are a basal
In phylogenetics, a basal clade is the earliest clade to branch in a larger clade; it appears at the base of a cladogram.A basal group forms an outgroup to the rest of the clade, such as in the following example:...
form of invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...
. They are the simplest in structure of all non-parasitic multicellular animals (Metazoa). They are generally classified as a single species, Trichoplax adhaerens
, although there is enough genetic diversity that it is likely that there are multiple, morphologically similar species. Although they were first discovered in 1883, a common name does not yet exist for the taxon; the scientific name literally means "flat animals".
is a small, flattened, animal around 1 millimetre across. Like an Amoeba
, 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
Epithelium is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. Epithelial tissues line the cavities and surfaces of structures throughout the body, and also form many glands. Functions of epithelial cells include secretion, selective...
enclosing a loose sheet of stellate cells resembling the mesenchyme
Mesenchyme, or mesenchymal connective tissue, is a type of undifferentiated loose connective tissue that is derived mostly from mesoderm, although some are derived from other germ layers; e.g. some mesenchyme is derived from neural crest cells and thus originates from the ectoderm...
of some more advanced animals. The epithelial cells bear flagella
A flagellum is a tail-like projection that protrudes from the cell body of certain prokaryotic and eukaryotic cells, and plays the dual role of locomotion and sense organ, being sensitive to chemicals and temperatures outside the cell. There are some notable differences between prokaryotic and...
, 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. It reproduces asexually, budding off smaller individuals, and the lower surface may also bud off eggs into the mesenchyme.
There is no convincing fossil record of the placozoa, although the Ediacaran
The Ediacaran Period , named after the Ediacara Hills of South Australia, is the last geological period of the Neoproterozoic Era and of the Proterozoic Eon, immediately preceding the Cambrian Period, the first period of the Paleozoic Era and of the Phanerozoic Eon...
biota (Precambrian) organism Dickinsonia
Dickinsonia is an iconic fossil of the Ediacaran biota. It resembles a bilaterally symmetrical ribbed oval. Its affinities are presently unknown; most interpretations consider it to be an animal, although others suggest it may be fungal, or a member of an "extinct kingdom".-Species variety:A...
appears to be closely allied with this phylum.
Traditionally, classification has been based on their level of organization: i.e. they possess no tissues or organs. However this may be as a result of secondary loss, so is inadequate to demark a clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
. 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
Eumetazoa is a clade comprising all major animal groups except sponges, placozoa and several other little known animals. Characteristics of eumetazoans include true tissues organized into germ layers, and an embryo that goes through a gastrula stage...
. In such a feature-poor phylum, molecular data are considered to provide the most reliable approximation of the placozoans' phylogeny.
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) and are thus considered a sister taxon to all other metazoans:
According to a functional-morphology model, all or most animals are descended from a gallertoid
, a free-living (pelagic) sphere in seawater, consisting of a single ciliated layer of cells supported by a thin, noncellular separating layer, the basal lamina
The basal lamina is a layer of extracellular matrix secreted by the epithelial cells, on which the epithelium sits. It is often confused with the basement membrane, and sometimes used inconsistently in the literature, see below....
. The interior of the sphere is filled with contractile fibrous cells and a gelatinous extracellular matrix
In biology, the extracellular matrix is the extracellular part of animal tissue that usually provides structural support to the animal cells in addition to performing various other important functions. The extracellular matrix is the defining feature of connective tissue in animals.Extracellular...
. Both the modern Placozoa and all other animals then descended from this multicellular beginning stage via two different processes:
- Infolding of the epithelium led to the formation of an internal system of ducts and thus to the development of a modified gallertoid from which the sponges (Porifera), Cnidaria
Cnidaria is a phylum containing over 9,000 species of animals found exclusively in aquatic and mostly marine environments. Their distinguishing feature is cnidocytes, specialized cells that they use mainly for capturing prey. Their bodies consist of mesoglea, a non-living jelly-like substance,...
and Ctenophora subsequently 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). 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 sides facing toward and away from the substrate, and between their orientation and the vertical direction perpendicular to the substrate. This results naturally in a selective advantage for flattening of the body, as of course can be seen in many benthic species. In the proposed functional-morphology model, the Placozoa, and possibly also several organisms known only from the fossil state, are descended from such a life form, which is now termed placuloid. Three different life strategies have accordingly led to three different 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 Ediacaran Period in geology. These are usually attributed to bilaterally symmetrical worms, 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 algae as photosynthetically active endosymbionts, i.e. primarily obtaining their nutrients from their partners in symbiosis, were accordingly responsible for the mysterious creatures of the Ediacara fauna that are not assigned to any modern animal taxon and lived during the Ediacaran Period, before the start of the Paleozoic. Recent work has shown that some of the Ediacaran assemblages (e.g. Mistaken Point) were in deep water, below the photic zone, and that the organisms were not dependent on endosymbiotic photosynthesisers.
- Animals that grazed on algal mats were ultimately the direct ancestors of the Placozoa. The advantages of an amoeboid multiplicity of shapes thus allowed a previously present basal lamina and a gelatinous extracellular matrix to be lost secondarily. Pronounced differentiation between the ventral surface facing the substrate and the dorsal, facing away from it, 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 the analysis presented above turn out to be correct, Trichoplax adhaerens
would be the oldest branch of the multicellular animals and a relic of the Ediacara fauna, or even the pre-Ediacara fauna. Due to the absence of extracellular matrix and basal lamina, the development potential of these animals, very successful in their ecological niche, was of course limited, which would explain the low rate of evolution, referred to as bradytely
, of their phenotype, their outward form as adults.
This hypothesis was supported by a recent analysis of the Trichoplax adhaerens
mitochondrial genome in comparison to those of other animals, The hypothesis was, however, rejected in a 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, but only at the p=0.07
level, which indicates a marginal level of statistical significance.
Functional-morphology hypotheses are not undisputed among scientists and are often rejected because of their highly theoretical character, which is not directly accessible to empirical study. Cladistics, a modern form of systematics research, is based exclusively on demonstrable features of living and fossil animal groups (taxa) for reconstructing the genealogy of a species or group.
The most important concept based on purely morphological characteristics pictures the Placozoa as the nearest relative of the animals with true tissues (Eumetazoa). The taxon they share, called the Epitheliozoa, is itself construed to be a sister group to the sponges (Porifera):
The principle support for such a relationship comes from special cell/cell junctions, the belt desmosomes, that occur not just in the Placozoa but in all animals except the sponges; they enable the cells to join together in an unbroken layer like the epitheloid of the Placozoa. Trichoplax adhaerens
also shares the ventral gland cells with most eumetazoans. Both characteristics can be considered apomorphies, i.e. evolutionarily derived features, and thus form the basis of a common taxon for all animals that possess them.
One possible scenario inspired by the proposed hypothesis starts with the idea that the monociliated cells of the epitheloid in Trichoplax adhaerens
evolved by reduction of the collars in the collar cells (choanocytes) of sponges as the ancestors of the Placozoa abandoned a filtering mode of life. The epitheloid would then have served as the precursor to the true epithelial tissue of the eumetazoans.
In contrast to the model based on functional morphology described earlier, in the Epitheliozoa concept 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 Bilateria may have developed from endoderm, whereas ectoderm is, among other things, the precursor to the external skin layer (epidermis). The interior space pervaded by a fiber syncytium in the Placozoa would then correspond to connective tissue in the other animals. It is uncertain whether the calcium ions stored in the syncytium are related to the lime skeletons of many cnidarians.
As noted above, this hypothesis was supported in a 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.
A third hypothesis, based primarily on molecular genetics, views the Placozoa as highly simplified eumetazoans. According to this, Trichoplax adhaerens
is descended from considerably more complex animals that already had muscles and nerve tissues. Both tissue types, as well as the basal lamina of the epithelium, were accordingly lost more recently by radical secondary simplification.
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:
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
All versions of this hypothesis were rejected with high confidence in a 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.