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It is widely accepted that multicellular animals (metazoans) constitute a monophyletic unit, deriving from ancestral choanoflagellate-like protists that gave rise to simple choanocyte-bearing metazoans. However, a re-assessment of molecular and histological evidence on choanoflagellates, sponge choanocytes, and other metazoan cells reveals that the status of choanocytes as a fundamental cell type in metazoan evolution is unrealistic. Rather, choanocytes are specialized cells that develop from non-collared ciliated cells during sponge cmbryogenesis. Although choanocytes of adult sponges have no obvious homologue among metazoans, larval cells transd rentiating into choanocytes at metamorphosis do have such homologues. The evidence reviewed here also indicates that sponge larvae are architecturally closer than adult sponges to the remaining metazoans. This may mean that the basic multicellular organismal architecture from which diploblasts evolved, that is, the putative planktonic archimetazoan, was more similar to a modern poriferan larva lacking choanocytes than to an adult sponge. Alternatively, it may mean that other metazoans evolved from a neotcnous larva of ancient sponges. Indeed, the Porifera possess some features of intriguing evolutionary significance: (1) widespread occurrence of internal lertilization and a notable diversity of' gastrulation modes, (2) dispersal through architecturally complcx lecithotrophic larvae, in which an ephemeral archenteron (in dispherula larvae) and multiciliated and syncytial cells (in trichimella larvae) occur, (3) acquisition of direct development by some groups, and (4) replacement of choanocyte-based lilter-feeding by carnivory in some sponges. Together, thcse features strongly suggest that the Porifera may have a longer and more complicated evolutionary history than traditionally assumed, and also that the simple anatomy of modern adult spongcs may have resulted from a secondary simplification. This makes the idea of a neotenous evolution less likely than that of a larva-like choanocyte-lacking archimetazoan. From this perspective, the view that choanoflagellates may be simplified sponge-derived metazoans, rather than protists, emerges as a viable alternative hypothesis. This idea neither conflicts with the available evidence nor can be disproved by it, and must be specifically re-examined by further approaches combining morphological and molecular information. Interestingly, several microbial lincages lacking choanocyte-like morphology, such as Corallochytrea, Cristidiscoidea, Ministeriida, and Mesomycetozoea, have recently been placed at the boundary between fungi and animals, becoming a promising source of information in addition to the choanoflagellates in the search for the unicellular origin of animal multicellularity.
A cladistic analysis of 87 morphological and life history characters of medusozoan cnidarians, rooted with Anthozoa, results in the phylogenetic hypothesis (Anthozoa (Hydrozoa (Scyphozoa (Staurozoa, Cubozoa)))). Staurozoa is a new class of Cnidaria consisting of Stauromedusae and the fossil group Conulatae. Scyphozoa is redefined as including those medusozoans characterized by strobilation and ephyrae (Coronatae, Semaeostomeae, and Rhizostomeae). Within Hydrozoa, Limnomedusae is identified as either the earliest diverging hydrozoan lineage or as the basal group of either Trachylina (Actinulida (Trachymedusae (Narcomedusae, Laingiomedusae))) or Hydroidolina (Leptothecata (Siphonophorae, Anthoathecata)). Cladistic results are highly congruent with recently published phylogenetic analyses based on 18S molecular characters. We propose a phylogenetic classification of Medusozoa that is consistent with phylogenetic hypotheses based on our cladistic results, as well as those derived from 18S analyses. Optimization of the characters presented in this analysis are used to discuss evolutionary scenarios. The ancestral cnidarian probably had a sessile biradial polyp as an adult form. The medusa is inferred to be a synapomorphy of Medusozoa. However, the ancestral process (metamorphosis of the apical region of the polyp or lateral budding involving an entocodon) could not be inferred unequivocally. Similarly, character states for sense organs and nervous systems could not be inferred for the ancestral medusoid of Medusozoa.
After feeding, polyps of colonial hydroids contract regularly, dispersing food throughout the colony via the gastrovascular fluid. Such contractions may trigger signaling pathways that allow colonies to grow in an adaptive manner, i.e., to initiate development of more polyps in food-rich areas and to suppress polyp development in food-poor areas. In this context, we investigated the structure and potential signaling of the junction between polyps and stolons in colonies of the hydroid Podocoryna carnea. Using transmission electron microscopy, we found that the density of mitochondrion-rich epitheliomuscular cells was low in polyp and stolon tissues except at or near the polyp-stolon junction, where many of these mitochondrion-rich cells occur in ectodermal tissue. In vivo fluorescence microscopy suggests that these mitochondria are a principal source of the metabolic signals of the colony. Both native fluorescence of NAD(P)H and fluorescence from peroxides (visualized with H2DCFDA) co-localize to this region of the polyp. Rhodamine 123 fluorescence suggests that both these metabolic signals emanate from mitochondria. To test whether such metabolic signals may be involved in colony pattern formation, inbred lines of P. carnea were used. Colonies of a runner-like inbred line grow with widely spaced polyps and long stolonal connections, much like wild-type colonies in a food-poor environment. Colonies of a sheet-like inbred line grow with closely spaced polyps and short stolonal connections, similar to wild-type colonies in a food-rich environment. Polyp-stolon junctions in runner-like and sheet-like colonies were imaged for the fluorescence of H2DCFDA. Densitometric analysis of this signal indicates that the mitochondria in epitheliomuscular cells of runner-like polyps emit greater amounts of peroxides. Because peroxides and other reactive oxygen species are frequently intermediaries in metabolic signaling pathways, we suspect that such signaling may indeed occur at polyp-stolon junctions, affecting colony pattern formation in these inbred lines and possibly in hydroid colonies in general.
The reproductive biology, development, and planula behavior of the gorgonian Pseudnpterogorgia etisabethae were studied at 2 sites in the Bahamas between 1996 and 2001. Colonies were gonochoric, and females brooded planulae on the colony surface. Gonads were observed only in colonies 18 cm high or larger. Spawning was asynchronous within and between sites but was concentrated 2-10 days after the new moons from late November through early January. Fertilized eggs developed into planulae over 1-2 days and the planulae remained attached to the surface of the female colony for an additional 2-4 days. Planulae were negatively buoyant and field observations suggest that larvae may settle within tens of meters of the maternal colony. P. elisabethue is harvested for natural products, and information on the reproduction of this commercially important species is crucial to the understanding of its population biology and to the development of management plans for the conservation of the species.
The monogeneans Decacotyle lymmae and D. tetrakordyle (Monocotylidae: Decacotylinae), from gills of the dasyatid stingrays Tueniuru lymma and Pustinachus sephen, respectively, have a single aperture for adhesive secretion on each side of the anterior ventrolateral region. Rod-shaped bodies (S1) and electron-dense spherical secretion (S2) exit through specialised ducts opening adjacent to one another within these apertures. The S1 bodies are 230 ± 11 nm wide and ≥ 4 µm long in D. lymmae and 240 ± 9 nm wide and ≥3.3 µm long in D. tetrakordyle. The S2 bodies have a diameter of 88 ± 7 nm in D. lymmue and 65 ± 6 nm in D. tetrakordyle. The apertures are unusual in being extremely small (internal diameter, 3–5 µm). Each aperture has a slit-like surface opening as small as 160 nm wide, surrounded by muscle fibres indicating that they may be opened and closed. The aperture is also surrounded and underlain by muscle fibres that may aid in secretion from, or even eversion of, the tissue within the aperture. Sensilldcilia are also found within the apertures. Additional secretions from anteromedian and anterolateral glands (body glands), each containing granular secretions, occur in profusion and exit anteriorly and posteriorly to the position of the apertures, through duct openings in the general body tegument. These granular secretions do not appear to be associated with anterior adhesion. Both species show similarities in aperture, underlying tissue, sense organ, and secretion detail, in accordance with findings from other monogenean genera, and which supports the importance of such data for phylogenetic studies.
Large knots (> 100 individuals) of adult horsehair worms (Gordius difficilis) were collected in several spring-fed habitats in Wisconsin and Minnesota. These aggregations were dominated numerically by males. Although distributions of length and diameter of males and females overlapped, females tended to be significantly longer and wider than males. Moreover, at any given length, females were wider than males. These findings confirm and add statistical support to previously published reports.
The albumen gland is a female accessory sex gland that synthesizes and secretes perivitelline fluid around pulmonate eggs. The perivitelline fluid is composed of mainly galactogen and proteins, and is thought to provide nourishment to the embryos during development. We have previously identified the major secretory protein of the albumen gland of the freshwater snail Helisoma duryi as a native glycoprotein of ∼288 kDa, consisting of four 66-kDa subunits. In this study, the major albumen gland protein in H. duryi was purified, cloned, and the full-length cDNA sequence determined. Nucleotide sequence analysis revealed that the albumen gland protein (HdAGP) shared 83% identity with a partial cDNA sequence from a developmentally regulated albumen gland protein in Biomphalaria glabrata. The HdAGP mRNA was detected by RT-PCR in the albumen gland, ovotestis, mantle and digestive gland. SDS-PAGE analysis of the albumen gland protein in egg masses at different stages of development showed that the amount of HdAGP steadily decreased during embryogenesis, suggesting its possible catabolism by the developing embryos. Protein domain searches suggested that the HdAGP shared limited sequence identity, and adopted a similar three-dimensional conformation to the bactericidal, permeability increasing, protein family, raising the possibility of a potential bactericidal function for this important reproductive/developmental protein.
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