Homology of ciliary bands in Spiralian Trochophores

A number of hypotheses have been presented regarding the originsof the metazoans and, more specifically, the Bilateria. Usingvarious phylogenetic analyses, characteristics have been mappedon phylogenetic trees to infer ancestral body plans and lifehistory strategies of those ancestors. Many arguments on theevolution of the Bilateria are based on the presumed homologyof certain characteristics of extant larva and adults, includingvarious ciliated bands involved in feeding and locomotion. Thisarticle considers a recent study indicating that the second,downstream-collecting, ciliated band in the veliger larva ofthe gastropod mollusc, Crepidula fornicata, is actually derivedfrom secondary trochoblasts (derived from second quartet micromeres),that normally form part of the prototrochal band found in otherspiralian phyla (Hejnol et al. 2007). Despite previous arguments,these new findings suggest that the second ciliated band inthe veliger larva is not homologous to the metatroch found inthe trochophore larva of some other spiralians, such as theannelid, Polygordius lacteus. In the latter case, the metatrochwas reported to be formed by a different set of lineage precursors(derived from third quartet micromeres) (Woltereck 1904). Thesefindings have important implications for the interpretationof various hypotheses related to the evolution of metazoan phyla.     

Opposed bands of cilia in the nonfeeding larvae of the serpulid annelid Salmacina tribranchiata

The nonfeeding planktonic larvae of marine invertebrates typically lack larval feeding structures. One puzzling exception to this generalization is the annelid clade Sabellidae, in which nonfeeding larvae possess ciliary bands (specifically, food groove and metatroch) that, to the best of our knowledge, have no function other than in feeding. Nishi and Yamasu (1992b, Bulletin of the College of Sciences, University of the Ryukyus, 54 , 107–121) published a scanning electron micrograph showing that nonfeeding larvae of the serpulid annelid Salmacina dysteri also possess food groove and metatrochal cilia. Here I demonstrate that nonfeeding larvae of Salmacina tribranchiata also bear ciliary bands identifiable as food groove and metatroch by position. High‐speed video of ciliary beat patterns shows that, together with the prototrochal cilia, these bands function in an opposed band system. The presence of feeding structures in nonfeeding annelid larvae is thus more widely distributed than previously recognized. The presence of feeding structures may make evolutionary transitions to planktotrophy more likely, and may underlie an inferred origin of larval feeding in the common ancestor of one of the two major clades of serpulid annelids, Serpulinae.     

High-resolution fate map of the snail Crepidula fornicata: the origins of ciliary bands, nervous system, and muscular elements

The littorinimorph gastropod Crepidula fornicata shows a spiralian cleavage pattern and has been the subject of studies in experimental embryology, cell lineage, and the organization of the larval nervous system. To investigate the contribution of early blastomeres to the veliger larva, we used intracellular cell lineage tracers in combination with high-resolution confocal imaging. This study corroborates many features derived from other spiralian fate maps (such as the origins of the hindgut and mesoderm from the 4d mesentoblast), but also yields new findings, particularly with respect to the origins of internal structures, such as the nervous system and musculature that have never been described in detail. The ectomesoderm in C. fornicata is mainly formed by micromeres of the 3rd quartet (principally 3a and 3b), which presumably represents a plesiomorphic condition for molluscs. The larval central nervous system is mainly formed by the micromeres of the 1st and 2nd quartet, of which 1a, 1c, and 1d form the anterior apical ganglion and nerve tracks to the foot and velum, and 2b and 2d form the visceral loop and the mantle cell. Our study shows that both first and second velar ciliary bands are generated by the same cells that form the prototroch in other spiralians and apparently bear no homology to the metatroch found in annelids.     

Trochophore concepts: ciliary bands and the evolution of larvae in spiralian Metazoa

‘Trochophore’ is a term used in a strict sense for larvae having an opposed-band method of feeding, involving a prototroch and metatroch. Other ciliary bands such as a telotroch and neurotroch may be present. The trochophore has been proposed to represent the ancestral larval form for a group of metazoan phyla (including all members of the Spiralia). The name trochophore is also often applied to larvae that do not conform to the above definition. A cladistic analysis of spiralian taxa (with special reference to polychaete annelids), based on a suite of adult and larval characters, is used to assess several hypotheses: (1) that the trochophore (in a strict sense) is a plesiomorphic form for the Spiralia; (2) that die stricdy defined trochophore is plesiomorphic for members of the Spiralia such as the Polychaeta. The homology of each of the various separate ciliary bands of spiralian larvae, and features such as the apical tuft and protonephridia is also assessed. The results favour the conclusion that the trochophore, if defined as a feeding larval form using opposed bands, should not be regarded as an ancestral (= plesiomorphic) type for the Spiralia, or any other large taxon such as the Polychaeta or Mollusca. The evidence suggests that the various ciliary bands have differing evolutionary histories, and only the Echiura (possibly an annelid group) has members with the classical trochophore. The trochophore is re-defined as a larval form with a prototroch. This broad definition covers a wide variety of larvae, and matches the current usage more accurately than the restricted term. Features such as the neurotroch, telotroch and opposed-band feeding show convergence and reversals. The nature of the metatroch requires further investigation. The presence of a prototroch (and hence trochophore larvae) is used to identify an apomorphy-based taxon, Trochozoa, that includes the first ancestor to have evolved a prototroch and all its descendants. This minimally includes the Annelida [sensu lato), Echiura, Entoprocta, Mollusca and Sipuncula and is a less inclusive taxon than the Spiralia.     

Opposed ciliary bands in the feeding larvae of sabellariid annelids

The larvae of marine annelids capture food using an unusual diversity of suspension-feeding mechanisms. Many of the feeding mechanisms of larval annelids are poorly known despite the abundance and ecological significance of both larvae and adults of some annelid taxa. Here we show that larvae of two species of sabellariid annelids, Sabellaria cementarium and Phragmatopoma californica, bear prototrochal and metatrochal cilia that beat in opposition to each other. For larvae of S. cementarium, we provide evidence that these opposed bands of cilia are used to capture suspended particles. In video recordings, captured particles were overtaken by a prototrochal cilium and then moved with the cilium to the food groove, a band of cilia between the prototroch and metatroch. They were then transported by cilia of the food groove to the mouth. Lengths of the prototrochal cilia, lengths of the prototrochal ciliary band, size range of the particles captured, and estimated rates of clearance increased with larval age and body size. Confirmation of the presence of opposed bands in larvae of sabellariids extends their known occurrence in the annelids to members of 10 families. Opposed bands in these different taxa differ in the arrangements and spacing of prototrochal and metatrochal cilia, and in whether they are used in combination with other feeding mechanisms. Opposed bands appear to be particularly widespread among the larvae of sabellidan annelids (a clade that includes sabellariids, sabellids, and serpulids), even in some species whose larvae do not feed. A parsimony analysis suggests that opposed bands are ancestral in this clade of annelids.     

Foraging by the mud snail, Ilyanassa obsoleta (Say), modulates spatial variation in benthic community structure

We investigated the foraging behavior of the mud snail, Ilyanassa obsoleta, and its consequences for macrobenthic community structure on mud flats on Long Island, NY, USA. Field sampling demonstrated strong spatial heterogeneity in the population densities of I. obsoleta. We experimentally tested three hypotheses: (i) I. obsoleta are strongly attracted to areas with high levels of detritus; (ii) local abundances of deposit-feeding annelids are limited by detritus; and (iii) the foraging activities of I. obsoleta negatively affects annelid assemblages. We manipulated the density of mud snails using inclusion fences and the levels of detritus using dried Ulva. Results showed that high densities of I. obsoleta were attracted to areas enriched with Ulva detritus. In addition, high densities of snails negatively affected abundances of annelids, with the opportunistic species, Capitella spp. and Paranais litoralis, being most affected. The addition of Ulva detritus had more specific effects on annelid assemblages. Only Capitella spp. showed a significant positive response, although previous evidence has demonstrated that higher experimental detrital inputs stimulated growth of other species of annelids and microphytobenthos. In an experimental treatment with enhanced detritus and low densities of snails, we found population abundances of opportunistic annelids (up to 200,000 m⁻²) substantially larger than has ever been recorded in 5 years of sampling. Because mud snails in natural areas actively search, locate and exploit areas with enhanced detritus and their foraging negatively affects abundances of opportunistic worms, I. obsoleta probably controls the upper limits of annelid abundance in the field. Foraging behavior of I. obsoleta therefore modulates spatial variation in benthic community structure in an environment where limiting resources are patchily distributed.     

Embryonic and post-embryonic development of the polyclad flatworm Maritigrella crozieri; implications for the evolution of spiralian life history traits

Background

Planktonic life history stages of spiralians share some muscular, nervous and ciliary system characters in common. The distribution of these characters is patchy and can be interpreted either as the result of convergent evolution, or as the retention of primitive spiralian larval features. To understand the evolution of these characters adequate taxon sampling across the Spiralia is necessary. Polyclad flatworms are the only free-living Platyhelminthes that exhibit a continuum of developmental modes, with direct development at one extreme, and indirect development via a trochophore-like larval stage at the other. Here I present embryological and larval anatomical data from the indirect developing polyclad Maritrigrella crozieri, and consider these data within a comparative spiralian context.

Results

After 196 h hours of embryonic development, M. crozieri hatches as a swimming, planktotrophic larva. Larval myoanatomy consists of an orthogonal grid of circular and longitudinal body wall muscles plus parenchymal muscles. Diagonal body wall muscles develop over the planktonic period. Larval neuroanatomy consists of an apical plate, neuropile, paired nerve cords, a peri-oral nerve ring, a medial nerve, a ciliary band nerve net and putative ciliary photoreceptors. Apical neural elements develop first followed by posterior perikarya and later pharyngeal neural elements. The ciliated larva is encircled by a continuous, pre-oral band of longer cilia, which follows the distal margins of the lobes; it also possesses distinct apical and caudal cilia.

Conclusions

Within polyclads heterochronic shifts in the development of diagonal bodywall and pharyngeal muscles are correlated with life history strategies and feeding requirements. In contrast to many spiralians, M. crozieri hatch with well developed nervous and muscular systems. Comparisons of the ciliary bands and apical organs amongst spiralian planktonic life-stages reveal differences; M. crozieri lack a distinct ciliary band muscle and flask-shaped epidermal serotonergic cells of the apical organ. Based on current phylogenies, the distribution of ciliary bands and apical organs between polyclads and other spiralians is not congruent with a hypothesis of homology. However, some similarities exist, and this study sets an anatomical framework from which to investigate cellular and molecular mechanisms that will help to distinguish between parallelism, convergence and homology of these features.     

The Larval Nervous System of Polygordius lacteus Scheinder, 1868 (Polygordiidae,Polychaeta): Immunocytochemical Data

The nervous system of the planktotrophic trochophore larva of Polygordius lacteus has been investigated using antibodies to serotonin (5-HT) and the neuropeptide FMRFamide. The apical ganglion contains three 5-HT-ir neurons, many FMRFamide-ir neurons and a tripartate 5-HT-ir and FMRFamide-ir neuropil. A lateral nerve extends from each side of the apical ganglion across the episphere and the ventral hyposphere, where the two nerves combine to form the paired ventral nerve cord. These nerves have both 5-HT-ir and FMRFamide-ir processes. Three circumferential nerves are associated with the ciliary bands: two prototroch and one metatroch nerve. All contain 5-HT-ir and FMRFamide-ir processes. An oral nerve plexus also contain both 5-HT-ir and FMRFamide-ir processes develops from the metatroch nerve, and an esophageal ring of FMRFamide-ir processes develops in later larval stages. In young stages the ventral ganglion contains two 5-HT-ir and two FMRFamide-ir perikarya; during development the ventral ganglion grows caudally and adds additional 5-HR-ir and FMRFamide-ir perikarya. These are the only perikarya that could be found along the lateral nerve and ventral nerve cord. The telotroch nerve develops from the ventral nerve cord. The 5-HT-ir and FMRFamide-ir part of the nervous system is strictly bilateral symmetric. and much of the system (i.e. apical ganglion, lateral nerves ventral nerve cord, dorsal nerve and oral plexus) is retained in the adult.     

Ciliary band gene expression patterns in the embryo and trochophore larva of an indirectly developing polychaete

The trochophore larvae of indirectly developing spiralians have ciliary bands with motor and feeding functions. The preoral prototroch ciliary band is the first differentiating organ in annelid and mollusk embryos. Here we report the expression of several ciliary band markers during embryogenesis and early larval stages of the indirectly developing polychaete Hydroides elegans. Genes with similarity to caveolin, beta-tubulin, alpha-tubulin, and tektin are expressed in the eight primary prototroch precursors, 1q(221) and 1q(212). Blastomeres 1q(221) and 1q(212) locate at the same equatorial latitude after the complementary asymmetric division of their 1q(22) and 1q(21) precursors. In addition, caveolin and alpha-tubulin are expressed in the metatroch and adoral ciliary zone. Caveolin is expressed in foregut ciliated cells, and alpha-tubulin is expressed in apical tuft ciliated cells. The expression of a beta-thymosin homolog is restricted to 1q(122) and 1q(121) blastomeres, which locate just above and in close association with the eight primary prototroch cells 1q(221) and 1q(212). In addition, the beta-thymosin homolog has a transient expression in the hindgut and apical zone. The expression of all these genes provides a landmark for the early specification of ciliary bands and other ciliated organs.     

Too cold to prosper—winter mortality prevents population increase of the introduced American slipper limpet Crepidula fornicata in northern Europe

The distribution of the introduced American slipper limpet Crepidula fornicata (L.) on the Atlantic coast of Europe shows a clear latitudinal gradient. In the south, C. fornicata may reach abundances of several thousand individuals per meter squared accompanied by diverse ecological and economic effects. In contrast, abundances at northern waters as in Germany, Denmark and Norway barely reach 100 ind. m⁻², indicating one or several limiting factors for population increase. By studying a population in the northern Wadden Sea of Germany, we examined four factors that we regarded as likely to potentially limit population increase of C. fornicata: (1) high predation by the main benthic mollusc predators, (2) high infestation by parasites, (3) low reproductive output and growth in cold waters and (4) high winter mortality during freezing winters. Our results do not indicate biotic restrictions: The main benthic mollusc predators, shore crabs (Carcinus maenas) and sea stars (Asterias rubens) strongly preferred the dominant blue mussel (Mytilus edulis) in choice experiments and no infestation with parasitic trematodes were observed. Further, there was no indication of limitation in reproduction and growth since abundant larvae were observed in the water column and the period of reproduction (April to September) as well as growth rates (recruits: 9-14 mm first summer) match with data from southern areas. In contrast, population dynamics of C. fornicata was strongly affected by cold winters: During the two winters investigated, mortality amounted to 56-64% with up to 97% on single mussel beds contrasting to 11-14% yearly mortality in areas without frost in southern Europe. Negative winter effects are also suggested by remarkably low larval abundances after an exceptionally severe winter. Our results suggest that winter mortality is the main limiting factor for population increase of C. fornicata in the study area. We propose that milder winters as a corollary of global warming may allow for an increase in the abundance of northern populations combined with a northward shift of the concomitant negative ecological and economic effects.     

The ‘organizing’ role of the D quadrant in an equal-cleaving spiralian,Lymnaea stagnalis as studied by UV laser deletion of macromeres at intervals between third and fourth quartet formation

Summary

In the spiralian embryos studied which display unequal-cleavage at the first two cleavages (either by a polar lobe or an asymmetric cleavage mechanism) the D quadrant is determined at the four cell stage by an unequal segregation of cytoplasmic stuffs. The normal formation of eyes, foot, and shell by overlying micromeres in these forms requires the inductive interaction with the D quadrant before the formation of the third quartet of micromeres. In equal-cleaving spiralians the D quadrant (3D macromere) becomes determined as a result of inductive interactions with first quartet derivatives (animal-vegetal interaction) sometime after the production of the third quartet of micromeres. This paper investigates the exact timing of D quadrant determination and the inductive role of third-order macromeres on the development of micromere derived structures in an equal-cleaving spiralian. Deletions of third-order macromeres, and their derivatives, were performed without rupturing the egg capsule membrane of the Lymnaea embryo with a UV laser microbeam. Virtually normal snails were produced when the 3A, 3B, 3C, or 4D macromere was irradiated. Juvenile snails lacking all mesodermal structures but possessing eyes, foot, and shell were obtained when the mesentoblast (4d) or its progenitor (3D) were deleted. Furthermore, ‘mesoderm-less’ snails were produced by deleting one of the two possible 3D candidates (cross furrow macromeres) as early as 20 min after third quartet formation. These results indicate that the 3D macromere begins to become determined at, or soon after, animal-vegetal interaction; before the 3D macromere becomes visibly distinguishable from the 3B macromere. The results also demonstrate that normal pattern formation in the overlying micromeres does not require the ‘prolonged’ interaction with an asymmetrically positioned 3D macromere. Possible adhesive differences between the 3D macromere and the remaining three macromeres are also revealed.     

Infestation of the invasive mollusc <Emphasis Type="Italic">Crepidula fornicata</Emphasis> by the native shell borer <Emphasis Type="Italic">Cliona celata</Emphasis>: a case of high parasite load without detrimental effects

The temporal prevalence of the widespread boring sponge Cliona celata and its effects were analysed in a population of the invasive mollusc Crepidula fornicata. This mollusc produces extra shell material when infested with the endolithic sponge, suggesting that infestation may be detrimental for C. fornicata growth and/or reproduction. For 37 months, size, sex, female reproductive status and sponge-infestation stage were recorded for 300 individuals sampled every month in the Bay of Morlaix (France). In the 12,049 individuals examined, the prevalence of C. celata was high with a monthly average of 43.1% of the individuals hosting the sponge. The relative proportion of heavily infested individuals generally increased over time. Nevertheless, a cyclic decrease occurred every 10 months, suggesting putative episodes of mortality of heavily infested individuals. The gregarious behaviour of the mollusc seemed to promote the high prevalence of the sponge, which may propagate through contact between neighbouring C. fornicata individuals. Due to the sex-size relationship in protandrous C. fornicata, females were far more infested than males. We did not find evidence for a cost of producing extra shell material on somatic growth or on female fertility, and the boring sponge is unlikely to substantially affect the sex-change patterns in C. fornicata. The limited effects of the endolithic sponge on C. fornicata contrasts with the documented damage to some local species, including commercially exploited shellfish, suggesting that C. fornicata may alter the infestation dynamics in the surrounding native community. Dedicated studies are now needed to investigate the extent and mechanisms of these species interactions.     

Embryonic Development of the Phoronid Phoronis ijimai

We studied the embryonic development of the phoronid Phoronis ijimai Oka, 1897. The egg cleavage is radial. The fourth and fifth cleavage furrows extend along the meridian of the egg. The blastula is flattened. Gastrulation occurs by a combination of epiboly, bending, and invagination. The mesoderm originates from two sources. The anterior mesoderm arises through immigration and gives rise to the first and second coeloms. The third coelomic mesoderm originates enterocoelically from the hindgut. The newly hatched larva has preoral and postoral ciliary bands, which can be compared with the corresponding ciliary bands of dipleurula and with the prototroch and metatroch of trochophore larvae.     

Development of the prototroch in embryogenesis of Nereis virens (polychaeta)

Prototroch formation was studied in the polychaete Nereis virens using light, scanning electron, and confocal laser microscopy. Cell lineage of trochoblasts was followed and chronology of their appearance was determined. The prototroch ciliary ring is formed by twelve descendants of micromere 1m ². The remaining four primary trochoblasts have no cilia and, together with descendants of accessory trochoblasts, become anterior supporting cells of the prototroch. Posterior supporting cells are formed by secondary trochoblasts, which are derived from the second micromere quartet 2m. The results obtained make it possible to analyze one of the ancient programs of animal development.     

Swimming Speed of Larval Snail Does Not Correlate with Size and Ciliary Beat Frequency

Many marine invertebrates have planktonic larvae with cilia used for both propulsion and capturing of food particles. Hence, changes in ciliary activity have implications for larval nutrition and ability to navigate the water column, which in turn affect survival and dispersal. Using high-speed high-resolution microvideography, we examined the relationship between swimming speed, velar arrangements, and ciliary beat frequency of freely swimming veliger larvae of the gastropod Crepidula fornicata over the course of larval development. Average swimming speed was greatest 6 days post hatching, suggesting a reduction in swimming speed towards settlement. At a given age, veliger larvae have highly variable speeds (0.8–4 body lengths s⁻¹) that are independent of shell size. Contrary to the hypothesis that an increase in ciliary beat frequency increases work done, and therefore speed, there was no significant correlation between swimming speed and ciliary beat frequency. Instead, there are significant correlations between swimming speed and visible area of the velar lobe, and distance between centroids of velum and larval shell. These observations suggest an alternative hypothesis that, instead of modifying ciliary beat frequency, larval C. fornicata modify swimming through adjustment of velum extension or orientation. The ability to adjust velum position could influence particle capture efficiency and fluid disturbance and help promote survival in the plankton.     

The epitome of hand waving? Larval feeding and hypotheses of metazoan phylogeny

SUMMARY The homology of larval forms, and particularly their feeding methods, has been a major element in some recent discussions about animal phylogeny. "Downstream feeding" is one of two main larval-feeding modes and is usually equated to an opposed-band system with ciliary bands called the prototroch and metatroch. Feeding in larvae is reviewed here and the homology hypothesis of downstream larval feeding is expanded, encompassing any feeding involving the prototroch. It is often argued that the presence of planktotrophic larvae using downstream feeding is plesiomorphic among spiralian animals, and that there is a bias in transformations, such that feeding larvae tend to be lost rather than gained. These hypotheses are assessed using cladistic parsimony methodology, in relation to Spiralia, Trochozoa, and with particular reference to polychaete annelids. Methods adopted for the possibility of a bias in transformations toward loss of downstream larval feeding include: expanded primary homology arguments, character reconstructions favoring reversals, and polymorphic terminals coded as having downstream larval feeding. Nevertheless, all analyses show that downstream larval feeding appears to have evolved multiple times from a lecithotrophic condition. The results support a conclusion that the prototroch was primarily locomotory, and has become associated with feeding a number of times. Hypotheses of metazoan phylogeny predicated on the assumption that downstream-feeding larvae are plesiomorphic are re-assessed.     

Growth Inhibition of Phytopathogenic Fungi and Oomycetes by Basidiomycete Irpex lacteus and Identification of its Antimicrobial Extracellular Metabolites

In dual culture confrontation assays, basidiomycete Irpex lacteus efficiently antagonized Fusarium spp., Colletotrichum spp., and Phytophthora spp. phytopathogenic strains, with growth inhibition percentages between 16.7–46.3%. Antibiosis assays evaluating the inhibitory effect of soluble extracellular metabolites indicated I. lacteus strain inhibited phytopathogens growth between 32.0–86.7%. Metabolites in the extracellular broth filtrate, identified by UPLC-QTOF mass spectrometer, included nine terpenes, two aldehydes, and derivatives of a polyketide, a quinazoline, and a xanthone, several of which had antifungal activity. I. lacteus strain and its extracellular metabolites might be valuable tools for phytopathogenic fungi and oomycete biocontrol of agricultural relevance.     

Quartet Analysis of Putative Horizontal Gene Transfer in Crenarchaeota

Horizontal gene transfers (HGT) between four Crenarchaeota species (Metallosphaera cuprina Ar-4^T, Acidianus hospitalis W1^T, Vulcanisaeta moutnovskia 768-28^T, and Pyrobaculum islandicum DSM 4184^T) were investigated with quartet analysis. Strong support was found for individual genes that disagree with the phylogeny of the majority, implying genomic mosaicism. One such gene, a ferredoxin-related gene, was investigated further and incorporated into a larger phylogeny, which provided evidence for HGT of this gene from the Vulcanisaeta lineage to the Acidianus lineage. This is the first application of quartet analysis of HGT for the phylum Crenarchaeota. The results have shown that quartet analysis is a powerful technique to screen homologous sequences for putative HGTs and is useful in visually describing genomic mosaicism and HGT within four taxa.     

Some aspects of spiralian development

Nielsen, C. 2010. Some aspects of spiralian development. —Acta Zoologica (Stockholm) 91 : 20–28 Spiralian development is not only a characteristic early cleavage pattern, with shifting orientations of the cleavage planes, but also highly conserved cell lineages, where the origin of several organs can be traced back to identifiable cells in the lineage. These patterns are well documented in annelids, molluscs, nemertines, and platyhelminths and are considered ancestral of a bilaterian clade including these phyla. Spiral cleavage has not been documented in ecdysozoans, and no trace of the spiral development pattern is seen in phoronids and brachiopods. Origin of the spatial organization in spiralian embryos is puzzling, but much of the information appears to be encoded in the developing oocyte. Fertilization and “pseudofertilization” apparently provides the information defining the secondary, anterior‐posterior body axis in many species. The central nervous system consists of three components: an apical organ, derived from the apical blastomeres 1a¹¹¹‐1d¹¹¹, which degenerates before or at metamorphosis; the cerebral ganglia derived from other blastomeres of the first micromere quartet and retained in the adult as a preoral part of the brain; and the originally circumblastoporal nerve cord, which has become differentiated into a perioral part of the brain, the paired or secondarily fused ventral nerve cords, and a small perianal nerve ring.     

What studies of turbellarian embryos can tell us about the evolution of development mechanisms

In spiralian embryos determination of the axes of bilateral symmetry is associated with D quadrant specification. This can occur late through equal cleavage and cell interactions (conditional specification) or by the four-cell stage through unequal cleavage and cytoplasmic localization (autonomous specification). Freeman & Lundelius (1992) suggest that in spiralian coelomates the former method is ancestral and the latter derived, with evolutionary pressure to shorten metamorphosis resulting in early D quadrant determination through unequal cleavage and appearance of adult features in the larvae. Because of the key phylogenetic position of the turbellarian platyhelminthes, understanding the method of axis specification in this group is important in evaluating the hypothesis. Polyclad development, with equal quartet spiral cleavage, is believed to represent the most primitive condition among living turbellarians and has been examined experimentally in Hoploplana inquilina. Blastomere deletions at the two and four-cell stage produce larvae that are abnormal in morphology and symmetry, indicating that early development is not regulative, and also establish that the embryo does not have an invariant cell lineage. Deletions of micromeres and macromeres at the eight-cell stage indicate that cell interactions are involved in dorso-ventral axis determination, with cross-furrow macromeres playing a more significant role than non-cross-furrow cells. The results support the idea that conditional specification is the primitive developmental mode that characterized the common ancestor of the turbellarians and spiralian coelomates. Evolutionary trends in development in polyclads and other turbellarian orders are discussed.