Upregulation of DNA repair genes and cell extrusion underpin the remarkable radiation resistance of Trichoplax adhaerens.

Trichoplax adhaerens is the simplest multicellular animal with tissue differentiation and somatic cell turnover. Like all other multicellular organisms, it should be vulnerable to cancer, yet there have been no reports of cancer in T. adhaerens or any other placozoan. We investigated the cancer resistance of T. adhaerens, discovering that they are able to tolerate high levels of radiation damage (218.6 Gy). To investigate how T. adhaerens survive levels of radiation that are lethal to other animals, we examined gene expression after the X-ray exposure, finding overexpression of genes involved in DNA repair and apoptosis including the MDM2 gene. We also discovered that T. adhaerens extrudes clusters of inviable cells after X-ray exposure. T. adhaerens is a valuable model organism for studying the molecular, genetic, and tissue-level mechanisms underlying cancer suppression.

Placozoa.

Schierwater & DeSalle introduce the enigmatic phylum Placozoa.

Getting Nervous: An Evolutionary Overhaul for Communication.

The evolution of a nervous system as a control system of the body's functions is a key innovation of animals. Its fundamental units are neurons, highly specialized cells dedicated to fast cell-cell communication. Neurons pass signals to other neurons, muscle cells, or gland cells at specialized junctions, the synapses, where transmitters are released from vesicles in a Ca2+-dependent fashion to activate receptors in the membrane of the target cell. Reconstructing the origins of neuronal communication out of a more simple process remains a central challenge in biology. Recent genomic comparisons have revealed that all animals, including the nerveless poriferans and placozoans, share a basic set of genes for neuronal communication. This suggests that the first animal, the Urmetazoan, was already endowed with neurosecretory cells that probably started to connect into neuronal networks soon afterward. Here, we discuss scenarios for this pivotal transition in animal evolution.

Dendrogramma, new genus, with two new non-bilaterian species from the marine bathyal of southeastern Australia (Animalia, Metazoa incertae sedis)--with similarities to some medusoids from the Precambrian Ediacara.

A new genus, Dendrogramma, with two new species of multicellular, non-bilaterian, mesogleal animals with some bilateral aspects, D. enigmatica and D. discoides, are described from the south-east Australian bathyal (400 and 1000 metres depth). A new family, Dendrogrammatidae, is established for Dendrogramma. These mushroom-shaped organisms cannot be referred to either of the two phyla Ctenophora or Cnidaria at present, because they lack any specialised characters of these taxa. Resolving the phylogenetic position of Dendrogramma depends much on how the basal metazoan lineages (Ctenophora, Porifera, Placozoa, Cnidaria, and Bilateria) are related to each other, a question still under debate. At least Dendrogramma must have branched off before Bilateria and is possibly related to Ctenophora and/or Cnidaria. Dendrogramma, therefore, is referred to Metazoa incertae sedis. The specimens were fixed in neutral formaldehyde and stored in 80% ethanol and are not suitable for molecular analysis. We recommend, therefore, that attempts be made to secure new material for further study. Finally similarities between Dendrogramma and a group of Ediacaran (Vendian) medusoids are discussed.

Animal evolution: looking for the first nervous system.

The human brain is easily the most baffling bit of biology on the planet. How did the nervous system evolve? What came first: neurons or synaptic proteins? A new paper studying the pancake-shaped Trichoplax suggests it was not the neurons.

Novel cell types, neurosecretory cells, and body plan of the early-diverging metazoan Trichoplax adhaerens.

BACKGROUND: Trichoplax adhaerens is the best-known member of the phylum Placozoa, one of the earliest-diverging metazoan phyla. It is a small disk-shaped animal that glides on surfaces in warm oceans to feed on algae. Prior anatomical studies of Trichoplax revealed that it has a simple three-layered organization with four somatic cell types. RESULTS: We reinvestigate the cellular organization of Trichoplax using advanced freezing and microscopy techniques to identify localize and count cells. Six somatic cell types are deployed in stereotyped positions. A thick ventral plate, comprising the majority of the cells, includes ciliated epithelial cells, newly identified lipophil cells packed with large lipid granules, and gland cells. Lipophils project deep into the interior, where they alternate with regularly spaced fiber cells whose branches contact all other cell types, including cells of the dorsal and ventral epithelium. Crystal cells, each containing a birefringent crystal, are arrayed around the rim. Gland cells express several proteins typical of neurosecretory cells, and a subset of them, around the rim, also expresses an FMRFamide-like neuropeptide. CONCLUSIONS: Structural analysis of Trichoplax with significantly improved techniques provides an advance in understanding its cell types and their distributions. We find two previously undetected cell types, lipohil and crystal cells, and an organized body plan in which different cell types are arranged in distinct patterns. The composition of gland cells suggests that they are neurosecretory cells and could control locomotor and feeding behavior.

Global diversity of the Placozoa.

The enigmatic animal phylum Placozoa holds a key position in the metazoan Tree of Life. A simple bauplan makes it appear to be the most basal metazoan known and genetic evidence also points to a position close to the last common metazoan ancestor. Trichoplax adhaerens is the only formally described species in the phylum to date, making the Placozoa the only monotypic phylum in the animal kingdom. However, recent molecular genetic as well as morphological studies have identified a high level of diversity, and hence a potential high level of taxonomic diversity, within this phylum. Different taxa, possibly at different taxonomic levels, are awaiting description. In this review we firstly summarize knowledge on the morphology, phylogenetic position and ecology of the Placozoa. Secondly, we give an overview of placozoan morphological and genetic diversity and finally present an updated distribution of placozoan populations. We conclude that there is great potential and need to erect new taxa and to establish a firm system for this taxonomic tabula rasa.

Chasing the urmetazoon: striking a blow for quality data?

The ever-lingering question: "What did the urmetazoan look like?" has not lost its charm, appeal or elusiveness for one and a half centuries. A solid amount of organismal data give what some feel is a clear answer (e.g. Placozoa are at the base of the metazoan tree of life (ToL)), but a diversity of modern molecular data gives almost as many answers as there are exemplars, and even the largest molecular data sets could not solve the question and sometimes even suggest obvious zoological nonsense. Since the problems involved in this phylogenetic conundrum encompass a wide array of analytical freedom and uncertainty it seems questionable whether a further increase in molecular data (quantity) can solve this classical deep phylogeny problem. This review thus strikes a blow for evaluating quality data (including morphological, molecule morphologies, gene arrangement, and gene loss versus gene gain data) in an appropriate manner.

Ultrastructural analyses support different morphological lineages in the phylum Placozoa Grell, 1971.

The morphology and ultrastructure of 10 clonal placozoan lineages were studied. We scored several morphological characters at a cellular and intracellular level and identified a number of morphological differences among clones. Some differences appear clone specific and allow recognizing five distinct lineages based on morphological criteria only. These data will be crucial for a yet to be established placozoan systematics. Furthermore, we here describe three new diagnostic morphological characters for Placozoa: a new structure in the upper epithelium, called "concave disc," two distinct subpopulations of fiber cells, and especially small cells in the body margin. Besides the fiber cells appear to be arranged in several layers forming a complex, three-dimensional net not previously described. We also describe the marginal cells as the formerly suggested potential stem-cell type. The basic morphology is revised.

[From Vendian to Cambrian: the beginning of morphological disparity of modern Metazoan phyla].

Continuity of the transition from Precambrian to the Phanerozoic metazoan fauna at the phylum level is analyzed. The discrete traces of feeding on bacterial mats by Dickinsonia and similar organisms are explained by extracorporeal digestion, characteristic ofa placozoan level of organization, as in the extant Trichoplax. The absence of a morphologically developed anterior end of the body, of food-gathering appendages of any kind, and of appendages responsible for movement suggest that these animals were at this level of organization. Probably, an expanded placozoan level of organization can be assumed for most vendian animals. Against this background, new characters emerged in the Cambrian to be included in the body plan of extant animal phyla. The relationships between the morphological archetype (body plan) and morphogenetic archetype are considered. It is shown that major features of a morphogenetic archetype could be retained from the time the taxon was formed.

Concatenated analysis sheds light on early metazoan evolution and fuels a modern "urmetazoon" hypothesis.

For more than a century, the origin of metazoan animals has been debated. One aspect of this debate has been centered on what the hypothetical "urmetazoon" bauplan might have been. The morphologically most simply organized metazoan animal, the placozoan Trichoplax adhaerens, resembles an intriguing model for one of several "urmetazoon" hypotheses: the placula hypothesis. Clear support for a basal position of Placozoa would aid in resolving several key issues of metazoan-specific inventions (including, for example, head-foot axis, symmetry, and coelom) and would determine a root for unraveling their evolution. Unfortunately, the phylogenetic relationships at the base of Metazoa have been controversial because of conflicting phylogenetic scenarios generated while addressing the question. Here, we analyze the sum of morphological evidence, the secondary structure of mitochondrial ribosomal genes, and molecular sequence data from mitochondrial and nuclear genes that amass over 9,400 phylogenetically informative characters from 24 to 73 taxa. Together with mitochondrial DNA genome structure and sequence analyses and Hox-like gene expression patterns, these data (1) provide evidence that Placozoa are basal relative to all other diploblast phyla and (2) spark a modernized "urmetazoon" hypothesis.