Warnowiaceae

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Warnowiaceae
Bmc evol bio hoppenrath proterythropsis ocelloid fig1a.png
A light micrograph of an ocelloid-containing dinoflagellate from the genus Proterythropsis. The nucleus is marked n, the ocelloid is indicated with a double arrowhead, and a posterior cell extension is indicated with an arrow; scale bar = 10 µm. [1]
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Clade: Diaphoretickes
Clade: SAR
Clade: Alveolata
Phylum: Myzozoa
Superclass: Dinoflagellata
Class: Dinophyceae
Order: Gymnodiniales
Family: Warnowiaceae
Genera

The Warnowiaceae are a family of athecate dinoflagellates (a diverse group of unicellular eukaryotes). Members of the family are known as warnowiids. The family is best known for a light-sensitive subcellular structure known as the ocelloid, a highly complex arrangement of organelles with a structure directly analogous to the eyes of multicellular organisms. The ocelloid has been shown to be composed of multiple types of endosymbionts, namely mitochondria and at least one type of plastid. [2]

Contents

Habitat and life cycle

Warnowiids are found in marine plankton but are very rare in most plankton samples. Little is known about their life histories because they cannot be cultured in the laboratory, and samples obtained from the natural environment do not survive well under laboratory conditions. Studies of wild samples have found evidence of distinctive structures called trichocysts in warnowiid cell vacuoles, suggesting that their prey might be other dinoflagellates. Despite the complexity of the ocelloid, the experimental difficulty of working with the cells has prevented experimental study of light-directed behavior such as phototaxis. [2]

Taxonomy

The family contains eight recognized genera. [3] Descriptions of genera and species within the family have been complicated by complex morphological changes during the life cycle and in response to the environment, and the systematics of this group is not currently well defined. [1]

Subcellular structures

The warnowiids as a group possess unusually complex subcellular structures. The ocelloid light sensitive structure is recognized as a synapomorphic character of warnowiids. [1]

Other complex subcellular structures, such as nematocysts, trichocysts, and pistons, are present in some (but not all) warnowiids and are shared with the polykrikoid dinoflagellates, the closest extant relatives as defined by molecular phylogenetics. [1] [4]

Ocelloid function and origin

The ocelloid of warnowiids functions similarly to eyes found in much larger organisms, containing structures similar to a retina and lens. It is receptive only to the polarized light that is created as light passes through the thecal plates of other dinoflagellates. [2] Because dinoflagellates are the main source of food for warnowiids, this trait is particularly useful for locating prey.


A gene fragment that is expressed in the rhodopsin of the retinal body of the ocelloid has been shown to be most closely related to those of bacteria, suggesting a bacterial endosymbiont as the origin of the organelle. [5]

Related Research Articles

In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name organelle comes from the idea that these structures are parts of cells, as organs are to the body, hence organelle, the suffix -elle being a diminutive. Organelles are either separately enclosed within their own lipid bilayers or are spatially distinct functional units without a surrounding lipid bilayer. Although most organelles are functional units within cells, some function units that extend outside of cells are often termed organelles, such as cilia, the flagellum and archaellum, and the trichocyst.

<span class="mw-page-title-main">Dinoflagellate</span> Unicellular algae with two flagella

The dinoflagellates are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered protists. Dinoflagellates are mostly marine plankton, but they also are common in freshwater habitats. Their populations vary with sea surface temperature, salinity, and depth. Many dinoflagellates are photosynthetic, but a large fraction of these are in fact mixotrophic, combining photosynthesis with ingestion of prey.

<i>Synechococcus</i> Genus of bacteria

Synechococcus is a unicellular cyanobacterium that is very widespread in the marine environment. Its size varies from 0.8 to 1.5 µm. The photosynthetic coccoid cells are preferentially found in well–lit surface waters where it can be very abundant. Many freshwater species of Synechococcus have also been described.

<span class="mw-page-title-main">Trichocyst</span> Organelle

A trichocyst is an organelle found in certain ciliates and dinoflagellates.

<i>Symbiodinium</i> Genus of dinoflagellates (algae)

Symbiodinium is a genus of dinoflagellates that encompasses the largest and most prevalent group of endosymbiotic dinoflagellates known. These unicellular microalgae commonly reside in the endoderm of tropical cnidarians such as corals, sea anemones, and jellyfish, where the products of their photosynthetic processing are exchanged in the host for inorganic molecules. They are also harbored by various species of demosponges, flatworms, mollusks such as the giant clams, foraminifera (soritids), and some ciliates. Generally, these dinoflagellates enter the host cell through phagocytosis, persist as intracellular symbionts, reproduce, and disperse to the environment. The exception is in most mollusks, where these symbionts are intercellular. Cnidarians that are associated with Symbiodinium occur mostly in warm oligotrophic (nutrient-poor), marine environments where they are often the dominant constituents of benthic communities. These dinoflagellates are therefore among the most abundant eukaryotic microbes found in coral reef ecosystems.

<span class="mw-page-title-main">Gymnodiniales</span> Order of single-celled organisms

The Gymnodiniales are an order of dinoflagellates, of the class Dinophyceae. Members of the order are known as gymnodinioid or gymnodinoid. They are athecate, or lacking an armored exterior, and as a result are relatively difficult to study because specimens are easily damaged. Many species are part of the marine plankton and are of interest primarily due to being found in algal blooms. As a group the gymnodinioids have been described as "likely one of the least known groups of the open ocean phytoplankton."

<i>Ornithocercus</i> Genus of single-celled organisms

Ornithocercus is a genus of planktonic dinoflagellate that is known for its complex morphology that features considerable lists growing from its thecal plates, giving an attractive appearance. Discovered in 1883, this genus has a small number of species currently categorized but is widespread in tropical and sub-tropical oceans. The genus is marked by exosymbiotic bacteria gardens under its lists, the inter-organismal dynamics of which are a current field of research. As they reside only in warm water, the genus has been used as a proxy for climate change and has potential to be an indicator species for environmental change if found in novel environments.

<span class="mw-page-title-main">Marine microorganisms</span> Any life form too small for the naked human eye to see that lives in a marine environment

Marine microorganisms are defined by their habitat as microorganisms living in a marine environment, that is, in the saltwater of a sea or ocean or the brackish water of a coastal estuary. A microorganism is any microscopic living organism or virus, that is too small to see with the unaided human eye without magnification. Microorganisms are very diverse. They can be single-celled or multicellular and include bacteria, archaea, viruses and most protozoa, as well as some fungi, algae, and animals, such as rotifers and copepods. Many macroscopic animals and plants have microscopic juvenile stages. Some microbiologists also classify viruses as microorganisms, but others consider these as non-living.

<i>Polykrikos kofoidii</i> Species of single-celled organism

Polykrikos kofoidii is a species of phagotrophic marine pseudocolonial dinoflagellates that can capture and engulf other protist prey, including the toxic dinoflagellate, Alexandrium tamarense. P. kofoidii is of scientific interest due to its status as a predator of other dinoflagellates, a behavior that is significant in the control of algal blooms. It has a complex life cycle of both vegetative (asexual) and sexual reproduction complicated by its pseudocolonial structure.

<i>Mesodinium rubrum</i> Species of single-celled organism

Mesodinium rubrum is a species of ciliates. It constitutes a plankton community and is found throughout the year, most abundantly in spring and fall, in coastal areas. Although discovered in 1908, its scientific importance came into light in the late 1960s when it attracted scientists by the recurrent red colouration it caused by forming massive blooms, that cause red tides in the oceans.

<span class="mw-page-title-main">Ocelloid</span>

An ocelloid is a subcellular structure found in the family Warnowiaceae (warnowiids), which are members of a group of unicellular organisms known as dinoflagellates. The ocelloid is analogous in structure and function to the eyes of multicellular organisms, which focus, process and detect light. The ocelloid is much more complex than the eyespot, a light-sensitive structure also found in unicellular organisms, and is in fact one of the most complex known subcellular structures. It has been described as a striking example of convergent evolution.

<span class="mw-page-title-main">Nematocyst (dinoflagellate)</span>

A nematocyst is a subcellular structure or organelle containing extrusive filaments found in two families of athecate dinoflagellates, the Warnowiaceae and Polykrikaceae. It is distinct from the similar subcellular structures found in the cnidocyte cells of cnidarians, a group of multicellular organisms including jellyfish and corals; such structures are also often called nematocysts, and cnidocytes are sometimes referred to as nematocytes. It is unclear whether the relationship between dinoflagellate and cnidarian nematocysts is a case of convergent evolution or common descent, although molecular evidence has been interpreted as supporting an endosymbiotic origin for cnidarian nematocysts.

<span class="mw-page-title-main">Piston (subcellular structure)</span>

A piston is a complex contractile organelle found in some dinoflagellates, namely the Erythropsidinium and Greuetodinium genera of the family Warnowiaceae. This group is also well known for possessing other unusually complex subcellular structures such as the ocelloid and nematocyst. Observations of Erythropsidinium samples reveal that the length of the piston is highly variable across specimens. The piston is known to be capable of repetitive and dramatic contractile motion; although its function is unknown, roles in locomotion, prey capture, and defense have been suggested.

<span class="mw-page-title-main">Polykrikaceae</span> Family of single-celled organisms

The Polykrikaceae are a family of athecate dinoflagellates of the order Gymnodiniales. Members of the family are known as polykrikoids. The family contains two genera: Polykrikos and Pheopolykrikos.

<i>Erythropsidinium</i> Genus of single-celled organisms

Erythropsidinium is a genus of dinoflagellates of the family Warnowiaceae.

<i>Polykrikos</i> Genus of single-celled organisms

Polykrikos is one of the genera of family Polykrikaceae that includes athecate pseudocolony-forming dinoflagellates. Polykrikos are characterized by a sophisticated ballistic apparatus, named the nematocyst-taeniocyst complex, which allows species to prey on a variety of organisms. Polykrikos have been found to regulate algal blooms as they feed on toxic dinoflagellates. However, there is also some data available on Polykrikos being toxic to fish.

Durinskia is a genus of dinoflagellate that can be found in freshwater and marine environments. This genus was created to accommodate its type species, Durinskia baltica, after major classification discrepancies were found. While Durinskia species appear to be typical dinoflagellates that are armored with cellulose plates called theca, the presence of a pennate diatom-derived tertiary endosymbiont is their most defining characteristic. This genus is significant to the study of endosymbiotic events and organelle integration since structures and organelle genomes in the tertiary plastids are not reduced. Like some dinoflagellates, species in Durinskia may cause blooms.

<span class="mw-page-title-main">Symbiosome</span>

A symbiosome is a specialised compartment in a host cell that houses an endosymbiont in a symbiotic relationship.

<span class="mw-page-title-main">Marine protists</span> Protists that live in saltwater or brackish water

Marine protists are defined by their habitat as protists that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Life originated as marine single-celled prokaryotes and later evolved into more complex eukaryotes. Eukaryotes are the more developed life forms known as plants, animals, fungi and protists. Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are mostly single-celled and microscopic. The term protist came into use historically as a term of convenience for eukaryotes that cannot be strictly classified as plants, animals or fungi. They are not a part of modern cladistics because they are paraphyletic.

Warnowia is a genus of athecate dinoflagellates, characterized by having a very sophisticated photoreceptor organelle called the ocelloid. This genus is dispersed worldwide but is scarce and difficult to find and nearly impossible to culture. As a result, the history and taxonomy of this genus are confusing at best, and many basic characteristics like its life cycle are still unknown. Still, Warnowia has drawn scientific interest as a unicellular organism with a fascinatingly complex photoreceptor system.

References

  1. 1 2 3 4 5 Hoppenrath, M; Bachvaroff, TR; Handy, SM; Delwiche, CF; Leander, BS (25 May 2009). "Molecular phylogeny of ocelloid-bearing dinoflagellates (Warnowiaceae) as inferred from SSU and LSU rDNA sequences". BMC Evolutionary Biology. 9 (1): 116. Bibcode:2009BMCEE...9..116H. doi: 10.1186/1471-2148-9-116 . PMC   2694157 . PMID   19467154.
  2. 1 2 3 Gregory S. Gavelis; Shiho Hayakawa; Richard A. White III; Takashi Gojobori; Curtis A. Suttle; Patrick J. Keeling; Brian S. Leander (2015). "Eye-like ocelloids are built from different endosymbiotically acquired components". Nature. 523 (7559): 204–7. Bibcode:2015Natur.523..204G. doi:10.1038/nature14593. hdl: 10754/566109 . PMID   26131935. S2CID   4462376.
  3. Guiry, M.D.; Guiry, G.M. (2021). "Warnowiaceae". AlgaeBase. National University of Ireland, Galway. Retrieved 15 January 2021.
  4. GÓMEZ, FERNANDO; LÓPEZ-GARCÍA, PURIFICACIÓN; MOREIRA, DAVID (September 2009). "Molecular Phylogeny of the Ocelloid-Bearing Dinoflagellates and (Warnowiaceae, Dinophyceae)". Journal of Eukaryotic Microbiology. 56 (5): 440–445. doi:10.1111/j.1550-7408.2009.00420.x. PMID   19737196. S2CID   41132911.
  5. Hayakawa, Shiho; Takaku, Yasuharu; Hwang, Jung Shan; Horiguchi, Takeo; Suga, Hiroshi; Gehring, Walter; Ikeo, Kazuho; Gojobori, Takashi (2015-03-03). "Function and Evolutionary Origin of Unicellular Camera-Type Eye Structure". PLOS ONE. 10 (3): e0118415. Bibcode:2015PLoSO..1018415H. doi: 10.1371/journal.pone.0118415 . ISSN   1932-6203. PMC   4348419 . PMID   25734540.