Auswahl der wissenschaftlichen Literatur zum Thema „Ascidies coloniaux“

Although ascidians form a conspicuous part of sessile assemblages in the Caribbean, no specialized inventories have been developed in Venezuela, except for a list of 15 species reported from Margarita Island (1984). Here we present the results of a taxonomic workshop held in the Universidad de Oriente, Boca del Rio, Margarita Island during April 20-25 of 2009, sponsored by the NaGISA-Caribbean Sea program. La Restinga National park was surveyed and we found 29 species belonging to 19 genera and 10 families. The most abundant colonial species were Clavelina oblonga, Aplidium accarense, Polyclinum constellatum, Distaplia bermudensis, Symplegma rubra, S. brakenhielmi, Botrylloides nigrum and Ecteinascidia turbinata. Among the solitary ascidians Phallusia nigra, Ascidia curvata, Microcosmus exasperatus, Styela canopus, Styela sp.1 and Styela sp. 2 were the most abundant. The ascidian diversity in this lagoon is one of the highest for similar habitats in the Caribbean and seven species are new registers for Venezuela (Ascidia curvata, Ecteinascidia styeloides, Aplidium accarense, Distaplia stylifera, Trididemnum orbiculatum, Symplegma rubra, and S. brakenhielmi). One point of concern for the conservation of la Restinga National Park is the presence of possible introduced species, and some management procedures are suggested.

Non-native ascidians have long dominated the artificial structures in southern California’s (United States) marinas and harbors. To determine the change in ascidian abundance and community composition over the last several decades, in 2019–2020 we replicated surveys from 1994–2000. We then created nMDS plots using the abundance data collected in the 1994–2000 and 2019–2020 surveys to compare the two groups. Range and average abundance per species were analyzed to determine trends and changes in ascidian community composition. Of the species used for comparison, four are native, three are cryptogenic, and 12 are non-native. As predicted by Lambert and Lambert, non-native species have persisted in southern California; however, ranges and abundances have changed. The only native species found consistently in both sets of surveys, Ascidia ceratodes, remained rare in 2019–2020, with an unchanged average abundance. Several non-native species increased in abundance or remained common. The non-native colonial species Polyandrocarpa zorritensis had the greatest influence on the dissimilarity between the surveys, increasing from rare in 1994–2000 to more common in 2019–2020, and spreading north to Santa Barbara. Several non-native species confined to San Diego in the 1994–2000 surveys have also spread north, such as Botrylloides giganteus and Styela canopus which were found in Santa Barbara in 2019–2020. A formerly unidentified Aplidium sp. has now been identified as the non-native Aplidium accarense. There have also been additional introductions since 2000, including Ascidia cf. virginea and the first report of Ascidiella aspersa in the NE Pacific. The overwhelming trends of the surveys indicate that we will continue to see an increase and persistence of newly introduced non-natives in Southern California marinas, with possible continued northward expansion.

The seasonality of sexual reproduction was studied in two Prochloron-bearing ascidians, Trididemnum cyclops and Lissoclinum bistratum, on a subtropical coral reef off Okinawajima Island, Japan. These colonial ascidians had testes and/or eggs/embryos from spring to summer. Embryos with tails occurred in summer. Whereas many photosymbiotic didemnids are thought to be sexually mature throughout the year in the tropics, sexual reproduction of the same species in subtropical waters may be limited to spring and summer. The subtropical winter may be too cold for gonad formation. A histological study of sexually mature colonies showed no Prochloron cells attached to ascidian larvae in the pre-hatch stage.

A new species of colonial Styelid ascidian of the genus Polyzoa was found attached to the test of solitary ascidians collected by divers off the coast of Georgia, U.S.A. The colony of this new species, consisting of small sandy zooids, is cryptic and difficult to detect. The species is characterized by three longitudinal branchial vessels on each side of the body and is the second species of the genus reported for the Atlantic.

To characterize the ultraviolet (UV) light-absorbing function of the ascidian tunic, the light absorption spectrum was compared in 22 ascidian species collected from tropical and temperate waters. Non-photosymbiotic ascidians (17 species) had transparent tunics transmitting both visible and UV light, or pigmented or opaque tunics equally absorb both UV and visible light. However, a prominent absorption peak around 320 nm was exclusively found in the colonial ascidians hosting the algal symbiont Prochloron sp., and this absorption peak corresponded to absorption of UV-A (320–400 nm) and UV-B (280–320 nm). The UV-absorbing substances were extracted with methanol. They were mycosporine-like amino acids (MAAs): mycosporine-glycine, palythine, shinorine, and porphyra-334. The MAAs are thought to be synthesized by the shikimic acid pathway that does not occur in animals. Since the isolated Prochloron cells contain MAAs, the symbionts are the most possible candidates as the source of the MAAs in the tunic. In Diplosoma virens, the composition of MAAs was different between isolated Prochloron cells and colony residue from which Prochloron cells were extracted.

Protochordate reproduction and development are influenced by many kinds of environmental factors. For example, spawning, sexual and asexual reproduction, larval behaviour, and life-cycle transitions (metamorphosis) are key processes known to be affected by environmental factors. This review must be restricted primarily to only one group of protochordates, the ascidians or "sea squirts", because information on the reproductive ecology of hemichordates and cephalochordates is limited to only a few studies. Topics discussed in the present review include (i) environmental factors that regulate larval settlement, (ii) how pelagic embryos avoid damage to DNA caused by UV radiation, (iii) the effect of water temperature and food availability on sexual reproduction in colonial ascidians, (iv) environmental regulation of asexual budding, (v) environmental regulation of metamorphosis, and (vi) the possible role of the environment in the evolution of direct-developing ascidians. A novel role for HSP90 and nitric oxide signaling in the integration of environmental factors with cell signaling pathways in ascidians is discussed near the end of this review. Throughout this review, the multiple roles of environmental stress on ascidian reproduction and development are emphasized.

The swimming larva represents the dispersal phase of ascidians, marine invertebrates belonging to tunicates. Due to its adhesive papillae, the larva searches the substrate, adheres to it, and undergoes metamorphosis, thereby becoming a sessile filter feeding animal. The larva anatomy has been described in detail in a few species, revealing a different degree of adult structure differentiation, called adultation. In the solitary ascidian Halocynthia roretzi, a species reared for commercial purposes, embryogenesis has been described in detail, but information on the larval anatomy is still lacking. Here, we describe it using a comparative approach, utilizing 3D reconstruction, as well as histological/TEM observations, with attention to its papillae. The larva is comparable to those of other solitary ascidians, such as Ciona intestinalis. However, it displays a higher level of adultation for the presence of the atrium, opened outside by means of the atrial siphon, and the peribranchial chambers. It does not reach the level of complexity of the larva of Botryllus schlosseri, a phylogenetically close colonial ascidian. Our study reveals that the papillae of H. roretzi, previously described as simple and conform, exhibit dynamic changes during settlement. This opens up new considerations on papillae morphology and evolution and deserves to be further investigated.

Simple and colonial ascidians were collected at different depths at Arraial do Cabo, Rio de Janeiro, between 2000 and 2003. The collection here presented includes representatives of the families Clavelinidae (Clavelina oblonga), Polycitoridae (Cystodytes dellechiajei), Polyclinidae (Polyclinum constellatum and Polyclinum molle sp. nov.), Holozoidae (Distaplia bermudensis), Ascidiidae (Ascidia sydneiensis and Phallusia nigra), Styelidae (Botrylloides giganteum, Botrylloides nigrum, Symplegma brakenhielmi, Symplegma rubra, Polyandrocarpa anguinea, Eusynstyela floridana, Eusynstyela tincta and Styela plicata), Pyuridae (Herdmania pallida and Microcosmus exasperatus). Didemnids were also collected in the area but were not considered here. Of the 17 species found, one (Polyclinum molle sp. nov.) is a new species in the family Polyclinidae and the others are all species with tropical distribution.

This review covers 25 years of progress on structural, functional, and developmental neurobiology of adult tunicates. The focus is on ascidians rather than pelagic species. The ascidian brain and peripheral nervous system are considered from the point of view of ultrastructure, neurotransmitters, regulatory peptides, and electrical activity. Sensory reception and effector control are stressed. Discussion of the dorsal strand plexus centres on its relationship with photoreceptors, the presence in it of gonadotropin-releasing hormone and its role in reproductive control. In addition to hydrodynamic sense organs based on primary sensory neurons (cupular organs), ascidians are now known to have coronal sense organs based on axonless hair cells resembling those of the vertebrate acustico-lateralis system. The peripheral nervous system is remarkable in that the motor neuron terminals are apparently interconnected synaptically, providing the equivalent of a nerve net. Development of the neural complex in ascidians is reviewed, highlighting recent embryological and molecular evidence for stomodeal, neurohypophyseal, and atrial placodes. The nervous system forms similarly during embryogenesis in the oozooid and blastogenesis in colonial forms. The regeneration of the brain in Ciona intestinalis (L., 1767) is discussed in relation to normal neurogenesis. Finally, the viviparous development of salps is considered, where recent work traces the early development of the brain, outgrowth of nerve roots, and the targetting of motor nerves to the appropriate muscles.

Six new polyaromatic alkaloids, lamellarin I (3), J (6), K (7), L (9), M (11) and the triacetate (13) of lamellarin N, and four known alkaloids, lamellarin A (I), B (15), C (16) and the triacetate (14) of lamellarin D, have been isolated from the marine ascidian Didemnum sp. The structures were deduced by high-field n.m.r. spectroscopy including 13C-1H shift-correlated two-dimensional (2D) n.m.r. experiments and n.O.e . measurements. The triacetates (8) and (10) of two of the compounds, lamellarin K (7) and lamellarin L (9), were dehydrogenated in high yield by treatment with 2,3-dichloro-5,6-dicyano-1,4-benzoquinone in ethanol heated under reflux. The products obtained were identical in all respects to the triacetates of lamellarin M and N, respectively. The re-isolation of lamellarins A-D, which were previously obtained from the prosobranch mollusc Lamellaria sp., lends further support for the idea that these molluscs sequester the compounds from ascidians as food sources.

Les tuniciers coloniaux peuvent générer un nouveau corps par reproduction asexuée et par la régénération entière du corps, deux formes de développement non-embryonnaire (DNE). Les différents modes de DNE sont définis en fonction de la nature des tissus organogénétiques. Curieusement, cette capacité est dispersée au sein du sous-phylum, qui contient des espèces capables de DNE (colonial) proches phylogénétiquement d’espèces ou les capacités régénératives sont absentes ou réduites (solitaire). Cela suggère que le DNE a été acquis et perdu plusieurs fois au sein du groupe. L’espèce coloniale Polyandrocarpa zorritensis semble avoir indépendamment acquis la capacité de DNE. Au cours de ma thèse, j’ai caractérisé le DNE dans cette espèce, en identifiant les étapes de DNE en conditions de laboratoire ainsi que les tissus et cellules mises en jeu. J’ai mis en évidence la participation des cellules mesenchymales et de l’épithélium vasculaire dans ce type de DNE. Ça n’a été pas décrit auparavant, et nous avons décidé de l’appeler ‘bourgeonnement vasale’. J’ai observé des cellules mesenchymales non-différenciées se regrouper et proliférer au point de régénération. J’ai décrit les cellules mesenchymales, en identifiant dans les cellules qui prolifèrent un morphotype non-différencié, les hémoblastes, aussi connues comme étant des cellules-souches putatives chez d’autres ascidies coloniales. De plus, j’ai défini la présence d’une étape de quiescence, la sphérule, dans le cycle de vie de P. zorritensis et j’ai caractérisé les variables environnementales et les mécanismes moléculaires mis en jeu dans la quiescence de cette espèce et dans une espèce éloignée, Clavelina lepadiformis
Colonial tunicates can generate a new adult body by asexual reproduction and whole body regeneration, two forms of non-embryonic development (NED). Different modes of NED are defined depending on the nature of the organogenetic tissues. Interestingly, this capacity is scattered across the sub-phylum, with species able of NED (colonial) closely related to species where regenerative capabilities are absent or reduced (solitary). This suggests that NED has been acquired or lost several times among the group. In recent phylogeny of family Styelidae, the colonial species Polyandrocarpa zorritensis seems to have acquired independently the capability of NED. During my PhD, I characterized the NED in this species, identifying the stages of NED under laboratory conditions and the tissues/cells involved. By histological and ultrastructural analyses, I highlighted the participation to NED of vascular epithelium and mesenchymal cells. This type of NED was undescribed before, and we decided to call it “vasal budding”. During the early stages of vasal budding I observed undifferentiated mesenchymal cells cluster and proliferate at the regenerative point; their distribution varies during vasal budding, increasing in the developing areas. I described the mesenchymal cells, identifying in the proliferating cells an undifferentiated morphotype, the hemoblasts, known as putative stem cells in other colonial ascidian. In addition, I defined the presence of a dormant stage, the spherule, in the life cycle of P. zorritensis and I characterized the environmental variable and the molecular mechanisms involved in dormancy in this species and in a distantly related species, Clavelina lepadiformis

Chez l’ascidie coloniale Botryllus schlosseri, en plus de l’embryogénèse existent deux voies de développement aboutissant à la production de la même structure : l’organisme adulte ou zooide. Ces développements alternatifs ont lieu lors de processus biologiques distincts : le bourgeonnement palléal (BP) et le bourgeonnement vasculaire (BV). Le BP est un processus de multiplication asexuée présentant une ontogénèse stéréotypée. En revanche, le BV est un phénomène régénératif, induit dans les vaisseaux sanguins de la colonie par l’ablation de tous les zooides et bourgeons palléaux. Mes travaux de recherche ont eu pour objectif de caractériser les bases moléculaires et cellulaires régissant le BP et le BV chez B. schlosseri. L’étude de gènes marqueurs des lignées méso-, endo- et ectodermiques a révélé l’existence de territoires présomptifs pour chacune de ces lignées, dès les premiers stades du BV et du BP, et suggéré l’existence d’un programme unique aux deux processus. Les lignées neurales et musculaires ont été étudiées plus en détail lors du BP, indiquant un double rôle potentiel, neuro- et myo-génétique, au tube dorsal, une structure jusqu’à présent uniquement associée au système nerveux. Une caractérisation morphologique poussée a mené à l’identification de stades précoces stéréotypés du BV lors de la régénération. Enfin, l’analyse de transcriptomes de différents stades du BP et de la régénération ont initié l’étude non biaisée des bases moléculaires du bourgeonnement chez Botryllus. L’objectif à long terme de ces travaux est de décrypter les bases moléculaires et génétiques facilitant, chez les métazoaires, l’évolution de voies de développement alternatives
In addition to embryogenesis, the colonial ascidians Botryllus schlosseri evolved two alternative developmental pathways leading to the same final structure: the adult body, or zooid. These non-embryonic ontogenesis occur during distinct biological processes: palleal budding (PB) and vascular budding (VB). PB is a process of asexual propagation, with a very stereotyped morphogenesis. Conversely, VB is a purely regenerative phenomenon, induced in the vascular system of the colony by the ablation of all zooids and palleal buds. My research work followed the objective to characterize the molecular and cellular basis of both PB and VB in B. schlosseri. The study of meso-, endo- and ectodermal lineage marker genes revealed the existence of presumptive territories of these lineages in the early palleal and vascular buds and that a single developmental program was launched in both VB and PB. Neural and muscle fates were studied in more detail for PB, indicating a potential double function, both neuro- and myo-genic for the dorsal tube, a structure so far associated with the nervous system only. A detailed morphological description of VB allowed to identify stereotyped stages during early regeneration. Eventually, a transcriptomic characterization of early VB and PB processes initiated an unbiased study of the molecular basis underlying the budding phenomenon in Botryllus. The overall goal of these research works is to unravel the molecular and genetic basis that facilitated, in Botryllus and globally in metazoan, the evolution of alternative developmental pathways

Chez l’ascidie coloniale Botryllus schlosseri, en plus de l’embryogénèse existent deux voies de développement aboutissant à la production de la même structure : l’organisme adulte ou zooide. Ces développements alternatifs ont lieu lors de processus biologiques distincts : le bourgeonnement palléal (BP) et le bourgeonnement vasculaire (BV). Le BP est un processus de multiplication asexuée présentant une ontogénèse stéréotypée. En revanche, le BV est un phénomène régénératif, induit dans les vaisseaux sanguins de la colonie par l’ablation de tous les zooides et bourgeons palléaux. Mes travaux de recherche ont eu pour objectif de caractériser les bases moléculaires et cellulaires régissant le BP et le BV chez B. schlosseri. L’étude de gènes marqueurs des lignées méso-, endo- et ectodermiques a révélé l’existence de territoires présomptifs pour chacune de ces lignées, dès les premiers stades du BV et du BP, et suggéré l’existence d’un programme unique aux deux processus. Les lignées neurales et musculaires ont été étudiées plus en détail lors du BP, indiquant un double rôle potentiel, neuro- et myo-génétique, au tube dorsal, une structure jusqu’à présent uniquement associée au système nerveux. Une caractérisation morphologique poussée a mené à l’identification de stades précoces stéréotypés du BV lors de la régénération. Enfin, l’analyse de transcriptomes de différents stades du BP et de la régénération ont initié l’étude non biaisée des bases moléculaires du bourgeonnement chez Botryllus. L’objectif à long terme de ces travaux est de décrypter les bases moléculaires et génétiques facilitant, chez les métazoaires, l’évolution de voies de développement alternatives
In addition to embryogenesis, the colonial ascidians Botryllus schlosseri evolved two alternative developmental pathways leading to the same final structure: the adult body, or zooid. These non-embryonic ontogenesis occur during distinct biological processes: palleal budding (PB) and vascular budding (VB). PB is a process of asexual propagation, with a very stereotyped morphogenesis. Conversely, VB is a purely regenerative phenomenon, induced in the vascular system of the colony by the ablation of all zooids and palleal buds. My research work followed the objective to characterize the molecular and cellular basis of both PB and VB in B. schlosseri. The study of meso-, endo- and ectodermal lineage marker genes revealed the existence of presumptive territories of these lineages in the early palleal and vascular buds and that a single developmental program was launched in both VB and PB. Neural and muscle fates were studied in more detail for PB, indicating a potential double function, both neuro- and myo-genic for the dorsal tube, a structure so far associated with the nervous system only. A detailed morphological description of VB allowed to identify stereotyped stages during early regeneration. Eventually, a transcriptomic characterization of early VB and PB processes initiated an unbiased study of the molecular basis underlying the budding phenomenon in Botryllus. The overall goal of these research works is to unravel the molecular and genetic basis that facilitated, in Botryllus and globally in metazoan, the evolution of alternative developmental pathways

Le but de mon projet était de décrire les molécules impliquées dans le développement asexué de l'ascidie coloniale Botryllus schlosseri et d'en déduire les mécanismes associés. Les ascidies appartiennent au sous embranchement des Tuniciers, groupe frère des Vertébrés. Elles sont les organismes les plus proches de l'humain capable de se reproduire de manière asexuée par bourgeonnement. Le travail a été réalisé sur l'ascidie coloniale Botryllus qui est composée de plusieurs individus (zoïde) réunis dans une tunique commune. Lors du développement asexué de Botryllus le zoïde peut sauter les stades embryonnaires et larvaires pour directement former un nouveau corps adulte incluant toute la musculature et le système nerveux en bourgeonnant. Dans le but d'étudier l'origine cellulaire et les mécanismes sous-jacents potentiels à la myogenèse non embryonnaire, j'ai suivi l'expression de ces gènes myogéniques lors du bourgeonnement de Botryllus et j'ai reconstruit la dynamique des précurseurs des muscles. Les orthologues de ces molécules ne sont pas exprimés seulement chez les ascidies mais également lors du développement du muscle cardio-pharyngé chez les Vertébrés et l'origine commune du cœur et des muscles pharyngés dans le champ cardio-pharyngé était déjà présente chez l'ancêtre commun des Tuniciers et des Vertébrés. En étudiant l'origine cellulaire lors de la neurogenèse j'ai pu observer que des gènes larvaires qui déterminent l'axe antérieur postérieur (AP) ont été cooptés dans une structure transitoire neuronale qui donne le ganglion cérébroïde. Ils divisent la structure en trois parties, évoquant ainsi de la tripartition de système nerveux des Vertébrés
The aim of this work is to describe via molecular-biological methods the asexual form of development of the colonial ascidian Botryllus schlosseri focusing on the formation of different tissues, namely muscles and nervous system, as well as exploring the potential presence of structures or cells homologous to the neural crest. Ascidians belong to the subphylum tunicates, sister group of vertebrates and are the closest relatives to man that can reproduce asexually, by budding. As colonial ascidian, the metamorphosis of Botryllus schlosseri specimen is followed by a lifelong, recurring, highly coordinated budding process, where multiple individuals (zooids) are connected and embedded in a common tunic. During asexual development, zooids can develop in a direct manner without embryonic and larval stages. To study the cellular origin and mechanisms of non-embryonic myogenesis I followed the expression pattern and dynamics of myogenic genes during asexual development and reconstructed muscle precursors. Orthologs of these genes are not only expressed during muscle formation via larval development but also during the formation of cardio-pharyngeal muscles in the vertebrate embryogenesis. I further drew a comparison of the regionalization of a transitory neurogenic structure, the dorsal tube, along the anteroposterior axis during budding with its larval counterpart, the neural tube, thus adding a RNA-expression profile of neural genes hereby proposing a scenario of cerebral ganglion formation by delamination. To better understand the nature of the dorsal tube and gangliogenesis I investigated potential involvement gene orthologs implemented in vertebrate neural crest formation

Orientador : A. Cecilia Z. Amaral
Dissertação (mestrado) - Universidade Estadual de Campinas, Instituto de Biologia
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Resumo: Apesar dos Ascidiacea constituirem um grupo muito comum nos costões rochosos do litoral de São Paulo, foram muito pouco estudados, principalmente no que se refere aos aspectos da biologia e ecologia das espécies. No Canal de São Sebastião existe uma fauna diversificada de ascídias coloniais que recobre a face inferior de pequenos blocos de rocha, tocas e frestas. O interesse em estudar alguns aspectos da biologia das espécies, suas interações na comunidade e a influência dos mesmos na colonização e manutenção do espaço foi o que levou à execução deste trabalho. O estudo do período reprodutivo foi feito através da imersão mensal de placas de recrutamento de lajota natural. O estudo da forma e velocidade de crescimento e da sucessão de ascídias coloniais, incluindo observações sobre interações entre as espécies, foi feito através do acompanhamento de placas artificiais durante 9 meses, imersas em duas épocas diferentes (verão e inverno). As cinco espécies estudadas em maior detalhe foram: Didemnum speciosum, Diplosoma macdonaldi, Clavelina oblonga, Symplegma viride e Botrylloides nigrum, pois eram as mais frequente no substrato artificial. Estas espécies apresentaram período reprodutivo longo com interrupção nos meses mais frios, observada em C. oblonga e B. nigrum. ...Observação: O resumo, na íntegra, poderá ser visualizado no texto completo da tese digital
Abstract: Although ascidians are very abundant on rocky shores of the Southeastern Brazilian coast (Sao Paulo State), there is little information about the species biology and ecology. In the São Sebastião Channel occurs a diversified fauna of compound ascidians, which cover the underside surface of boulders, crevices and cracks. To fulfill this gap of knowledge, the study of some life-history features of the ascidian species their interrelations in the community and how these aspects influence their colonization and space maintenance ability was undertaken. The reproductive period was determined by observations of one-month submerged artificial plates throughout one year, and by dissecting colonies to look for larvae. The ascidian community succession starting on clean plates were monthly observed on 9 month submerged plates, .on which the shape and growth rate of the colonies was also analyzed. Five species were discussed: Didemnum speciosum, Diplosoma macdonaldi, Clavelina oblonga, Symplegma viride e Botrylloides nigrum. They showed a long reprodutive period with a short interruption in C. oblonga and B. nigrum during winter. ...Note: The complete abstract is available with the full electronic digital thesis or dissertations
Mestrado
Mestre em Ecologia

Orientadores: Luiz Francisco Lembo Duarte, Vera Nisaka Solferini
Dissertação (mestrado) - Universidade Estadual de Campinas, Instituto de Biologia
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Mestrado
Ecologia
Mestre em Ecologia

The hermaphrodite, colonial ascidian Diplosoma listerianum (Chordata: Urochordata) mates by releasing sperm that disperse to neighbours, where they fertilise eggs that have been retained internally rather than spawned. The species is able to utilise highly dilute sperm: comparison with published information on a sea urchin, which released both eggs and sperm for external fertilisation, showed that D. listerianum maintained comparable levels of fertilisation at sperm concentrations two or three orders of magnitude lower than the echinoderm. Laboratory clones of D. listerianum displayed surprisingly high levels of sexual incompatibility. Fecundities of numerous pairwise crosses varied widely and suggested a continuous scale of computability. Although correlations of computability between reciprocal crosses appeared positive, considerable noise was present in the data and some crosses showed strongly asymmetrical compatibility. Patterns of sperm precedence with a five-day mating internal showed clear initial bias towards the first of two acting males. The proportion of second-male paternity (P₂) subsequently increased with time. Estimated P₂ for entire progeny arrays was consistently greater than 0.5, but varied widely. When mating interval was reduced, mate order effects appeared to be moderated. In competition with an alternative sperm source, acting males fathered more progeny if previously mated to a particular female than if no mating history existed, an advantage probably derived from fertilisations by stored sperm. When virgin acting female colonies were given mixtures of sperm at widely divergent concentrations, offspring were shared between the two sperm sources in approximately the ratio of each mixture. However, there existed a small but statistically significant deviation from the fair raffle model, in that sperm at the lower concentration consistently achieved a greater than expected share of paternity. Environmentally-determined fixed female preferences could be responsible for this negative frequency dependence ('rare male effect').

This dissertation examines two issues related to the ecological and evolutionary consequences of sexual and asexual reproduction in colonial marine invertebrates. The first two chapters explore the extent to which the planktonic larval phase limits the distribution and abundance of a colonial ascidian, Oiplosoma similis. The third chapter examines some of the fitness consequences of alterations in the pattern of asexual reproduction by colony fragmention in similis. All research was carried out on the fringing coral reef surrounding Coconut Island which is located in Kaneohe Bay, Oahu, Hawaii.
Typescript. Thesis (Ph. D.)--University of Hawaii at Manoa, 1989. Includes bibliographical references.

In the last years, several studies are addressed to the investigation of mechanisms that permitted the appearance and evolution of those structures considered as extremely important in the vertebrates radiation. The rise of vertebrates was accompanied by the acquisition of a great complexity in the structural plan of the organisms that is related to the evolution of features associated with the nervous system, such as neural crests, cranial placodes and an elaborated brain. The aim of the doctoral project is inserted in this line of research. Particularly, the attention is addressed to those characters, which in the non- vertebrate chordates can be interpreted as crucial for the subsequent evolution of the vertebrate body-plan. The starting point is represented by previous morphological studies that evidenced the presence, in the tunicate embryo, of transitory ectodermal and multipotential territories located at the neural plate border. Our research, carried out using different approaches, is focused on the characterisation and description of structures that differentiate from these domains. In this regard, it is investigated the organisation of the larval papillae and their formation from the rostral placode. These structures play a pivotal role in triggering the mechanisms and changes that characterise the metamorphosis, which in ascidians constitute the lost of the chordate body-plan of the larva and the begin of the sessile post-embryonic phase. The analysis of the sensory components in ascidians, finalised to the identification of homologies with the structures that derive form the placodes in vertebrates, is also extended to the coronal organ. The coronal organ is recently discovered and possesses morphological, positional and ultrastructural features that, together with the presence of hair cells, are comparable to the lateral line and inner ear of vertebrates, which components derive from the acoustic-lateral placodes. A substantial part of the work is dedicated to the investigation, with a molecular approach, of the presence of structures comparable to the neural placodes in ascidians. The attention is focused on the colonial ascidian Botryllus schlosseri that permits a comparative study on the mechanisms and genic networks involved in both embryogenetic and blastogenetic development. We characterised orthologues of placodal genes and constructed probes for in situ hybridisation experiments. During the stages of bud differentiation, we localised territories interested by expression of genes normally involved in the placodal induction and specification in vertebrates. These regions, for their position and differentiative potentialities, are comparable to other embryonic domains of B. schlosseri that are considered as homologues to the neural placodes. The larva of ascidians possesses a striated symmetrical musculature sharing some features with that of vertebrates and it flanks the dorsal tube and the notochord, representing a peculiar propriety of chordates. The acquisition of this new locomotory system probably required the parallel appearance of a sophisticated control of the coordination. The nervous system established new interactions and differentiated sensory structures that permitted the rapid perception of the environment by the mobile organism. At metamorphosis, the larval musculature is reabsorbed, while the cardiac and unstriated muscle fibres differentiate de novo from circulating mesenchymal cells. This plasticity constitutes the source of the evolutive potentiality of ascidians and thus we investigate its molecular and morphological bases. We isolated and characterised transcripts coding for muscle-specific genes, analysing the expression during the blastogenetic cycle of Botryllus, from the bud appearance to the adult regression. The use of different methods allowed the description of the organisation and differentiation of the unstriated muscle, confirming its unique proprieties. Taking together, our results contribute to the understanding of the origin and development of structures that represent an important starting point in the evolution and radiation of vertebrates.
Negli ultimi anni numerosi studi si sono rivolti all’approfondimento di quei meccanismi che hanno permesso la comparsa ed evoluzione di strutture ritenute di estrema importanza nella radiazione dei vertebrati. La comparsa dei vertebrati è stata accompagnata da un enorme balzo nella complessità del piano strutturale degli organismi, largamente ascrivibile all’evoluzione di strutture associate al sistema nervoso come le creste neurali, i placodi craniali ed un cervello elaborato. La tematica trattata durante lo svolgimento del progetto di dottorato si inserisce in questa attuale linea di ricerca. In particolare, l’attenzione è stata rivolta a quei caratteri che nei cordati non-vertebrati possono essere letti come cruciali per la successiva evoluzione del piano strutturale dei vertebrati. Il punto di partenza è rappresentato da precedenti studi morfologici che hanno evidenziato la presenza, nell’embrione dei tunicati, di territori ectodermici transitori e multipotenti localizzati al confine con la piastra neurale. Il nostro studio, svolto mediante l'utilizzo di vari approcci metodologici, si è rivolto alla caratterizzazione e descrizione delle strutture che queste aree sono in grado di differenziare. A questo proposito, è stata analizzata l’organizzazione delle papille larvali e la loro formazione a partire dal placode rostrale. Queste strutture giocano un ruolo primario nell’innescare i meccanismi e i cambiamenti che caratterizzano la metamorfosi, ovvero quel processo che nelle ascidie segna la perdita del piano corporeo da cordato della larva e il passaggio alla fase post-embrionale sessile. L’analisi riguardante strutture sensoriali presenti nelle ascidie, allo scopo di identificare eventuali omologie con le corrispondenti strutture derivanti dai placodi nei vertebrati, è stata poi estesa all’organo coronale. L’organo coronale è stato scoperto solo recentemente e presenta caratteristiche morfologiche generali, posizionali e ultrastrutturali tali, come la presenza di cellule capellute, che lo rendono comparabile alla linea laterale ed all’orecchio interno dei vertebrati, i cui componenti derivano dai placodi acustico-laterali. Una parte consistente del lavoro è stata dedicata all’indagine, da un punto di vista molecolare, della presenza di strutture accomunabile ai placodi neurali nelle ascidie. L’attenzione è stata rivolta all’ascidia coloniale Botryllus schlosseri, che permette di svolgere uno studio comparativo sui meccanismi e reti geniche che intervengono sia durante lo sviluppo embriogenetico che blastogenetico. Abbiamo caratterizzato specifici geni e prodotto sonde utilizzate in esperimenti di ibridazione in situ. Durante le fasi di differenziamento della gemma sono stati individuati specifici territori caratterizzati da espressione di alcuni geni normalmente coinvolti nell’induzione e specificazione placodale nei vertebrati. Grazie alla loro posizione e potenzialità differenziativa, queste stesse regioni sono apparse confrontabili con altri territori embrionali di B. schlosseri e di altre ascidie considerati omologhi a placodi neurali dei vertebrati. La larva delle ascidie presenta una muscolatura simmetrica striata, con caratteri comuni a quella dei vertebrati, la quale fiancheggia il tubo dorsale e la notocorda e che rappresenta una proprietà peculiare dei cordati. L’acquisizione di questo nuovo sistema locomotorio ha verosimilmente richiesto la comparsa parallela di un sofisticato sistema di controllo della coordinazione. Il sistema nervoso ha stabilito nuove interazioni e differenziato strutture sensoriali che hanno permesso all’organismo mobile la rapida percezione dell’ambiente circostante. Alla metamorfosi, la muscolatura larvale viene completamente riassorbita, mentre le fibre muscolari non striate della parete del corpo e quelle cardiache si differenziano de novo da cellule mesenchimali circolanti. Questa plasticità sta alla base della potenzialità evolutiva delle ascidie e quindi ne abbiamo indagato le basi molecolari e morfologiche. Abbiamo quindi isolato e caratterizzato trascritti e geni muscolo-specifici, studiandone l’espressione durante il ciclo blastogenetico di Botryllus, dalla comparsa della gemma alla regressione dell’adulto. L ’utilizzo di vari approcci ha permesso la descrizione dell’organizzazione e differenziamento della muscolatura non striata, confermandone le caratteristiche uniche. Nel complesso i diversi risultati rappresentano contributi significativi per la conoscenza delle origini e sviluppo quelle strutture che hanno rappresentato un punto di partenza importante nell’evoluzione e radiazione dei vertebrati.

AbstractAscidians are sessile tunicates, that is, marine animals belonging to the phylum Chordata and considered the sister group of vertebrates. They are widespread in all the seas, constituting abundant communities in various ecosystems. Among chordates, only tunicates are able to reproduce asexually, forming colonies. The high regenerative potentialities enabling tunicates to regenerate damaged body parts, or the whole body, represent a peculiarity of this taxon. Here we review the methodological approaches used in more than a century of biological studies to induce regeneration in both solitary and colonial species. For solitary species, we refer to the regeneration of single organs or body parts (e.g., siphon, brain, gonad, tunic, viscera). For colonial species, we review a plethora of experiments regarding the surgical manipulation of colonies, the regeneration of isolated colonial entities, such as single buds in the tunic, or part of tunic and its circulatory system.

AbstractThe decrease in sequencing costs and technology improvements has led to the adoption of RNA-sequencing to profile transcriptomes from further non-traditional regeneration model organisms such as the colonial ascidian Botrylloides leachii. The relatively unbiased way in which transcripts are identified and quantified makes this technique suitable to detect large-scale changes in expression, and the identification of novel transcripts and isoforms. Of particular interest to many researchers is the discovery of differentially expressed transcripts across different treatment conditions or stages of regeneration. This protocol describes a workflow starting from processing raw sequencing reads, mapping reads, assembly of transcripts, and measuring their abundance, creating lists of differentially expressed genes and their biological interpretation using gene ontologies. All programs used in this protocol are open-source software tools and freely available.

Abstract The three thaliacean groups, pyrosomas, salps and doliolids, differ from each other not only in their anatomy, but also in their locomotion, buoyancy, sensory systems and embryonic development. They share, however, metagenetic life-cycles involving alternation of generations (although one generation is only transient in pyrosomas) and essentially similar stolonic budding. Traditionally grouped together, a grouping retained here for convenience, it now seems probable that the Thaliacea are polyphyletic, and thus that their grouping together is artificial (Chapter 16).Pyrosomas are colonial animals, primarily warmwater forms, rather common in tropical and warm temperate waters except in the Red Sea. They are mainly caught in the epipelagic and upper mesopelagic layers, although specimens have occasionally been observed at greater depths (Tregouboff, 1956). In contrast to the other pelagic Tunicates, pyrosomas consist of permanent tubular hollow colonies; the blastozooids resulting from asexual propagation are essentially independent but remain embedded side by side in a common tunic or test, as in the aplousobranchiate Ascidians. These blastozooids are hermaphrodite and blastogenic and are responsible for the propagation of the species and the growth of the colony, whilst the oozooid is only a shortlived transient and blastogenic stage.