Placozoa
Placozoans Temporal range: [1]
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Trichoplax adhaerens
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Scientific classification | |
Domain: | Eukaryota |
(unranked): | Filozoa |
Kingdom: | Animalia |
Phylum: | Placozoa Grell, 1971 |
Type species | |
Trichoplax adhaerens | |
Classes[2] | |
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Placozoa (
The first known placozoan,
History
Trichoplax was discovered in 1883 by the German zoologist
In 1893, Italian zoologist Francesco Saverio Monticelli described another animal which he named Treptoplax, the specimens of which he collected from Naples. He gave the species name T. reptans in 1896.[19] Monticelli did not preserve them and no other specimens were found again, as a result of which the identification is ruled as doubful, and the species rejected.[20][21]
Schulze's description was opposed by other zoologists. For instance, in 1890, F.C. Noll argued that the animal was a flat worm (Turbellaria).
The development of
Biology
Placozoans do not have well-defined body plan much like
Placozoans have only three anatomical parts as tissue layers inside its body: the upper, intermediate (middle) and lower
The body axes of Hoilungia and Trichoplax are overtly similar to the oral–aboral axis of
Trichoplax is a small, flattened, animal around 1 mm (0.039 in) across. An amorphous multi-celled body, analogous to a single-celled
The lower surface engulfs small particles of organic detritus, on which the animal feeds. All placozoa can reproduce asexually, budding off smaller individuals, and the lower surface may also bud off eggs into the mesenchyme.[11] Sexual reproduction has been reported to occur in one clade of placozoans,[38][39] whose strain H8 was later found to belong to genus Cladtertia,[2] where intergenic recombination was observed as well as other hallmarks of sexual reproduction.
Some Trichoplax species contain Rickettsiales bacteria as endosymbionts.[40]
One of the at least 20 described species turned out to have two bacterial endosymbionts;
Studies suggests that aragonite crystals in crystal cells have the same function as statoliths, allowing it to use gravity for spatial orientation.[43]
Located in the dorsal epithelium there are lipid granules called shiny spheres which release a cocktail of venoms and toxins as an anti-predator defense, and can induce paralysis or death in some predators. Genes has been found in Trichoplax with a strong resemblance to the venom genes of some poisonous snakes, like the American copperhead and the West African carpet viper.[44][45]
The Placozoa show substantial evolutionary radiation in regard to sodium channels, of which they have 5–7 different types, more than any other invertebrate species studied to date.[47]
Three modes of population dynamics depended upon feeding sources, including induction of social behaviors, morphogenesis, and reproductive strategies. [48]
In addition to fission, representatives of all species produced “swarmers” (a separate vegetative reproduction stage), which could also be formed from the lower epithelium with greater cell-type diversity.[49]
Evolutionary relationships
There is no convincing fossil record of the placozoa, although the Ediacaran biota (Precambrian, 550 million years ago) organism Dickinsonia appears somewhat similar to placozoans.[50] Knaust (2021) reported preservation of placozoan fossils in a microbialite bed from the Middle Triassic Muschelkalk (Germany).[1]
Traditionally, classification was based on their level of organization, i.e., they possess no tissues or organs. However this may be as a result of secondary loss and thus is inadequate to exclude them from relationships with more complex animals. More recent work has attempted to classify them based on the DNA sequences in their genome; this has placed the phylum between the sponges and the eumetazoa.[51] In such a feature-poor phylum, molecular data are considered to provide the most reliable approximation of the placozoans' phylogeny.
Their exact position on the phylogenetic tree would give important information about the origin of neurons and muscles. If the absence of these features is an original trait of the Placozoa, it would mean that a nervous system and muscles evolved three times should placozoans and cnidarians be a sister group; once in the Ctenophora, once in the Cnidaria and once in the Bilateria. If they branched off before the Cnidaria and Bilateria split, the neurons and muscles would have the same origin in the two latter groups.
Functional-morphology hypothesis
On the basis of their simple structure, the Placozoa were frequently viewed as a model organism for the transition from unicellular organisms to the multicellular animals (
Metazoa |
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According to a functional-morphology model, all or most animals are descended from a
- Infolding of the Porifera), Cnidaria and Ctenophorasubsequently developed.
- Other gallertoids, according to this model, made the transition over time to a benthic mode of life; that is, their habitat has shifted from the open ocean to the floor (benthic zone). This results naturally in a selective advantagefor flattening of the body, as of course can be seen in many benthic species.
While the probability of encountering food, potential sexual partners, or predators is the same in all directions for animals floating freely in the water, there is a clear difference on the seafloor between the functions useful on body sides facing toward and away from the substrate, leading their sensory, defensive, and food-gathering cells to differentiate and orient according to the vertical – the direction perpendicular to the substrate. In the proposed functional-morphology model, the Placozoa, and possibly several similar organisms only known from the fossils, are descended from such a life form, which is now termed placuloid.
Three different life strategies have accordingly led to three different possible lines of development:
- Animals that live interstitially in the sand of the ocean floor were responsible for the fossil crawling traces that are considered the earliest evidence of animals; and are detectable even prior to the dawn of the bilaterally symmetricalworms, but the hypothesis presented here views animals derived from placuloids, and thus close relatives of Trichoplax adhaerens, to be the producers of the traces.
- Animals that incorporated Mistaken Point) were in deep water, below the photic zone, and hence those individuals could not dependent on endosymbiotic photosynthesisers.
- Animals that grazed on dorsal) accordingly led to the physiologically distinct cell layers of Trichoplax adhaerens that can still be seen today. Consequently, these are analogous, but not homologous, to ectoderm and endoderm– the "external" and "internal" cell layers in eumetazoans – i.e. the structures corresponding functionally to one another have, according to the proposed hypothesis, no common evolutionary origin.
Should any of the analyses presented above turn out to be correct, Trichoplax adhaerens would be the oldest branch of the multicellular animals, and a relic of the
This hypothesis was supported by a recent analysis of the Trichoplax adhaerens
Epitheliozoa hypothesis
A concept based on purely morphological characteristics pictures the Placozoa as the nearest relative of the animals with true tissues (
The above view could be correct, although there is some evidence that the
This is now a disputed classification.[56] Placozoans are estimated to have emerged 750–800 million years ago, and the first modern neuron to have originated in the common ancestor of cnidarians and bilaterians about 650 million years ago (many of the genes expressed in modern neurons are absent in ctenopheres, although some of these missing genes are present in placozoans).[57][58]The principal support for such a relationship comes from special cell to cell junctions – belt
One possible scenario inspired by the proposed hypothesis starts with the idea that the monociliated cells of the epitheloid in
In contrast to the model based on functional morphology described earlier, in the Epitheliozoa hypothesis, the ventral and dorsal cell layers of the Placozoa are homologs of endoderm and ectoderm — the two basic embryonic cell layers of the eumetazoans. The digestive gastrodermis in the Cnidaria or the gut epithelium in the bilaterally symmetrical animals (Bilateria) may have developed from endoderm, whereas ectoderm is the precursor to the external skin layer (epidermis), among other things. The interior space pervaded by a fiber syncytium in the Placozoa would then correspond to connective tissue in the other animals. It is unclear whether the calcium ions stored in the syncytium would be related to the lime skeletons of many cnidarians.[citation needed]
As noted above, this hypothesis was supported in a statistical analysis of the Trichoplax adhaerens whole genome sequence, as compared to the whole-genome sequences of six other animals and two related non-animal species.[51]
Eumetazoa hypothesis
A third hypothesis, based primarily on molecular genetics, views the Placozoa as highly simplified
Various studies in this regard so far yield differing results for identifying the exact sister group: in one case the Placozoa would qualify as the nearest relatives of the Cnidaria, while in another they would be a sister group to the Ctenophora, and occasionally they are placed directly next to the Bilateria. Currently, they are typically placed according to the cladogram below:[60]
Metazoa
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In this cladogram the
An argument raised against the proposed scenario is that it leaves morphological features of the animals completely out of consideration. The extreme degree of simplification that would have to be postulated for the Placozoa in this model, moreover, is known only for parasitic organisms but would be difficult to explain functionally in a free-living species like Trichoplax adhaerens.[citation needed]
This version is supported by statistical analysis of the Trichoplax adhaerens whole genome sequence in comparison to the whole genome sequences of six other animals and two related non-animal species. However, ctenophora was not included in the analyses, placing the placozoas outside of the sampled Eumetazoans.[51][61]
Cnidaria-sister hypothesis
DNA comparisons suggest that placozoans are related to Cnidaria, derived from planula larva (as seen in some Cnidaria).[62] The Bilateria also are thought to be derived from planuloids.[63][64][65][66][67][68][69][70] The Cnidaria and Placozoa body axis are overtly similar, and Placozoan and Cnidarian cells are responsive to the same neuropeptide antibodies despite extant Placozoa's not developing any neurons.[71][72]
Choanozoa |
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950 mya |
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External links
- The Trichoplax adhaerens Grell-BS-1999 v1.0 Genome Portal at the DOE Joint Genome Institute
- The Trichoplax Genome Project at the Yale Peabody Museum
- A Weird Wee Beastie: Trichoplax adhaerens
- Research articles from the ITZ, TiHo Hannover
- Information page from the University of California at Berkeley
- Ender A, Schierwater B (January 2003). "Placozoa are not derived cnidarians: evidence from molecular morphology". Mol. Biol. Evol. 20 (1): 130–4. analysis and phylogeny of Trichoplax adhaerens
- Historical overview of Trichoplax research
- Science Daily:Genome Of Simplest Animal Reveals Ancient Lineage, Confounding Array Of Complex Capabilities
- Vicki Buchsbaum Pearse, and Oliver Voigt, 2007. "Field biology of placozoans (Trichoplax): distribution, diversity, biotic interactions. Integrative and Comparative Biology", .