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 , possible relatives of Cnidaria. Trichoplax are very flat organisms commonly less than 4 mm in diameter,
Trichoplax feed by absorbing food particles—mainly —with their underside. They generally reproduce asexually, by Cell division or budding, but can also reproduce sexually. Though Trichoplax has a small genome in comparison to other animals, nearly 87% of its 11,514 predicted protein-coding genes are identifiably similar to known genes in other animals.
Although from the very beginning most researchers who studied Trichoplax in any detail realized that it had no close relationship to other animal phyla, the zoologist Thilo Krumbach published a hypothesis that Trichoplax is a form of the planula larva of the sea anemone-like hydrozoan Eleutheria krohni in 1907. 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. In the 1960s and 1970s a new interest among researchers led to acceptance of Placozoa as a new animal phylum. Among the new discoveries was study of the early phases of the animals' embryonic development and evidence that the animals that people had been studying are adults, not larvae. This newfound interest also included study of the organism in nature (as opposed to aquariums).
Trichoplax lacks tissues and organs; there is also no manifest body symmetry, so it is not possible to distinguish anterior from posterior or left from right. It is made up of a few thousand cells of six types in three distinct layers: dorsal epithelia cells and ventral epithelia cells, each with a single cilium ("monociliate"), ventral gland cells, syncytial fiber cells, lipophils, and crystal cells (each containing a birefringent crystal, arrayed around the rim). Lacking sensory and muscle cells, it moves using Cilium on its external surface. The collective movements of the cilia are completely coordinated by mechanical interactions.
A mature individual consists of up to a thousand cells that can be divided into four different cell types. The monociliated cells of the dorsal epitheloid are flattened and contain lipid bodies. The cells on the ventral side likewise possess a single cilium, while their elongated columnar shape, with a small cross section at the surface, packs them very closely together, causing the cilia to be very closely spaced on the ventral side and to form a ciliated "crawling sole". Interspersed among these ventral epithlioid cells are unciliated gland cells thought to be capable of synthesizing .
On both sides of the septa are liquid-filled capsules that cause the septa to resemble , i.e. nerve-cell junctions that occur in fully expressed form only in animals with tissues (Eumetazoa). Striking accumulations of calcium ions, which may have a function related to the propagation of stimuli, likewise suggest a possible role as protosynapses. This view is supported by the fact that fluorescent antibodies against cnidarian neurotransmitters, i.e. precisely those signal carriers that are transferred in synapses, bind in high concentrations in certain cells of Trichoplax adhaerens, and thus indicate the existence of comparable substances in the Placozoa. The fibre syncytium also contains molecules of actin and probably also of myosin, which occur in the muscle cells of eu . In the placozoans, they ensure that the individual fibres can relax or contract and thus help determine the animals' shape.
In this way, the fibre syncytium assumes the functions of nerve and muscle tissues. Moreover, at least a portion of digestion occurs here. On the other hand, no gelatinous extracellular matrix exists of the kind observed, in mesoglea, in and .
Pluripotent cells, which can differentiate into other cell types, have not yet been demonstrated unambiguously in T. adhaerens, in contrast to the case of the Eumetazoa. The conventional view is that dorsal and ventral epithelioid cells arise only from other cells of the same type.
All nuclei of placozoan cells contain six pairs of that are only about two to three micrometres in size. Three pairs are metacentric, meaning that the centromere, the attachment point for the spindle fibers in cell division, is located at the center, or acrocentric, with the centromere at an extreme end of each chromosome. The cells of the fiber syncytium can be tetraploid, i.e. contain a quadruple complement of chromosomes.
A single complement of chromosomes in Trichoplax adhaerens contains a total of fewer than fifty million base pairs and thus forms the smallest animal genome; the number of base pairs in the intestinal bacterium Escherichia coli is smaller by a factor of only ten.
The genetic complement of Trichoplax adhaerens has not yet been very well researched; it has, however, already been possible to identify several genes, such as Brachyury and TBX2/TBX3, which are homologous to corresponding base-pair sequences in eumetazoans. Of particular significance is Trox-2, a placozoan gene known under the name Cnox-2 in cnidarians and as Gsx in the bilaterally symmetrical Bilateria. As a homeobox or Hox gene it plays a role in organization and differentiation along the axis of symmetry in the embryonic development of eumetazoans; in cnidarians, it appears to determine the position of mouth-facing (oral) and opposite-facing (aboral) sides of the organism. Since placozoans possess no axes of symmetry, exactly where the gene is transcribed in the body of Trichoplax is of special interest. Antibody studies have been able to show that the gene's product occurs only in the transition zones of the dorsal and ventral sides, perhaps in a fifth cell type that has not yet been characterized. It is not yet clear whether these cells, contrary to traditional views, are , which play a role in cell differentiation. In any case, Trox-2 can be considered a possible candidate for a proto-Hox gene, from which the other genes in this important family could have arisen through gene duplication and variation.
Initially, molecular-biology methods were applied unsuccessfully to test the various theories regarding Placozoa's position in the Metazoa system. No clarification was achieved with standard markers such as 18S rDNA/RNA: the marker sequence was apparently "garbled", i.e. rendered uninformative as the result of many mutations. Nevertheless, this negative result supported the suspicion that Trichoplax might represent an extremely primitive lineage of metazoans, since a very long period of time had to be assumed for the accumulation of so many mutations.
Of the 11,514 genes identified in the six chromosomes of Trichoplax, 87% are identifiably similar to genes in cnidarians and bilaterians. In those Trichoplax genes for which equivalent genes can be identified in the human genome, over 80% of the introns (the regions within genes that are removed from RNA molecules before their sequences are translated in protein synthesis) are found in the same location as in the corresponding human genes. The arrangement of genes in groups on chromosomes is also conserved between the Trichoplax and human genomes. This contrasts to other model systems such as fruit flies and soil nematodes that have experienced a paring down of non-coding regions and a loss of the ancestral genome organizations.
Field specimens tend to be found in the coastal of tropical and subtropical seas, on such substrates as the trunks and roots of mangroves, shells of molluscs, fragments of stony corals or simply on pieces of rock. One study was able to detect seasonal population fluctuations, the causes of which have not yet been deduced.
Entire single-celled organisms can also be ingested through the upper epitheloid (that is, the "dorsal surface" of the animal). This mode of feeding could be unique in the animal kingdom: the particles, collected in a slime layer, are drawn through the intercellular gaps (cellular interstices) of the epitheloid by the fibre cells and then digested by phagocytosis ("cell-eating"). Such "collecting" of nutrient particles through an intact tegument is only possible because some "insulating" elements (specifically, a basal lamina under the epitheloid and certain types of cell-cell junctions) are not present in the Placozoa.
When the concentrations of algae are high the animals are more likely to engage in social feeding behavior.
Not all bacteria in the interior of Placozoa are digested as food: in the endoplasmic reticulum, an organelle of the fibre syncytium, bacteria are frequently found that appear to live in symbiosis with Trichoplax adhaerens. These , which are no longer able to survive outside its host, are transferred from one generation to the next through both vegetative and sexual reproduction.
It has been possible to demonstrate a close connection between body shape and the speed of locomotion, which is also a function of available food:
Since the transition is not smooth but happens abruptly, the two modes of extension can be very clearly separated from one another. The following is a qualitative explanation of the animal's behavior:
The actual direction in which Trichoplax moves each time is random: if we measure how fast an individual animal moves away from an arbitrary starting point, we find a linear relationship between elapsed time and mean square distance between starting point and present location. Such a relationship is also characteristic of random Brownian motion of molecules, which thus can serve as a model for locomotion in the Placozoa.
Small animals are also capable of swimming actively with the aid of their cilia. As soon as they come into contact with a possible substrate, a dorsoventral response occurs: the dorsal cilia continue to beat, whereas the cilia of ventral cells stop their rhythmic beating. At the same time, the ventral surface tries to make contact with the substrate; small protrusions and invaginations, the microvilli found on the surface of the columnar cells, help in attaching to the substrate via their adhesive action.
Using T. adhaerens as a model, 0.02–0.002 Hz oscillations in locomotory and feeding patterns were observed, and taken as evidence of complex multicellular integration, dependent on endogenous secretion of signal molecules. Evolutionarily conserved low-molecular-weight transmitters (glutamate, aspartate, glycine, GABA, and ATP) acted as coordinators of distinct locomotory and feeding patterns. Specifically, L-glutamate induced and partially mimicked endogenous feeding cycles, whereas glycine and GABA suppressed feeding. ATP-modified feeding is complex, first causing feeding-like cycles and then suppressing feeding. Trichoplax locomotion was modulated by glycine, GABA, and, surprisingly, by animals’ own mucus trails. Mucus triples locomotory speed compared to clean substrates. Glycine and GABA increased the frequency of turns.
Sexual reproduction is thought to be triggered by excessive population density. As a result, the animals absorb liquid, begin to swell, and separate from the substrate so that they float freely in the water. In the protected interior space, the ventral cells form an ovum surrounded by a special envelope, the fertilisation membrane; the ovum is supplied with nutrients by the surrounding syncytium, allowing energy-rich yolk to accumulate in its interior. Once maturation of the ovum is complete, the rest of the animal degenerates, liberating the ovum itself. Small, unciliated cells that form at the same time are interpreted to be spermatozoa. It has not yet been possible to observe fertilisation itself; the existence of the fertilisation membrane is currently taken to be evidence, however, that it has taken place.
Putative eggs have been observed, but they degrade, typically at the 32–64 cell stage. Neither embryonic development nor sperm have been observed. Despite lack of observation of sexual reproduction in the lab, the genetic structure of the populations in the wild is compatible with the sexual reproduction mode, at least for species of the analysed genotype H5.
Usually even before its liberation, the ovum initiates cleavage processes in which it becomes completely pinched through at the middle. A ball of cells characteristic of animals, the blastula, is ultimately produced in this manner, with a maximum of 256 cells. Development beyond this 256-cell stage has not yet been observed.
Trichoplax lack a homologue of the Boule protein that appears to be ubiquitous and conserved in males of all species of other animals tested. If its absence implies the species has no males, then perhaps its "sexual" reproduction may be a case of the above-described process of regeneration, combining cells from two separate organisms into one.
Due to the possibility of its cloning itself by asexual propagation without limit, the life span of Placozoa is infinite; in the laboratory, several lines descended from a single organism have been maintained in culture for an average of 20 years without the occurrence of sexual processes.
Its ability to fight cancer through a combination of aggressive DNA repair and ejection of damaged cells makes it a promising organism for cancer research.
In addition to basic research, this animal could also be suitable for studying wound-healing and regeneration processes; as yet unidentified metabolic products should be researched. Finally, Trichoplax adhaerens is also being considered as an animal model for testing compounds and antibacterial drugs.
The related lineage Trichoplax sp. H2 has been suggested to be a more suitable model organism than T. adhaerens, due to its abundance and ease of culture.
Significant genetic differences have been observed between collected specimens matching the morphological description of T. adhaerens, leading scientists to suggest in 2004 that it may be a cryptic species complex. At least twenty haplotypes have since been assigned based on the 16S mitochondrial DNA fragment, with T. adhaerens being equated to the lineage H1.
While most haplotypes have not been formally described as species, they have been (with the exception of the morphologically distinct H0, Polyplacotoma mediterranea) provisionally placed into seven distinct clades. The genus Trichoplax was redefined in 2021 as comprising clades I and II, including haplotypes H1, H2, H3 and H17. A 2022 study defined Trichoplax more restrictively as only clade I (haplotypes H1, H2 and H17), with H3 being suggested to belong to a separate undescribed genus in the family Trichoplacidae.
Trichoplax sp. H2 also show differences in cell differentiation, having 29 cell types compared to H1's 28 when analyzed using graph-based 2D projections and gene expression patterns. It has a population of "concave-disk" epithelia upper-like cells unique to this haplotype, first described in 2023. It also has an additional "unknown" cell type characterized by β-secretase expression; this type is found in other Placozoans besides H1. Other relatively minor differences make up for the difference in number.
H17 has been imaged using mmunofluorescence imaging.
Role as a model organism
Systematics
Haplotype H2
Haplotype H17
Comparative genomics
Further reading
Scientific literature
First descriptions
External links
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