smith, FRS, FRSC, NERC Professorial Fellow (21 October 1942 – 19 March 2021), was a professor of evolutionary biology in the Department of Zoology, at the University of Oxford.
His research has led to discov ...
in 1981 to differentiate some protists from typical protozoans and plants.
According to Cavalier-Smith, the kingdom originally included only algae, but his later analysis indicated that many protozoa 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
''Phytophthora infestans'' is an oomycete or water mold, a fungus-like microorganism that causes the serious potato and tomato disease known as late blight or potato blight. Early blight, caused by ''Alternaria solani'', is also often called "po ...
,
dinoflagellates
The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates a ...
, ''
Paramecium
''
''Paramecium'' ( , ; also spelled ''Paramoecium'') is a genus of eukaryotic, unicellular ciliates, commonly studied as a representative of the ciliate group. ''Paramecia'' are widespread in freshwater, brackish, and marine environments and a ...
'', the brain parasite ''
Toxoplasma
''Toxoplasma gondii'' () is an obligate intracellular parasitic protozoan (specifically an apicomplexan) that causes toxoplasmosis. Found worldwide, ''T. gondii'' is capable of infecting virtually all warm-blooded animals, but felids, such as d ...
,'' and the malarial parasite ''
Plasmodium
''Plasmodium'' is a genus of unicellular eukaryotes that are obligate parasites of vertebrates and insects. The life cycles of ''Plasmodium'' species involve development in a blood-feeding insect host which then injects parasites into a vert ...
''.
Biology
Members of Chromista are single-celled and multicellular eukaryotes having basically either or both features:
[
#plastid(s) that contain chlorophyll ''c'' and lie within an extra (periplastid) membrane in the lumen of the rough endoplasmic reticulum (typically within the perinuclear cisterna);
#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.] Molecular evidence indicate that the plastids in chromists were derived from red algae through secondary symbiogenesis
Symbiogenesis (endosymbiotic theory, or serial endosymbiotic theory,) is the leading evolutionary theory of the origin of eukaryotic cells from prokaryotic organisms. The theory holds that mitochondria, plastids such as chloroplasts, and possi ...
in a single event. (In contrast, plants acquired their plastids from cyanobacteria through primary symbiogenesis.) 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
Membrane proteins are common proteins that are part of, or interact with, biological membranes. Membrane proteins fall into several broad categories depending on their location. Integral membrane proteins are a permanent part of a cell membrane ...
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. There was also additional symbiogenesis of green algae
The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
, the genes of which are retained in some members (such as heterokonts), as well as bacterial chlorophyll (indicated by the presence of ribosomal protein L36 gene, ''rpl36'') in haptophytes and cryptophytes.
History and groups
The name Chromista was first introduced by Cavalier-Smith
Thomas (Tom) Cavalier-Smith, FRS, FRSC, NERC Professorial Fellow (21 October 1942 – 19 March 2021), was a professor of evolutionary biology in the Department of Zoology, at the University of Oxford.
His research has led to disc ...
in 1981;[ the earlier names Chromophyta, Chromobiota and Chromobionta correspond to roughly the same group.
It has been described as consisting of three different groups:]
*Heterokont
Heterokonts are a group of protists (formally referred to as Heterokonta, Heterokontae or Heterokontophyta). The group is a major line of eukaryotes. Most are algae, ranging from the giant multicellular kelp to the unicellular diatoms, which ...
s or stramenopiles: brown alga
Brown algae (singular: alga), comprising the class (biology), class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown algae are the major seaweeds of the ...
e, diatom
A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s, water mould
Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the r ...
s, etc.
*Haptophyte
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for '' Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at ...
s
*Cryptomonad
The cryptomonads (or cryptophytes) are a group of algae, most of which have plastids. They are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape, with an anteri ...
s
Cavalier-Smith later (in 2009) stated his reason for making a new kingdom, saying:
Since then Chromista has been defined in different ways at different times. In 2010, Cavalier-Smith himself indicated his desire to move Alveolata, Rhizaria
The Rhizaria are an ill-defined but species-rich supergroup of mostly unicellular eukaryotes. Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthethic, but many foramini ...
and Heliozoa
Heliozoa, commonly known as sun-animalcules, are microbial eukaryotes ( protists) with stiff arms ( axopodia) radiating from their spherical bodies, which are responsible for their common name. The axopodia are microtubule-supported projections f ...
into Chromista.
Some examples of classification of the Chromista and related groups are shown below.
Chromophycées (Chadefaud, 1950)
The Chromophycées (Chadefaud, 1950), renamed Chromophycota (Chadefaud, 1960), included the current Ochrophyta
The ochrophytes, subphylum Ochrophytina, is a group of mostly photosynthetic heterokonts. Their plastid is of red algal origin.
The classification of the group is still being worked out. Originally, the ochrophytes were regarded as a phylum den ...
(autotrophic Stramenopile
Stramenopile is a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have be ...
s), Haptophyta
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for ''Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at t ...
(included in Chrysophyceae until Christensen, 1962), Cryptophyta
The cryptophyceae are a class of algae, most of which have plastids. About 220 species are known, and they are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape, ...
, Dinophyta
The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
, Euglenophyceae and Choanoflagellida (included in Chrysophyceae until Hibberd, 1975).
Chromophyta (Christensen 1962, 1989)
The Chromophyta (Christensen 1962, 2008), defined as algae with chlorophyll c
Chlorophyll ''c'' is a form of chlorophyll found in certain marine algae, including the photosynthetic Chromista (e.g. diatoms and brown algae) and dinoflagellates.
It has a blue-green color and is an accessory pigment, particularly significant i ...
, included the current Ochrophyta
The ochrophytes, subphylum Ochrophytina, is a group of mostly photosynthetic heterokonts. Their plastid is of red algal origin.
The classification of the group is still being worked out. Originally, the ochrophytes were regarded as a phylum den ...
(autotrophic Stramenopile
Stramenopile is a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have be ...
s), Haptophyta
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for ''Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at t ...
, Cryptophyta
The cryptophyceae are a class of algae, most of which have plastids. About 220 species are known, and they are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape, ...
, Dinophyta
The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
and Choanoflagellida. The Euglenophyceae were transferred to the Chlorophyta
Chlorophyta or Prasinophyta is a taxon of green algae informally called chlorophytes. The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it refers to ...
.
Chromophyta (Bourrelly, 1968)
The Chromophyta (Bourrelly, 1968) included the current Ochrophyta
The ochrophytes, subphylum Ochrophytina, is a group of mostly photosynthetic heterokonts. Their plastid is of red algal origin.
The classification of the group is still being worked out. Originally, the ochrophytes were regarded as a phylum den ...
(autotrophic Stramenopile
Stramenopile is a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have be ...
s), Haptophyta
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for ''Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at t ...
and Choanoflagellida. The Cryptophyceae
The cryptophyceae are a class of algae, most of which have plastids. About 220 species are known, and they are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape ...
and the Dinophyceae
Dinophyceae is a class of dinoflagellates.
Taxonomy
* Class Dinophyceae Pascher 1914 eridinea Ehrenberg 1830 stat. nov. Wettstein; Blastodiniphyceae Fensome et al. 1993 orthog. emend.** Order Haplozoonales aplozooidea Poche 1913*** Family Ha ...
were part of Pyrrhophyta
The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
(= Dinophyta).
Chromista (Cavalier-Smith, 1981)
The Chromista (Cavalier-Smith, 1981) included the current Stramenopile
Stramenopile is a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have be ...
s, Haptophyta
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for ''Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at t ...
(grouped together as Chromophyta) and Cryptophyta
The cryptophyceae are a class of algae, most of which have plastids. About 220 species are known, and they are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape, ...
.
Chromalveolata (Adl ''et al''., 2005)
The Chromalveolata
Chromalveolata was a eukaryote supergroup present in a major classification of 2005, then regarded as one of the six major groups within the eukaryotes.
It was a refinement of the kingdom Chromista, first proposed by Thomas Cavalier-Smith in 1 ...
(Cavalier-Smith, 1981) included Stramenopile
Stramenopile is a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have be ...
s, Haptophyta
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for ''Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at t ...
, Cryptophyta
The cryptophyceae are a class of algae, most of which have plastids. About 220 species are known, and they are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape, ...
and Alveolata.
Classification
In his original classification in 1981, Cavalier-Smith included three phyla under Chromista:[
#Heterokonta
#Haptophyta
#Cryptomonada
In 2010, Cavalier-Smith reorganised Chromista to include ]SAR supergroup
The SAR supergroup, also just SAR or Harosa, is a clade that includes stramenopiles (heterokonts), alveolates, and Rhizaria. The name is an acronym derived from the first letters of each of these clades; it has been alternatively spelled "RAS". T ...
(such as Stramenopiles
Stramenopile is a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have be ...
, Alveolata and Rhizaria
The Rhizaria are an ill-defined but species-rich supergroup of mostly unicellular eukaryotes. Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthethic, but many foramini ...
) and Hacrobia
The cryptomonads-haptophytes assemblage is a proposed monophyletic grouping of unicellular eukaryotes that are not included in the SAR supergroup. Several alternative names have been used for the group, including Hacrobia (derived from "ha-" refe ...
(Haptista
Haptista is a proposed group of protists made up of centrohelids and haptophytes. Phylogenomic studies indicate that Haptista, together with ''Ancoracysta twista'', forms a sister clade to the SAR+Telonemia supergroup, but it may also be siste ...
, Cryptista
Cryptista is a clade of algae-like eukaryotes. It is most likely related to Archaeplastida which includes plants and many algae, within the larger group Diaphoretickes.
Although it has sometimes placed along with Haptista in the group Hacrobia, ...
).
In 2015, 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 11 phyla, namely:
#Subkingdom Harosa (=Supergroup SAR)
##Phylum Ciliophora
##Phylum Miozoa
##Phylum Bigyra
##Phylum Ochrophyta
##Phylum Pseudofungi (=Oomycota)
##Phylum Cercozoa
##Phylum Retaria
#Subkingdom Hacrobia
##Phylum N.N. (=''nomen nominandum'', name unknown)—i.e. the classes Endohelea, Picomonadea and Telonemea
##Phylum Cryptista
##Phylum Haptophyta
##Phylum Heliozoa
Cavalier-Smith made a new analysis of Chromista in 2018 in which he classified all chromists into 8 phyla (Gyrista corresponds to the above phyla Ochrophyta and Pseudofungi, Cryptista corresponds to the above phyla Cryptista and "N.N.", Haptista corresponds to the above phyla Haptophyta and Heliozoa):
#Subkingdom Harosa
##Phylum Miozoa
##Phylum Ciliophora
##Phylum Gyrista
##Phylum Bigyra
##Phylum Cercozoa
##Phylum Retaria
#Subkingdom Hacrobia
##Phylum Cryptista
##Phylum Haptista
Phylogeny
Controversy
Molecular trees have had some difficulty resolving relationships between the different groups. All three may share a common ancestor with the alveolate
The alveolates (meaning "pitted like a honeycomb") are a group of protists, considered a major clade and Biological classification, superphylum within Eukarya. They are currently grouped with the stramenopiles and Rhizaria among the protists with ...
s (see chromalveolate
Chromalveolata was a eukaryote supergroup present in a major classification of 2005, then regarded as one of the six major groups within the eukaryotes.
It was a refinement of the kingdom Chromista, first proposed by Thomas Cavalier-Smith in 1 ...
s), 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. Cryptista specifically may be sister or part of Archaeplastida.
See also
* Cabozoa
In the classification of eukaryotes (living organisms with a cell nucleus), Cabozoa was a taxon proposed by Cavalier-Smith.
It was a putative clade comprising the Rhizaria and Excavata. More recent research tends to place the Rhizaria with the ...
* Cavalier-Smith's system of classification
The biological classification system of life introduced by British zoologist Thomas Cavalier-Smith involves systematic arrangements of all life forms on earth. Following and improving the classification systems introduced by Carl Linnaeus, Ernst ...
* List of Chromista by conservation status
, the International Union for Conservation of Nature (IUCN) has evaluated the conservation status of 15 species within Chromista.
The IUCN has not evaluated any protist species other than those in Chromista. No Chromista subspecies or subpopulati ...
References
External links
UCMP: Introduction to the Chromista
{{Taxonbar, from=Q862296
Obsolete eukaryote taxa
Kingdoms (biology)