The organization of the cnidarian nervous system has been widely documented in polyps and medusae, but little is known about the nervous system of planula larvae, which give rise to adult forms after settling and metamorphosis. We describe histological and cytological features of the nervous system in planulae of the hydrozoan Clava multicornis. These planulae do not swim freely in the water column but rather crawl on the substrate by means of directional, coordinated ciliary movement coupled to lateral muscular bending movements associated with positive phototaxis. Histological analysis shows pronounced anteroposterior regionalization of the planula's nervous system, with different neural cell types highly concentrated at the anterior pole. Transmission electron microscopy of planulae shows the nervous system to be unusually complex, with a large, orderly array of sensory cells at the anterior pole. In the anterior half of the planula, the basiectodermal plexus of neurites forms an extensive orthogonal network, whereas more posteriorly neurites extend longitudinally along the body axis. Additional levels of nervous system complexity are uncovered by neuropeptide-specific immunocytochemistry, which reveals distinct neural subsets having specific molecular phenotypes. Together these observations imply that the nervous system of the planula of Clava multicornis manifests a remarkable level of histological, cytological, and functional organization, the features of which may be reminiscent of those present in early bilaterian animals.
Thalia democratica is a cosmopolitan tunicate belonging to the Thaliacea class. To further investigate the anatomy of this species, immunohistochemical labelling was performed using anti-tubulin and anti-serotonin antibodies on specimens collected in the Mediterranean Sea. The anti-tubulin antibody stained the cilia of the endostyle, the pericoronal bands and of the gill bar, enabling a detailed description of these structures. Moreover, immunolabelling of the nervous system showed the presence of eight pairs of nerve fibres emerging from the neural ganglion. Serotonergic cells were observed in the distal tract of the intestine, along the pericoronal bands, and in the placenta of gravid blastozooids, as well as in the neural ganglion. The presence of serotonin in the central nervous system has also been reported in the larvae of ascidians and may be linked to the planktonic life of these animals, a condition shared by adult thaliaceans and ascidian larvae. This work improves our knowledge of the anatomy of T. democratica and demonstrates the presence of a complex serotonergic system.
Retinoic acid (RA) is a derivative of vitamin A known to be involved in the regulation of many developmental and physiological processes in chordates. Recently, evidence showing its presence and function in invertebrate deuterostomes and protostomes indicated the early evolutionary origin and conservation of the RA developmental machinery, with a key role also in neural differentiation. Moreover, it is known that retinoids can influence pattern specification in polyps of the hydroid Hydractinia echinata. The planula larvae of the hydrozoan Clava multicornis are characterized by a complex nervous system formed by a frontal neural plexus composed by two distinct peptidergic cell populations with antero‐posterior organization: a first, anterior‐most GLWamide immunoreactive (GLW‐IR) population and an RFamide immunoreactive (RF‐IR) cell belt just posterior to the first population. In contrast to most swimming planulae, C. multicornis larvae display a smooth gliding movement on the substrate, characterized by alternate bending of the anterior pole. We tested the effects of RA and of a RA antagonist on C. multicornis development by analyzing the nervous system organization and the photoresponsive behavior of larvae exposed to RA during their embryogenesis. After the exposure, the nervous system became completely disorganized, leading to the displacement of RFamide‐IR cells. Moreover, RA‐treated larvae did not exhibit the typical phototropic behavior of control specimens. Our results suggest that RA can alter the normal development of nervous elements in this hydroid species, supporting the hypothesis that the morphogenetic activity of RA predates the origin of chordates.
Music is believed to work as a bio-social tool enabling groups of people to establish joint action and group bonding experiences. However, little is known about the quality of the group members' interaction needed to bring about these effects. To investigate the role of interaction quality, and its effect on joint action and bonding experience, we asked dyads (two singers) to perform music in medieval "hocket" style, in order to engage their co-regulatory activity. The music contained three relative inter-onset-interval (IOI) classes: quarter note, dotted quarter note and eight note, marking time intervals between successive onsets (generated by both singers). We hypothesized that singers co-regulated their activity by minimizing prediction errors in view of stable IOI-classes. Prediction errors were measured using a dynamic Bayesian inference approach that allows us to identify three different types of error called fluctuation (micro-timing errors measured in milliseconds), narration (omission errors or misattribution of an IOI to a wrong IOI class), and collapse errors (macro-timing errors that cause the breakdown of a performance). These three types of errors were correlated with the singers' estimated quality of the performance and the experienced sense of joint agency. We let the singers perform either while moving or standing still, under the hypothesis that the moving condition would have reduced timing errors and increased We-agency as opposed to Shared-agency (the former portraying a condition in which the performers blend into one another, the latter portraying a joint, but distinct, control of the performance). The results show that estimated quality correlates with fluctuation and narration errors, while agency correlates (to a lesser degree) with narration errors. Somewhat unexpectedly, there was a minor effect of movement, and it was beneficial only for good performers. Joint agency resulted in a "shared," rather than a "we," sense of joint agency. The methodology and findings open up promising avenues for future research on social embodied music interaction.
Life and social sciences often focus on the social nature of music (and language alike). In biology, for example, the three main evolutionary hypotheses about music (i.e., sexual selection, parent-infant bond, and group cohesion) stress its intrinsically social character (Honing et al., 2015). Neurobiology thereby has investigated the neuronal and hormonal underpinnings of musicality for more than two decades (Chanda and Levitin, 2013; Salimpoor et al., 2015; Mehr et al., 2019). In line with these approaches, the present paper aims to suggest that the proper way to capture the social interactive nature of music (and, before it, musicality), is to conceive of it as an embodied language, rooted in culturally adapted brain structures (Clarke et al., 2015; D’Ausilio et al., 2015). This proposal heeds Ian Cross’ call for an investigation of music as an “interactive communicative process” rather than “a manifestation of patterns in sound” (Cross, 2014), with an emphasis on its embodied and predictive (coding) aspects (Clark, 2016; Leman, 2016; Koelsch et al., 2019). In the present paper our goal is: (i) to propose a framework of music as embodied language based on a review of the major concepts that define joint musical action, with a particular emphasis on embodied music cognition and predictive processing, along with some relevant neural underpinnings; (ii) to summarize three experiments conducted in our laboratories (and recently published), which provide evidence for, and can be interpreted according to, the new conceptual framework. In doing so, we draw on both cognitive musicology and neuroscience to outline a comprehensive framework of musical interaction, exploring several aspects of making music in dyads, from a very basic proto-musical action, like tapping, to more sophisticated contexts, like playing a jazz standard and singing a hocket melody. Our framework combines embodied and predictive features, revolving around the concept of joint agency (Pacherie, 2012; Keller et al., 2016; Bolt and Loehr, 2017). If social interaction is the “default mode” by which human brains communicate with their environment (Hari et al., 2015), music and musicality conceived of as an embodied language may arguably provide a route toward its navigation.
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