Abstract

BackgroundThe ground pattern underlying the nervous system of the last common ancestor in annelids was long thought to be settled, consisting of a dorsal brain, circumoesophageal connectives and a subepithelial, ladder-like ventral nerve cord with segmental ganglia connected by paired connectives. With the advent of immunocytochemical stainings and confocal laser scanning microscopy, it becomes evident that its architecture is extremely diverse, which makes the reconstruction of a ground pattern in annelida challenging. Whereas the nervous systems of many different families has already been described, only very few studies looked at the diversity of nervous systems within such clades to give a closer estimate on how plastic the annelid nervous system really is. So far, little is known on syllid nervous system architecture, one of the largest and most diverse groups of marine annelids.ResultsThe position of the brain, the circumoesophageal connectives, the stomatogastric nervous system, the longitudinal nerves that traverse each segment and the innervation of appendages are relatively uniform within the clade. Both the number of connectives within the ventral nerve cord and the number of segmental nerves, which in earlier studies were used to infer phylogenetic relationships and to reconstruct an annelid ground pattern, are highly diverse and differ between genera or even within a given genus. Differences in the distribution of somata of the brain, the nuchal innervation and its associated cell bodies were found between Syllinae and Exogoninae and may be subfamily-specific.ConclusionsThe nervous system morphology of syllids very likely depends on the taxon-specific ecological requirements. Thus, it is not surprising that in a clade, which occupies such diverse niches as the Annelida, we find similar patterns in phylogenetically widely separated species in similar niches and a high degree of modularity within a family. Only standardized protocols and staining methods can lead to comparable results, but so far different approaches have been taken to describe annelid nervous systems, making homologization of certain structures difficult. This study provides the first thorough description of the nervous system in the family Syllidae, allowing more detailed comparisons between annelid families in the future.

Highlights

  • The ground pattern underlying the nervous system of the last common ancestor in annelids was long thought to be settled, consisting of a dorsal brain, circumoesophageal connectives and a subepithelial, ladderlike ventral nerve cord with segmental ganglia connected by paired connectives

  • Overview of the nervous system in Syllidae According to Fauchald and Rouse and Pleijel [48, 49] the anterior end of syllids comprise a well-developed prostomium, a peristomium reduced to the lips and an achaetigerous first segment followed by the second, chaetigerous segment (Fig. 1b)

  • The brain is connected by the dorsal and ventral root of the circumoesophaegeal connectives to the ventral nerve cord

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Summary

Introduction

The ground pattern underlying the nervous system of the last common ancestor in annelids was long thought to be settled, consisting of a dorsal brain, circumoesophageal connectives and a subepithelial, ladderlike ventral nerve cord with segmental ganglia connected by paired connectives. With the advent of immunocytochemical stainings and confocal laser scanning microscopy, it becomes evident that its architecture is extremely diverse, which makes the reconstruction of a ground pattern in annelida challenging. The actual situation is more complex [25,26,27] and the underlying ground pattern is being discussed to date [27], with authors suggesting two [7, 28] or five [26] connectives present in the ventral nerve cord of the last common ancestor of annelids. The high diversity of nervous system architecures in recent annelid clades, including the likely reduction of features in early branching families such as the Oweniidae [30] and Sipunculidae [31], makes it difficult to reconstruct the ground pattern at present

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