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'''Chromalveolata''' is a [[eukaryote]] supergroup first proposed by [[Thomas Cavalier-Smith]] as a refinement of his [[kingdom (biology)|kingdom]] [[Chromista]], which was first proposed in 1981. As of 2006, it is often regarded as one of six major [[clade]]s of eukaryotes,<ref name="parfrey">{{cite journal | url = http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1713255 | title = Evaluating Support for the Current Classification of Eukaryotic Diversity | author = Laura Wegener Parfrey, Erika Barbero, Elyse Lasser, Micah Dunthorn, Debashish Bhattacharya, David J Patterson, and Laura A Katz | doi = 10.1371/journal.pgen.0020220 | journal = PLoS Genet. | date = 2006 December | volume = 2 | issue = 12 | pages = e220 | pmid = 17194223 }}</ref> although it is not yet clear whether it is indeed monophyletic<ref name="parfrey" /><ref>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. [http://www.botany.ubc.ca/keeling/PDF/05chromalvJSEM.pdf]</ref>. It may be considered a “kingdom”, though it is not given any formal taxonomical classification. It comprises a line descending from a [[bikont]] which underwent secondary [[endosymbiosis]] with a [[red algae|red alga]], and it includes these four major groups:
'''Chromalveolata''' is a [[eukaryote]] supergroup first proposed by [[Thomas Cavalier-Smith]] as a refinement of his [[kingdom (biology)|kingdom]] [[Chromista]], which was first proposed in 1981. It was proposed to represent the result of a single secondary [[endosymbiosis]] of a line descending from a [[bikont]] with a [[red algae|red alga]] that became the progenitor of [[chlorophyll]]-c containing [[plastid]]s. As of 2006, it was often regarded as one of six major [[clade]]s of eukaryotes,<ref name="parfrey">{{cite journal | url = http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1713255 | title = Evaluating Support for the Current Classification of Eukaryotic Diversity | author = Laura Wegener Parfrey, Erika Barbero, Elyse Lasser, Micah Dunthorn, Debashish Bhattacharya, David J Patterson, and Laura A Katz | doi = 10.1371/journal.pgen.0020220 | journal = PLoS Genet. | date = 2006 December | volume = 2 | issue = 12 | pages = e220 | pmid = 17194223 }}</ref> although it is hotly debated and does not appear to be monophyletic as originally proposed.<ref name="parfrey" /><ref>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. [http://www.botany.ubc.ca/keeling/PDF/05chromalvJSEM.pdf]</ref> Specifically, recent [[expressed sequence tag]] analysis has suggested that [[haptophyta]] and [[cryptophyta]] are less closely related, while [[Rhizaria]] groups in a clade dubbed the '''"SAR" super assemblage''' with [[Alveolata]] and [[stramenophile]]s.<ref>{{cite journal|url=http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0000790|title=Phylogenomics Reshuffles the Eukaryotic Supergroups|author=Fabien Burki, Kamran Shalchian-Tabrizi, Marianne Minge, Åsmund Skjæveland, Sergey I. Nikolaev, Kjetill S. Jakobsen, Jan Pawlowski|journal=PLoS ONE|volume=2|issue=8|doi=doi:10.1371/journal.pone.0000790}}</ref><ref>{{cite journal | doi = 10.1371/journal.pone.0002621 }}</ref>

Chromalveolates are not given any formal taxonomical classification, but may be considered a “kingdom”. They include the major groups:


*[[Heterokontophyta]]
*[[Heterokontophyta]]
*[[Alveolata]]
*[[Haptophyta]]
*[[Haptophyta]]
*[[Cryptophyta]]
*[[Cryptophyta]]
*[[Alveolata]]


Though several groups, such as the [[ciliate]]s and the [[water mold]]s, have lost the ability to photosynthesize, most are [[autotroph]]ic. All photosynthetic chromalveolates use [[chlorophyll]]s ''a'' and ''c'', and many use accessory pigments.
Though several groups, such as the [[ciliate]]s and the [[water mold]]s, have lost the ability to photosynthesize, most are [[autotroph]]ic. All photosynthetic chromalveolates use [[chlorophyll]]s ''a'' and ''c'', and many use accessory pigments.


==Evolutionary relationship==
==Evolutionary relationship==
<!-- Image with unknown copyright status removed: [[Image:Euktree.jpg|thumb|left|A graphic summary of the six major eukaryote groups.]] -->
Chromalveolata is part of the bikonts, which also comprise the [[Archaeplastida]], the [[Rhizaria]], the [[Excavata]], and some smaller, unresolved groups such as the [[Apusozoa]] and the [[Centrohelid]]a. As bikonts, they all descend from a [[heterotroph]]ic eukaryote with two [[flagella]]. It is also thought that the Chromalveolata share a closer relationship with the Archaeplastida than with the other groups, and some{{specify}} have proposed a clade called [[Corticata]] for this grouping.
Chromalveolata is part of the bikonts, which also comprise the [[Archaeplastida]], the [[Rhizaria]], the [[Excavata]], and some smaller, unresolved groups such as the [[Apusozoa]] and the [[Centrohelid]]a. As bikonts, they all descend from a [[heterotroph]]ic eukaryote with two [[flagella]]. It is also thought that the Chromalveolata share a closer relationship with the Archaeplastida than with the other groups, and some{{specify}} have proposed a clade called [[Corticata]] for this grouping.


Historically, many chromalveolates were considered [[plant]]s, with their cell walls, photosynthetic ability, and in some cases their morphological resemblance to the [[Embryophyta]]. However, when the five-kingdom system took prevalence over the animal-plant dichotomy, most chromalveolates were put into the kingdom [[Protista]], with the [[water mold]]s and [[Labyrinthulomycetes|slime nets]] put into the kingdom [[Fungi]], and the [[brown algae]] staying in the plant kingdom. Later, Chromalveolata was proposed as a monophyletic group, but the monophyly of this group is not yet established.<ref>{{cite journal | author = Burki F, Shalchian-Tabrizi K, Minge M, Skjæveland Å, Nikolaev SI, et al. | year = 2007 | title = Phylogenomics Reshuffles the Eukaryotic Supergroups | journal = PLoS ONE | volume = 2 | issue = 8: e790 | doi = 10.1371/journal.pone.0000790 | pages = e790}}</ref><ref name="parfrey" />
[[Image:Modeuktax.PNG|left|thumb|Modern eukaryote taxonomy.]]
Historically, many chromalveolates were considered [[plant]]s, with their cell walls, photosynthetic ability, and in some cases their morphological resemblance to the [[Embryophyta]]. However, when the five-kingdom system took prevalence over the animal-plant dichotomy, most chromalveolates were put into the kingdom [[Protista]], with the [[water mold]]s and [[Labyrinthulomycetes|slime nets]] put into the kingdom [[Fungi]], and the [[brown algae]] staying in the plant kingdom. After much research, Chromalveolata has been proposed as a monophyletic group, but the monophyly of this group is not yet established.<ref>{{cite journal | author = Burki F, Shalchian-Tabrizi K, Minge M, Skjæveland Å, Nikolaev SI, et al. | year = 2007 | title = Phylogenomics Reshuffles the Eukaryotic Supergroups | journal = PLoS ONE | volume = 2 | issue = 8: e790 | doi = 10.1371/journal.pone.0000790 | pages = e790}}</ref><ref name="parfrey" />


==Morphology==
==Morphology==

Revision as of 19:40, 6 January 2009

Chromalveolates
Ceratium hirundinella, a dinoflagellate
Clockwise from top-left: a haptophyte, some diatoms, a water mold, a cryptomonad, and Macrocystis, a phaeophyte
Scientific classification
Domain:
(unranked):
Kingdom:
Chromalveolata

Phyla

Chromalveolata is a eukaryote supergroup first proposed by Thomas Cavalier-Smith as a refinement of his kingdom Chromista, which was first proposed in 1981. It was proposed to represent the result of a single secondary endosymbiosis of a line descending from a bikont with a red alga that became the progenitor of chlorophyll-c containing plastids. As of 2006, it was often regarded as one of six major clades of eukaryotes,[1] although it is hotly debated and does not appear to be monophyletic as originally proposed.[1][2] Specifically, recent expressed sequence tag analysis has suggested that haptophyta and cryptophyta are less closely related, while Rhizaria groups in a clade dubbed the "SAR" super assemblage with Alveolata and stramenophiles.[3][4]

Chromalveolates are not given any formal taxonomical classification, but may be considered a “kingdom”. They include the major groups:

Though several groups, such as the ciliates and the water molds, have lost the ability to photosynthesize, most are autotrophic. All photosynthetic chromalveolates use chlorophylls a and c, and many use accessory pigments.

Evolutionary relationship

Chromalveolata is part of the bikonts, which also comprise the Archaeplastida, the Rhizaria, the Excavata, and some smaller, unresolved groups such as the Apusozoa and the Centrohelida. As bikonts, they all descend from a heterotrophic eukaryote with two flagella. It is also thought that the Chromalveolata share a closer relationship with the Archaeplastida than with the other groups, and some[specify] have proposed a clade called Corticata for this grouping.

Historically, many chromalveolates were considered plants, with their cell walls, photosynthetic ability, and in some cases their morphological resemblance to the Embryophyta. However, when the five-kingdom system took prevalence over the animal-plant dichotomy, most chromalveolates were put into the kingdom Protista, with the water molds and slime nets put into the kingdom Fungi, and the brown algae staying in the plant kingdom. Later, Chromalveolata was proposed as a monophyletic group, but the monophyly of this group is not yet established.[5][1]

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 Crytophyta, 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 in most cell walls

Since this is such a diverse group, it is difficult to summarize shared chromalveolate characteristics.

Ecological role

A potato plant infected with Phytophthora infestans.

Many chromalveolates affect our ecosystem in enormous ways. Some of these organisms can be very harmful. Dinoflagellates produce red tides which can devastate fish populations and intoxicate oyster harvests. Apicomplexans are some of the most successful specific parasites to animals. Water molds cause several plant diseases. In fact, it was a water mold, Phytophthora infestans, that caused the Irish potato famine.

A Californian kelp forest.

However, many chromalveolates are vital members of our ecosystem. Diatoms are one of the major photosynthetic producers, and produce much of the oxygen we breathe, and also take in much of the carbon dioxide that is thought to be a cause of global warming. Brown algae, most specifically kelps, 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 in brown algae is used a food thickener, most famously in ice cream. 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.

References

  1. ^ a b c Laura Wegener Parfrey, Erika Barbero, Elyse Lasser, Micah Dunthorn, Debashish Bhattacharya, David J Patterson, and Laura A Katz (2006 December). "Evaluating Support for the Current Classification of Eukaryotic Diversity". PLoS Genet. 2 (12): e220. doi:10.1371/journal.pgen.0020220. PMID 17194223. {{cite journal}}: Check date values in: |date= (help)CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  2. ^ 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. [1]
  3. ^ Fabien Burki, Kamran Shalchian-Tabrizi, Marianne Minge, Åsmund Skjæveland, Sergey I. Nikolaev, Kjetill S. Jakobsen, Jan Pawlowski. "Phylogenomics Reshuffles the Eukaryotic Supergroups". PLoS ONE. 2 (8). doi:doi:10.1371/journal.pone.0000790. {{cite journal}}: Check |doi= value (help)CS1 maint: multiple names: authors list (link)
  4. ^ . doi:10.1371/journal.pone.0002621. {{cite journal}}: Cite journal requires |journal= (help); Missing or empty |title= (help)CS1 maint: unflagged free DOI (link)
  5. ^ Burki F, Shalchian-Tabrizi K, Minge M, Skjæveland Å, Nikolaev SI; et al. (2007). "Phylogenomics Reshuffles the Eukaryotic Supergroups". PLoS ONE. 2 (8: e790): e790. doi:10.1371/journal.pone.0000790. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  • Patron, N. J., Rogers, M. B. and Keeling, P. J. 2004. Gene replacement of fructose-1,6-bisphophate aldolase supports the hypothesis of a single photosynthetic ancestor of chromalveolates. Eukaryot. Cell. 3, 1169-1174. [2]
  • Harper, J. T. & Keeling, P. J. 2003. Nucleus-encoded, plastid-targeted glyceraldehyde-3-phosphate dehydrogenase (GAPDH) indicates a single origin for chromalveolate plastids. Mol. Biol. Evol. 20, 1730-1735.
  • Archibald, J. M. & Keeling, P. J. 2002. Recycled plastids: a "green movement" in eukaryotic evolution. Trends Genet., 18, 577-584.
  • Fast, N. M., Kissinger, J. C., Roos, D. S., & Keeling, P. J. 2001. Nuclear-encoded, plastid-targeted genes suggest a single common origin for apicomplexan and dinoflagellate plastids. Mol. Biol. Evol., 18, 418-426.
  • Cavalier-Smith, T. 1999. Principles of protein and lipid targeting in secondary symbiogenesis: Euglenoid, dinoflagellate, and sporozoan plastid origins and the eukaryote family tree. J. Eukaryot. Microbiol. 46:347–66.