Hoilungia

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Hoilungia
Hoilungia hongkongensis.jpg
Microscopic image of Hoilungia hongkongensis. Scale bar is 0.2 mm.
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Placozoa
Class: Uniplacotomia
Order: Hoilungea
Family: Hoilungidae
Genus: Hoilungia
Eitel, Schierwater & Wörheide, 2018
Species:
H. hongkongensis
Binomial name
Hoilungia hongkongensis
Eitel, Schierwater & Wörheide, 2018

Hoilungia is a genus that contains one of the simplest animals and belongs to the phylum Placozoa. [1] [2] Described in 2018, it has only one named species, H. hongkongensis, although there are possible other species. [3] The animal superficially resembles another placozoan, Trichoplax adhaerens , but genetically distinct from it as mitochondrial DNA analysis revealed. [1]

Contents

Hoilungia was discovered in brackish water from mangrove swamps in Hong Kong. [4] These organisms are generally found in the biofilm surfaces in tropical and subtropical environments. Phylogenetically, they are placed closest to cnidarians. They are diploblastic animals and are believed to have dorso-ventral polarity along top and bottom body layers. Their body is overtly similar to oral-aboral axis of cnidarians. [5]

Discovery

Trichoplax adhaerens was discovered by the German zoologist Franz Eilhard Schulze in 1883. But its identification as to what kind of animal it was (systematic position) was not known. [6] Another German, Karl Gottlieb Grell, discovered the diversity of these animals and created a new phylum Placozoa, in 1971. Grell derived the name from the placula hypothesis, Otto Bütschli's notion on the origin of metazoans. [7]

The advent of molecular techniques allowed genetic analysis of placozoans. The first important report in 2004 by a team of zoologists at the Institute of Animal Ecology & Cell Biology in Hannover, Germany, led by Allen G. Collins and Bernd Schierwater, indicated that placozoans known under T. adhaerens could be genetically many species. [8] Schierwater, teaming up with Michael Eitel and Gert Wörheide from the Ludwig-Maximilians-Universität München, made further studies and found that the specimen H13 was a different placozoan animal for which they introduced the genus Hoilungia, and the species H. hongkongensis, in 2018. [1] [3]

Etymology

The genus name is derived from the phrase "hoi lung", which means "sea dragon" in Cantonese. The species name is after Hong Kong from where it was discovered. [1]

Biology

Structure

Ultrastructure of Hoilungia Hongkongensis. The upper epithelium (blue bar) with monociliated cells (light blue). The intermediate layer (green bar) consists of nonciliated fiber cells (labeled "fc" in light green). The lower epithelium (orange bar) is mostly made up of monociliated cylinder cells (light red). Scale bar is 2 mm. Hoilungia structure.jpg
Ultrastructure of Hoilungia Hongkongensis. The upper epithelium (blue bar) with monociliated cells (light blue). The intermediate layer (green bar) consists of nonciliated fiber cells (labeled “fc” in light green). The lower epithelium (orange bar) is mostly made up of monociliated cylinder cells (light red). Scale bar is 2 μm.

Hoilungia do not have well-defined body plan much like amoebas, unicellular eukaryotes. As Andrew Masterson reported, "[as other placozoans] they are as close as it is possible to get to being simply a little living blob." [9] An individual body measures about 0.55 mm in diameter. [3] There are no body parts; as Eitel described: "There's no mouth, there's no back, no nerve cells, nothing." [1]

As do other placozoans, Hoilungia has only three anatomical parts as tissue layers inside its body: the upper, intermediate (middle) and lower epithelia. There are at least six different cell types. [10] The upper epithelium is the thinnest portion and essentially comprises flat cells with their cell body hanging underneath the surface, and each cell having a cilium. [3] Crystal cells are sparsely distributed near the marginal edge. Few cells have unusually large number of mitochondria. [10] The middle layer is the thickest made up of numerous fiber cells, which contain mitochondrial complexes, vacuoles and endosymbiotic bacteria in the endoplasmic reticulum. The lower epithelium consists of numerous monociliated cylinder cells along with a few endocrine-like gland cells and lipophil cells. Each lipophil cell contains numerous middle-sized granules, one of which is a secretory granule. [3]

The body axes of Hoilungia and Trichoplax are overtly similar to the oral–aboral axis of cnidarians, [5] animals from another phylum with which they are most closely related. [11] Structurally, they can not be distinguished from other placozoans, so that identification is purely on genetic (mitochondrial DNA) differences. [3] Genome sequencing has shown that Hoilungia have 164 unique genes and 9 uniquely missing genes compared to other placozoans. [12]

Nutrition

Hoilungia feed on algae, bacteria, yeast and other byproducts of biofilms. They feed from lower tissue layer which has various peptidergic gland cells. [3]

Reproduction

Hoilungia reproduce asexually through binary fission and budding. They might also reproduce sexually. [3]

Evolutionary history

Hoilungia and Trichoplax are considered one of the earliest branching animal lineages, and have relatively simple morphologies their complexity of NO-cGMP-mediated signaling is greater to those in vertebrates. This evidence has been found in their DNA by experimentation using ultra-sensitive capillary electrophoresis assays. [3] The genomes of H. hongkongensis and other placozoans add support to the phylogenetic placement of the Placozoa as the most ancient (basal) animals in the tree of life. [12]

Related Research Articles

<span class="mw-page-title-main">Cnidaria</span> Aquatic animal phylum having cnydocytes

Cnidaria is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in fresh water and marine environments, including jellyfish, hydroids, sea anemones, corals and some of the smallest marine parasites. Their distinguishing features are a decentralized nervous system distributed throughout a gelatinous body and the presence of cnidocytes or cnidoblasts, specialized cells with ejectable flagella used mainly for envenomation and capturing prey. Their bodies consist of mesoglea, a non-living, jelly-like substance, sandwiched between two layers of epithelium that are mostly one cell thick. Cnidarians are also some of the only animals that can reproduce both sexually and asexually.

<span class="mw-page-title-main">Placozoa</span> Basal form of free-living invertebrate

Placozoa is a phylum of marine and free-living (non-parasitic) animals. They are simple blob-like animals without any body part or organ, and are merely aggregates of cells. Moving in water by ciliary motion, eating food by engulfment, reproducing by fission or budding, placozoans are described as "the simplest animals on Earth." Structural and molecular analyses have supported them as among the most basal animals, thus, constituting a primitive metazoan phylum.

<span class="mw-page-title-main">Mesozoa</span> Subkingdom of worm-like parasites of marine invertebrates

The Mesozoa are minuscule, worm-like parasites of marine invertebrates. Generally, these tiny, elusive creatures consist of a somatoderm of ciliated cells surrounding one or more reproductive cells.

<span class="mw-page-title-main">Ctenophora</span> Phylum of gelatinous marine animals

Ctenophora comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming, and they are the largest animals to swim with the help of cilia.

<span class="mw-page-title-main">Parazoa</span> Ancestral subkingdom of animals

Parazoa are a taxon with sub-kingdom category that is located at the base of the phylogenetic tree of the animal kingdom in opposition to the sub-kingdom Eumetazoa; they group together the most primitive forms, characterized by not having proper tissues or that, in any case, these tissues are only partially differentiated. They generally group a single phylum, Porifera, which lack muscles, nerves and internal organs, which in many cases resembles a cell colony rather than a multicellular organism itself. All other animals are eumetazoans, which do have differentiated tissues.

<span class="mw-page-title-main">Eumetazoa</span> Basal animal clade as a sister group of the Porifera

Eumetazoa, also known as diploblasts, Epitheliozoa or Histozoa, are a proposed basal animal clade as a sister group of Porifera (sponges). The basal eumetazoan clades are the Ctenophora and the ParaHoxozoa. Placozoa is now also seen as a eumetazoan in the ParaHoxozoa. The competing hypothesis is the Myriazoa clade.

<i>Trichoplax</i> Genus of Placozoa

Trichoplax adhaerens is one of the four named species in the phylum Placozoa. The others are Hoilungia hongkongensis, Polyplacotoma mediterranea and Cladtertia collaboinventa. Placozoa is a basal group of multicellular animals, possible relatives of Cnidaria. Trichoplax are very flat organisms commonly less than 4 mm in diameter, lacking any organs or internal structures. They have two cellular layers: the top epitheloid layer is made of ciliated "cover cells" flattened toward the outside of the organism, and the bottom layer is made up of cylinder cells that possess cilia used in locomotion, and gland cells that lack cilia. Between these layers is the fibre syncytium, a liquid-filled cavity strutted open by star-like fibres.

<span class="mw-page-title-main">Coelenterata</span> Term encompassing animal phyla Cnidaria and Ctenophora

Coelenterata is a term encompassing the animal phyla Cnidaria and Ctenophora. The name comes from Ancient Greek κοῖλος (koîlos) 'hollow', and ἔντερον (énteron) 'intestine', referring to the hollow body cavity common to these two phyla. They have very simple tissue organization, with only two layers of cells, along with a middle undifferentiated layer called mesoglea, and radial symmetry. Some examples are corals, which are typically colonial; hydrae, jellyfish, sea anemones, and Aurelia, which are solitary; Pennatula; Portuguese man o' war; Gorgonia; and Physalia. Coelenterata lack a specialized circulatory system, relying instead on diffusion across the tissue layers.

<span class="mw-page-title-main">Medusozoa</span> Clade of marine invertebrates

Medusozoa is a clade in the phylum Cnidaria, and is often considered a subphylum. It includes the classes Hydrozoa, Scyphozoa, Staurozoa and Cubozoa, and possibly the parasitic Polypodiozoa. Medusozoans are distinguished by having a medusa stage in their often complex life cycle, a medusa typically being an umbrella-shaped body with stinging tentacles around the edge. With the exception of some Hydrozoa, all are called jellyfish in their free-swimming medusa phase.

<span class="mw-page-title-main">Animal</span> Kingdom of living things

Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia, With few exceptions, animals consume organic material, breathe oxygen, have myocytes and are able to move, can reproduce sexually, and grow from a hollow sphere of cells, the blastula, during embryonic development. Animals form a clade, meaning that they arose from a single common ancestor.

The Urmetazoan is the hypothetical last common ancestor of all animals, or metazoans. It is universally accepted to be a multicellular heterotroph — with the novelties of a germline and oogamy, an extracellular matrix (ECM) and basement membrane, cell-cell and cell-ECM adhesions and signaling pathways, collagen IV and fibrillar collagen, different cell types, spatial regulation and a complex developmental plan, and relegated unicellular stages.

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

Planulozoa is a clade which includes the Placozoa, Cnidaria and the Bilateria. The designation Planulozoa may be considered a synonym to Parahoxozoa. Within Planulozoa, the Placozoa may be a sister of Cnidaria to the exclusion of Bilateria. The clade excludes basal animals such as the Ctenophora, and Porifera (sponges). Although this clade was sometimes used to specify a clade of Cnidaria and Bilateria to the exclusion of Placozoa, this is no longer favoured due to recent data indicating a sister group relationship between Cnidaria and Placozoa.

<span class="mw-page-title-main">ParaHoxozoa</span> Clade of all animals except sponges and comb jellies

ParaHoxozoa is a clade of animals that consists of Bilateria, Placozoa, and Cnidaria. The relationship of this clade relative to the two other animal lineages Ctenophora and Porifera is debated. Some phylogenomic studies have presented evidence supporting Ctenophora as the sister to Parahoxozoa and Porifera as the sister group to the rest of animals. Other studies have presented evidence supporting Porifera as the sister to Parahoxozoa and Ctenophora as the sister group to the rest of animals, finding that nervous systems either evolved independently in ctenophores and parahoxozoans, or were secondarily lost in poriferans. If ctenophores are taken to have diverged first, Eumetazoa is sometimes used as a synonym for ParaHoxozoa.

Polyplacotoma mediterranea is a species in the phylum Placozoa, only representative of the genus Polyplacotoma. They differ greatly from other species of placozoans with regards to their morphology and genetic makeup, and have been ranked in the separate class Polyplacotomia. P. mediterranea has the smallest mitogenome, the lowest GC content, and the smallest intergenic spacer regions of all placozoans. Their bodily structure consists of elongated polytomous body branches, as well as a maximum size that is greater than 10 mm in length. The mitochondrial genome of Polyplacotoma mediterranea is also very compact and contains overlapping protein and tRNA gene codes.

<span class="mw-page-title-main">Uniplacotomia</span> Class of Placozoa

Uniplacotomia is a class of placozoans encompassing the vast majority of the phylum, with the exception of Polyplacotoma. It was established in 2022. It comprises the orders Trichoplacea, Cladhexea and Hoilungea. Their morphology is consistent across the class, resembling the typical Trichoplax as mostly rounded, flat organisms rather than the polytomous, branching structure exhibited by Polyplacotomia.

<i>Cladtertia</i> Genus of placozoans

Cladtertia is a genus of placozoan discovered in 2022, whose only currently described species is Cladtertia collaboinventa. However, the genus is known to contain several other species, awaiting a formal description. Its closest described relative is Hoilungia hongkongensis, with whom it forms the order Hoilungea. After Trichoplax, Hoilungia and Polyplacotomia, it is the fourth described placozoan genus up to date.

Hoilungea is a recently created placozoan order comprising Cladtertia, Hoilungia, and other yet-undescribed species. Named in 2022, it is believed to be sister to Cladhexea, and corresponds to Clades III, IV, V and VII of the literature.

Cladhexea is a recently created placozoan order comprising yet-undescribed species. Named in 2022, it is believed to be sister to Hoilungea, and corresponds to Clade VI of the literature.

Polyplacotomia is a class of placozoans, to this date only comprising Polyplacotoma mediterranea. It was established in 2022. Their morphology is strikingly different from other placozoans in Uniplacotomia, exhibiting a highly ramified, branching structure with multiple amoeboid projections. It differs from Uniplacotomia by 76 uniquely present and 600 absent genes.

Hoilungidae is a recently created placozoan family comprising Hoilungia and other yet-undescribed species. Named in 2022, it is believed to be sister to Cladtertiidae, and corresponds to Clades IV, V and VII of the literature.

References

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  11. Laumer, Christopher E.; Fernández, Rosa; Lemer, Sarah; Combosch, David; Kocot, Kevin M.; Riesgo, Ana; Andrade, Sónia C. S.; Sterrer, Wolfgang; Sørensen, Martin V.; Giribet, Gonzalo (2019-07-10). "Revisiting metazoan phylogeny with genomic sampling of all phyla". Proceedings. Biological Sciences. 286 (1906): 20190831. doi:10.1098/rspb.2019.0831. ISSN   1471-2954. PMC   6650721 . PMID   31288696.
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