Placozoa

Placozoa ( PLAK-ə-ZOH-ə; lit. 'flat animals') is a phylum of free-living (non-parasitic) marine invertebrates. They are blob-like animals composed of aggregations of cells. Moving in water by ciliary motion, eating food by engulfment, and 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. The first known placozoan, Trichoplax adhaerens, was discovered in 1883 by the German zoologist Franz Eilhard Schulze (1840–1921). Recognizing its distinctive characteristics, the German zoologist Karl Gottlieb Grell (1912–1994) erected a new phylum, Placozoa, for it in 1971. Remaining a monotypic phylum for over a century, new species began to be added in 2018. So far, three other extant species have been described, in two distinct classes: Uniplacotomia (Hoilungia hongkongensis in 2018 and Cladtertia collaboinventa in 2022) and Polyplacotomia (Polyplacotoma mediterranea, the most basal, in 2019). A single putative fossil species is known, the Middle Triassic Maculicorpus microbialis.

History Trichoplax was discovered in 1883 by the German zoologist Franz Eilhard Schulze, in a seawater aquarium at the Zoological Institute in Graz, Austria. The generic name is derived from the classical Greek θρίξ (thrix), meaning "hair", and πλάξ (plax), "plate". The specific epithet adhaerens is Latin meaning "adherent", reflecting its propensity to stick to the glass slides and pipettes used in its examination. Schulze realized that the animal could not be a member of any existing phyla, and based on the simple structure and behaviour, concluded in 1891 that it must be an early metazoan. He also observed the reproduction by fission, cell layers and locomotion. 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. Monticelli did not preserve them and no other specimens were found again, as a result of which the identification is ruled as doubtful, and the species rejected. Schulze's description was opposed by other zoologists. For instance, in 1890, F.C. Noll argued that the animal was a flatworm (Turbellaria). In 1907, Thilo Krumbach published a hypothesis that Trichoplax is not a distinct animal but that it is a form of the planula larva of the anemone-like hydrozoan Eleutheria krohni. Although this was refuted in print by Schulze and others, Krumbach's analysis became the standard textbook explanation, and nothing was printed in zoological journals about Trichoplax until the 1960s. The development of electron microscopy in the mid-20th century allowed in-depth observation of the cellular components of organisms, following which there was renewed interest in Trichoplax starting in 1966. The most detailed descriptions were made by Karl Gottlieb Grell at the University of Tübingen beginning in 1971. That year, Grell revived Schulze's interpretation that the animals are unique and created a new phylum Placozoa. Grell derived the name from the placula hypothesis, Otto Bütschli's notion on the origin of metazoans.

Biology

Anatomy

Placozoans lack a well-defined body plan and have a simple, flattened body. As Andrew Masterson reported: "they are as close as it is possible to get to being simply a little living blob." An individual body measures about 0.55 mm in diameter. There are no body parts; as one of the researchers Michael Eitel described: "There's no mouth, there's no back, no nerve cells, nothing." Animals studied in laboratories have bodies consisting of everything from hundreds to millions of cells. Placozoans have 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. (A 2023 analysis of 4 species across 3 genera found 8, one with an unknown role.) 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. Crystal cells are sparsely distributed near the marginal edge. A few cells have unusually large number of mitochondria. 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. The body axes of Hoilungia and Trichoplax are similar to the oral–aboral axis of cnidarians, animals from another phylum with which they are most closely related. Structurally, they can not be distinguished from other placozoans, so that identification is purely on genetic (mitochondrial DNA) differences. Genome sequencing has shown that each species has a set of unique genes and several uniquely missing genes. Trichoplax is a small, flattened, animal around 1 mm (0.039 in) across. An amorphous multi-celled body, analogous to a single-celled amoebas, it has no regular outline, although the lower surface is somewhat concave, and the upper surface is always flattened. The body consists of an outer layer of simple epithelium enclosing a loose sheet of stellate cells resembling the mesenchyme of some more complex animals. The epithelial cells bear cilia, which the animal uses to help it creep along the seafloor. The lower surface engulfs small particles of organic detritus, on which the animal feeds. Studies suggest that aragonite crystals in crystal cells have the same function as statoliths, allowing it to use gravity for spatial orientation. Located in the dorsal epithelium there are lipid granules which release a cocktail of toxins as a means of defense, and can induce paralysis or death in some predators. Genes encoding for proteins which make up the poisonous secretions of Trichoplax have been found to resemble venom-associated genes present in the genomes of certain snakes, like the American copperhead and the West African carpet viper.

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