Alveolate

From Wikipedia, the free encyclopedia

The alveolates (meaning "with cavities")[2] are a group of protists,

Alveolata
considered a major clade[3] and superphylum[4] within Eukarya, and are
[1]
also called Alveolata.[5] Temporal range: Ediacaran -
Recent

Contents
1 Characteristics
2 Classification
2.1 Phylogeny
2.2 Taxonomy Ceratium furca
3 Development
4 Evolution Scientific classification
5 References Domain: Eukaryota
6 External links
(unranked): SAR
(unranked): Alveolata
Characteristics Cavalier-Smith, 1991

Phyla
The most notable shared characteristic is the presence of cortical (outer-
region) alveoli (sacs). These are flattened vesicles (sacs) packed into a
continuous layer just under the membrane and supporting it, typically Apicomplexa
forming a flexible pellicle (thin skin). In dinoflagellates they often form Chromerida
armor plates. Alveolates have mitochondria with tubular cristae (ridges), Ciliophora
and their flagellae or cilia have a distinct structure. Dinoflagellata

Almost all sequenced mitochondrial genomes of ciliates and

apicomplexia are linear.[6] The mitochondrial genome of Babesia microti is circular.[7] This species is also now
known not to belong to either of the genera Babesia or Theileria and a new genus will have to be created for it.

Classification
Alveolata comprises around 9 major and minor groups, which are very diverse in form, and are known to be
related by various ultrastructural and genetic similarities:[8]

Ciliates very common protozoa with many short cilia arranged in rows, and two nuclei
Acavomonidia[8]
Colponemidia[8]
Dinoflagellates s.l. mostly marine flagellates many of which have chloroplasts
Perkinsozoa
Chromerida a marine phylum of photosynthetic protozoa
Colpodellida
Voromonadida
Apicomplexa parasitic protozoa that lack axonemal locomotive structures except in gametes

The Acavomonidia and Colponemidia were previously grouped together as colponemids, a taxon now split
based on ultrastructural analysis. The Acavomonidia are closer to the dinoflagellate/perkinsid group than the
Colponemidia are.[9] As such, the informal term "colponemids", as it stands currently, covers two non-sister
groups within Alveolata: the Acavomonidia and the Colponemidia.[8]
The Apicomplexa and dinoflagellates may be more closely related to each other than to the ciliates. Both have
plastids, and most share a bundle or cone of microtubules at the top of the cell. In apicomplexans this forms
part of a complex used to enter host cells, while in some colorless dinoflagellates it forms a peduncle used to
ingest prey. Various other genera are closely related to these two groups, mostly flagellates with a similar apical
structure. These include free-living members in Oxyrrhis and Colponema, and parasites in Perkinsus,[10]
Parvilucifera, Rastrimonas and the ellobiopsids. In 2001, direct amplification of the rRNA gene in marine
picoplankton samples revealed the presence of two novel alveolate linages, called group I and II.[11][12] Group I
has no cultivated relatives, while group II is related to the dinoflagellate parasite Amoebophrya, which was
classified until now in the Syndiniales dinoflagellate order.

Relationships between some of these major groups were suggested during the 1980s, and a specific relationship
between all three was confirmed in the early 1990s by genetic studies, most notably by Gajadhar et al.[13]
Cavalier-Smith introduced the formal name Alveolata in 1991,[5] although at the time he actually considered
the grouping to be a paraphyletic assemblage, rather than a monophyletic group.

Some studies suggested the haplosporids, mostly parasites of marine invertebrates, might belong here, but they
lack alveoli and are now placed among the Cercozoa.

Phylogeny

Based on a compilation of the following works.[14][15]

Alveolata Ciliophora
Heterotrichea
Postciliodesmatophora

Karyorelictea

Mesodiniea
Intramacronucleata
Litostomatea

Lamellicorticata Armophorea

Cariacotrichea

Spirotrichea

Protocruziea

?Discotrichida

Colpodea

Ventrata Nassophorea

Phyllopharyngea
 Prostomatea

Plagiopylea

Oligohymenophorea
Miozoa
Colponemea

Acavomonadea
Myzozoa
?Voromonadida

Chromerida

Colpodellida
Apicomplexa s.l.
Blastogregarinida

Sporozoa Paragregarea

Coccidiomorphea

Gregarinomorphea

?Myzomonadea

?Squirmidea

Perkinsea
Dinoflagellata s.s.
?Acrocoelida

?Rastromonadida

Dinozoa
?Pronoctilucea

Ellobiophyceae

Psammosea

Oxyrrhea
 Syndinea

Noctiluciphyceae
Dinokaryota

Dinophyceae

Taxonomy

Alveolata Cavalier-Smith 1991 [Alveolatobiontes]

Phylum Ciliophora Doflein 1901 stat. n. Copeland 1956 [Ciliata Perty 1852 ; Infusoria Btschli 1887 ; Ciliae,
Ciliozoa, Cytoidea, Eozoa, Heterocaryota, Heterokaryota]
Class ?Mesodiniea Chen et al. 2015
Subphylum Postciliodesmatophora Gerassimova & Seravin 1976
Class Heterotrichea Stein 1859
Class Karyorelictea Corliss 1974
Subphylum Intramacronucleata Lynn 1996
Infraphylum Lamellicorticata
Class Litostomatea Small & Lynn 1981
Class Armophorea Lynn 2004
Class Cariacotrichea Orsi et al. 201 1
Class Spirotrichea Btschli 1889
Infraphylum Ventrata Cavalier-Smith 2004 [Conthreep Lynn 2012 ]
Order ?Discotrichida Chen et al. 2015
Class Protocruziea Chen et al. 2015 [Protocruziidia de Puytorac, Grain & Mignot 1987 ]
Class Colpodea Small & Lynn 1981
Class Nassophorea Small & Lynn 1981
Class Phyllopharyngea de Puytorac et al. 1974
Class Prostomatea Schewiakof f 1896
Class Plagiopylea Small & Lynn 1985 sensu L ynn 2008
Class Oligohymenophorea de Puytorac et al. 1974
Phylum Miozoa Cavalier-Smith 1987
Subphylum Colponemidia Tikhonenkov, Mylnikov & Keeling 2013
Class Colponemea Cavalier-Smith 1993
Subphylum Acavomonadia Tikhonenkov et al. 2014
Class Acavomonadea Tikhonenkov et al. 2014
Subphylum Myzozoa Cavalier-Smith 2004
Infraphylum Apicomplexa Levine 1970 emend. Adl et al. 2005
Order ?Voromonadida Cavalier-Smith & Chao 2004
Order Chromerida Moore et al. 2008
Order Colpodellida Patterson & Zlf fel 1991 [Spiromonadida Krylov & Mylnikov 1986 ]
Superclass Sporozoa Leuckart 1879 stat. nov . Cavalier-Smith 2013 [Gamontozoa]
Class Blastogregarinida Chatton & Villeneuve 1936 [Blastogregarinina;
Blastogregarinorina Chatton & Villeneuve 1936 ]
Class Paragregarea Cavalier-Smith 2014
Class Gregarinomorphea Grass 1953
Class Coccidiomorphea Doflein 1901
Infraphylum Dinozoa Cavalier-Smith 1981 emend. 2003
Class Myzomonadea Cavalier-Smith & Chao 2004 sensu Ruggiero et al. 2015
Class Squirmidea Norn 1999 stat. nov . Cavalier-Smith 2014
Superclass Perkinsozoa Norn et al. 1999 s.s.
Class Perkinsea Levine 1978 [Perkinsasida Levine 1978 ]
 Superclass Dinoflagellata Butschli 1885 stat. nov . Cavalier-Smith 1999 s ensu Cavalier -Smith
2013 [Dinozoa Cavalier-Smith 1981 ]
Order ?Acrocoelida Cavalier-Smith & Chao 2004
Order ?Rastromonadida Cavalier-Smith & Chao 2004
Class Pronoctilucea
Class Ellobiopsea Cavalier-Smith 1993 [Ellobiophyceae Loeblich III 1970 ;
Ellobiopsida Whisler 1990 ]
Class Psammosea
Class Oxyrrhea Cavalier-Smith 1987
Class Syndinea Chatton 1920 s.l. [Syndiniophyceae Loeblich III 1970 s.s. ; Syndina
Cavalier-Smith ]
Class Noctiluciphyceae Fensome et al. 1993 [Noctilucae Haeckel 1866 ; Noctilucea
Haeckel 1866 stat. nov .; Cystoflagellata Haeckel 1873 stat. nov . Butschli 1887 ]
Class Dinophyceae Pascher 1914 [Peridinea Ehrenberg 1830 stat. nov . Wettstein ]

Development
The development of plastids among the alveolates is intriguing. Cavalier-Smith proposed the alveolates
developed from a chloroplast-containing ancestor, which also gave rise to the Chromista (the chromalveolate
hypothesis). Other researchers have speculated that the alveolates originally lacked plastids and possibly the
dinoflagellates and Apicomplexa acquired them separately. However, it now appears that the alveolates, the
dinoflagellates, the Chromerida and the heterokont algae acquired their plastids from a red alga with evidence
of a common origin of this organelle in all these four clades.[16]

Evolution
A Bayesian estimate places the evolution of the alveolate group at ~850 million years ago.[17] The Alveolata
consist of Myzozoa, Ciliates, and Colponemids. In other words, the term Myzozoa, meaning "to siphon the
contents from prey", may be applied informally to the common ancestor of the subset of alveolates that are
neither ciliates nor colponemids. Predation upon algae is an important driver in alveolate evolution, as it can
provide sources for endosymbiosis of novel plastids. The term Myzozoa is therefore a handy concept for
tracking the history of the alveolate phylum.

The ancestors of the alveolate group may have been photosynthetic.[18] The ancestral alveolate probably
possessed a plastid. Chromerids, apicomplexans, and peridinin dinoflagellates have retained this organelle.[19]
Going one step even further back, the chromerids, the peridinin dinoflagellates and the heterokont algae possess
a monophyletic plastid lineage in common, i.e. acquired their plastids from a red alga,[16] and so it seems likely
that the common ancestor of alveolates and heterokonts was also photosynthetic.

In one school of thought the common ancestor of the dinoflagellates, apicomplexans, Colpodella, Chromerida,
and Voromonas was a myzocytotic predator with two heterodynamic flagella, micropores, trichocysts,
rhoptries, micronemes, a polar ring and a coiled open sided conoid.[20] While the common ancestor of
alveolates may also have possessed some of these characteristics, it has been argued that Myzocytosis was not
one of these characteristics, as ciliates ingest prey by a different mechanism.[8]

An ongoing debate concerns the number of membranes surrounding the plastid across apicomplexans and
certain dinoflagellates, and the origin of these membranes. This ultrastructural character can be used to group
organisms and if the character is in common, it can imply that phyla had a common photosynthetic ancestor. On
the basis that apicomplexans possess a plastid surrounded by 4 membranes, and that peridinin dinoflagellates
possess a plastid surrounded by 3 membranes, Petersen et al.[21] have been unable to rule our that the shared
stramenopile-alveolate plastid could have been recycled multiple times in the alveolate phylum, the source
being stramenopile-alveolate donors, through the mechanism of ingestion and endosymbiosis.
Ciliates are a model alveolate, having been genetically studied in great depth over the longest period of any
alveolate lineage. They are unusual among eukaryotes in that reproduction involves a micronucleus and a
macronucleus. Their reproduction is easily studied in the lab, and made them a model eukaryote historically.
Being entirely predatory and lacking any remnant plastid, their development as a phylum illustrates how
predation and autotrophy[18] are in dynamic balance and that the balance can swing one way or other at the
point of origin of a new phylum from mixotrophic ancestors, causing one ability to be lost.

Paramecium caudatum Vorticella (Ciliophora) Plasmodium falciparum Eimeria maxima

(Ciliophora) (left) (Apicomplexa) in blood (Apicomplexa)

Dinophysis acuminata
(Dinoflagellata)

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External links
Tree of Life: Alveolates

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