Chromalveolate diseases
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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 Kingdom commonly refers to: * A monarchy ruled by a king or queen * Kingdom (biology), a category in biological taxonomy Kingdom may also refer to: Arts and media Television * ''Kingdom'' (British TV series), a 2007 British television drama s ...
Chromista Chromista is a biological kingdom consisting of single-celled and multicellular eukaryotic species that share similar features in their photosynthetic organelles ( plastids). It includes all protists whose plastids contain chlorophyll ''c'', ...
, first proposed by
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. Chromalveolata was proposed to represent the organisms descended from a single secondary
endosymbiosis An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...
involving a
red alga 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 ...
and a
bikont A bikont ("two flagella") is any of the eukaryotic organisms classified in the group Bikonta. Many single-celled members of the group, and the presumed ancestor, have two flagella. Enzymes Another shared trait of bikonts is the fusion of two ge ...
. The plastids in these organisms are those that contain chlorophyll c. However, the
monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
of the Chromalveolata has been rejected. Thus, two papers published in 2008 have phylogenetic trees in which the chromalveolates are split up, and recent studies continue to support this view.


Groups and classification

Historically, many chromalveolates were considered
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclu ...
s, because of their cell walls, photosynthetic ability, and in some cases their morphological resemblance to the land plants (
Embryophyta The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as siste ...
). However, when the
five-kingdom system 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, Plan ...
(proposed in 1969) took prevalence over the animal–plant dichotomy, most of what we now call chromalveolates were put into the kingdom
Protista 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 exc ...
, but the
water mold 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 ...
s and slime nets were put into the kingdom
Fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
, while the
brown algae Brown algae (singular: alga), comprising the 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 temperate and p ...
stayed in the plant kingdom. These various organisms were later grouped together and given the name Chromalveolata by Cavalier-Smith. He believed them to be a monophyletic group, but this is not the case.Harper, J. T., Waanders, E. & Keeling, P. J. 2005. On the monophyly of chromalveolates using a six-protein phylogeny of eukaryotes. Int. J. System. Evol. Microbiol., 55, 487-496. In 2005, in a classification reflecting the consensus at the time, the Chromalveolata were regarded as one of the six major clades of eukaryotes. Although not given a formal taxonomic status in this classification, elsewhere the group had been treated as a Kingdom. The Chromalveolata were divided into four major subgroups: *
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 ...
*
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 ...
*
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 ...
(or Heterokontophyta) * Alveolata Other groups that may be included within, or related to, chromalveolates, are: *
Centrohelid The centrohelids or centroheliozoa are a large group of heliozoan protists. They include both mobile and sessile forms, found in freshwater and marine environments, especially at some depth. Characteristics Individuals are unicellular and spher ...
s * Katablepharids *
Telonemia Telonemia is a phylum of microscopic eukaryote, single-celled organisms. They were formerly classified as protists until that kingdom fell out of general use, and are suggested to have evolutionary significance in being a possible transitional fo ...
Though several groups, such as the
ciliate The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a differen ...
s and the
water mold 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 ...
s, have lost the ability to photosynthesize, most are
autotroph An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Wo ...
ic. All photosynthetic chromalveolates use chlorophylls ''a'' and ''c'', and many use
accessory pigment Accessory pigments are light-absorbing compounds, found in photosynthetic organisms, that work in conjunction with chlorophyll ''a''. They include other forms of this pigment, such as chlorophyll ''b'' in green algal and higher plant antennae, w ...
s. Chromalveolates share similar glyceraldehyde 3-phosphate dehydrogenase proteins. However, as early as 2005, doubts were being expressed as to whether Chromalveolata was monophyletic, and a review in 2006 noted the lack of evidence for several of the supposed six major eukaryote groups, including the Chromalveolata. In 2012, consensus emerged that the group is not monophyletic. The four original subgroups fall into at least two categories: one comprises the Stramenopiles and the Alveolata, to which the
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 forami ...
are now usually added to form the SAR group; the other comprises the Cryptophyta and the Haptophyta. A 2010 paper splits the Cryptophyta and Haptophyta; the former are a sister group to the SAR group, the latter cluster with the
Archaeplastida The Archaeplastida (or kingdom Plantae '' sensu lato'' "in a broad sense"; pronounced /ɑːrkɪ'plastɪdə/) are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor gr ...
(plants in the broad sense). The katablepharids are closely related to the cryptophytes and the telonemids and
centrohelid The centrohelids or centroheliozoa are a large group of heliozoan protists. They include both mobile and sessile forms, found in freshwater and marine environments, especially at some depth. Characteristics Individuals are unicellular and spher ...
s may be related to the haptophytes. A variety of names have been used for different combinations of the groups formerly thought to make up the Chromalveolata. * ''Halvaria'' Analyses in 2007 and 2008 agreed that the Stramenopiles and the Alveolata were related, forming a reduced chromalveolate clade, called
Halvaria Halvaria is a grouping that includes Alveolata and Heterokonta (Stramenopiles). Analyses in 2007 and 2008 revealed that the Stramenopiles and the Alveolata are related, and form a reduced chromalveolate clade. They group together with the Rhiza ...
. * ''SAR group'' The Rhizaria, which were originally not considered to be chromalveolates, belong with the Stramenopiles and Alveolata in many analyses, forming the SAR group, i.e. Halvaria plus Rhizaria. * ''Hacrobia'' The other two groups originally included in Chromalveolata, the Haptophyta and the Cryptophyta, were related in some analyses, forming a clade which has been called
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 ...
. Alternatively, the Hacrobia appeared to be more closely related to the
Archaeplastida The Archaeplastida (or kingdom Plantae '' sensu lato'' "in a broad sense"; pronounced /ɑːrkɪ'plastɪdə/) are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor gr ...
(plants in the very broad sense), being a sister group in one analysis, and actually nested inside this group in another. (Earlier,
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 ...
had suggested a clade called
Corticata Corticata ("one with a cortex"), in the classification of eukaryotes (living organisms with a cell nucleus), is a clade suggested by Thomas Cavalier-Smith to encompass the eukaryote supergroups of the following two groups: * Plantae, or Archaepl ...
for the grouping of ''all'' the chromalveolates and the Archaeplastida.) More recently, as noted above, Hacrobia has been split, with the Haptophyta being sister to the SAR group and the Cryptophyta instead related to the Archaeplastida.


Morphology

Chromalveolates, unlike other groups with multicellular representatives, do not have very many common morphological characteristics. Each major subgroup has certain unique features, including the alveoli of the Alveolata, the haptonema of the Haptophyta, the ejectisome of the Cryptophyta, and the two different flagella of the Heterokontophyta. However, none of these features are present in all of the groups. The only common chromalveolate features are these: *The shared origin of chloroplasts, as mentioned above *Presence of
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell w ...
in most cell walls Since this is such a diverse group, it is difficult to summarize shared chromalveolate characteristics.


Ecological role

Many chromalveolates affect our ecosystem in enormous ways. Some of these organisms can be very harmful. Dinoflagellates produce
red tide A harmful algal bloom (HAB) (or excessive algae growth) is an algal bloom that causes negative impacts to other organisms by production of natural algae-produced toxins, mechanical damage to other organisms, or by other means. HABs are sometimes ...
s, which can devastate fish populations and intoxicate oyster harvests. Apicomplexans are some of the most successful specific parasites to animals (including the genus ''
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 ver ...
'', the
malaria Malaria is a mosquito-borne infectious disease that affects humans and other animals. Malaria causes symptoms that typically include fever, tiredness, vomiting, and headaches. In severe cases, it can cause jaundice, seizures, coma, or death. S ...
parasites). Water molds cause several plant diseases - it was the water mold ''
Phytophthora infestans ''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 "p ...
'' that caused the Irish potato blight that led to the Great Irish Famine. However, many others are vital members of our ecosystem. Diatoms are one of the major photosynthetic producers, and as such produce much of the
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...
that we breathe, and also take in much of the
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...
from the atmosphere.
Brown algae Brown algae (singular: alga), comprising the 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 temperate and p ...
, most specifically
kelp Kelps are large brown algae seaweeds that make up the order Laminariales. There are about 30 different genera. Despite its appearance, kelp is not a plant - it is a heterokont, a completely unrelated group of organisms. Kelp grows in "underwa ...
s, create underwater "forest" habitats for many marine creatures, and provide a large portion of the diet of coastal communities. Chromalveolates also provide many products that we use. The
algin Alginic acid, also called algin, is a naturally occurring, edible polysaccharide found in brown algae. It is hydrophilic and forms a viscous gum when hydrated. With metals such as sodium and calcium, its salts are known as alginates. Its colour ...
in brown algae is used as a food thickener, most famously in
ice cream Ice cream is a sweetened frozen food typically eaten as a snack or dessert. It may be made from milk or cream and is flavoured with a sweetener, either sugar or an alternative, and a spice, such as cocoa or vanilla, or with fruit such as ...
. The siliceous shells of diatoms have many uses, such as in reflective paint, in toothpaste, or as a filter, in what is known as diatomaceous earth.


Chromalveolata viruses

Like other organisms, chromalveolata have viruses. In the case of ''
Emiliania huxleyi ''Emiliania huxleyi'' is a species of coccolithophore found in almost all ocean ecosystems from the equator to sub-polar regions, and from nutrient rich upwelling zones to nutrient poor oligotrophic waters. It is one of thousands of different ...
'' (a common
algal bloom An algal bloom or algae bloom is a rapid increase or accumulation in the population of algae in freshwater or marine water systems. It is often recognized by the discoloration in the water from the algae's pigments. The term ''algae'' encompass ...
chromalveolate), a virus believed to be specific to it causes mass death and the end of the bloom.


See also

*
Halvaria Halvaria is a grouping that includes Alveolata and Heterokonta (Stramenopiles). Analyses in 2007 and 2008 revealed that the Stramenopiles and the Alveolata are related, and form a reduced chromalveolate clade. They group together with the Rhiza ...


References


External links


Tree of Life Eukaryotes
{{Taxonbar, from=Q477950 Algae Obsolete eukaryote taxa Bikont unranked clades