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BackgroundFunctional inference on the attachment of acanthocephalans has generally been drawn directly from morphology. However, performance of structures is often non-intuitive and context-dependent, thus performance analysis should be included whenever possible to improve functional interpretation. In acanthocephalans, performance analysis of attachment is available only for Acanthocephalus ranae, a species that solely relies on the proboscis to attach. Here we compare body morphology and muscle arrangement in 13 species of Corynosoma, which use their spiny body as a fundamental holdfast. A basic performance analysis using live cystacanths of two representative species is also provided.MethodsAdults of 13 Corynosoma spp. were obtained from 11 marine mammal species. Specimens were cut and carefully cleaned to examine muscle arrangement through light and scanning electron microscopy. Live cystacanths of C. australe and C. cetaceum were selected for performance analysis. Video records of evagination-invagination cycles of the proboscis were obtained and analysed with a video editor.ResultsThe basic arrangement of proboscis retractors, trunk circular and longitudinal muscles, neck retractors and receptacle retractors, was conserved in all Corynosoma species. Interspecific variability was found in the relative development of disk muscles: minimum in C. enhydri, maximum in C. cetaceum; the distal insertion of the ventral neck retractor: ventro-lateral in C. cetaceum, C. hamannni and C. pseudohamanni and ventral in the other species; and the distal insertion of the receptacle retractors: more proximal in species with a longer hindtrunk. Performance analysis indicated striking similarities to that described for A. ranae except that (i) the foretrunk bends ventrally during the evagination-invagination cycles of the proboscis; (ii) disk muscles can flatten the tip of the foretrunk regardless of these cycles; and (iii) the receptacle bends ventrally and is driven to the hindtrunk by coordinated action of receptacle retractors.ConclusionsSpecies of Corynosoma are able to use up to six holfast mechanisms. Attachment relies on a similar performance to that described for A. ranae. However, structural ventral bending of an inflated, spiny foretrunk, with a parallel re-arrangement of foretrunk muscles, have generated unexpected novel functions that make attachment extremely effective in species of Corynosoma. Interspecific variability in trunk shape and muscle arrangement grossly correlates with the rheological conditions each species experiences in their microhabitats within the gut of marine mammals.Electronic supplementary materialThe online version of this article (10.1186/s13071-018-3165-1) contains supplementary material, which is available to authorized users.
BackgroundFunctional inference on the attachment of acanthocephalans has generally been drawn directly from morphology. However, performance of structures is often non-intuitive and context-dependent, thus performance analysis should be included whenever possible to improve functional interpretation. In acanthocephalans, performance analysis of attachment is available only for Acanthocephalus ranae, a species that solely relies on the proboscis to attach. Here we compare body morphology and muscle arrangement in 13 species of Corynosoma, which use their spiny body as a fundamental holdfast. A basic performance analysis using live cystacanths of two representative species is also provided.MethodsAdults of 13 Corynosoma spp. were obtained from 11 marine mammal species. Specimens were cut and carefully cleaned to examine muscle arrangement through light and scanning electron microscopy. Live cystacanths of C. australe and C. cetaceum were selected for performance analysis. Video records of evagination-invagination cycles of the proboscis were obtained and analysed with a video editor.ResultsThe basic arrangement of proboscis retractors, trunk circular and longitudinal muscles, neck retractors and receptacle retractors, was conserved in all Corynosoma species. Interspecific variability was found in the relative development of disk muscles: minimum in C. enhydri, maximum in C. cetaceum; the distal insertion of the ventral neck retractor: ventro-lateral in C. cetaceum, C. hamannni and C. pseudohamanni and ventral in the other species; and the distal insertion of the receptacle retractors: more proximal in species with a longer hindtrunk. Performance analysis indicated striking similarities to that described for A. ranae except that (i) the foretrunk bends ventrally during the evagination-invagination cycles of the proboscis; (ii) disk muscles can flatten the tip of the foretrunk regardless of these cycles; and (iii) the receptacle bends ventrally and is driven to the hindtrunk by coordinated action of receptacle retractors.ConclusionsSpecies of Corynosoma are able to use up to six holfast mechanisms. Attachment relies on a similar performance to that described for A. ranae. However, structural ventral bending of an inflated, spiny foretrunk, with a parallel re-arrangement of foretrunk muscles, have generated unexpected novel functions that make attachment extremely effective in species of Corynosoma. Interspecific variability in trunk shape and muscle arrangement grossly correlates with the rheological conditions each species experiences in their microhabitats within the gut of marine mammals.Electronic supplementary materialThe online version of this article (10.1186/s13071-018-3165-1) contains supplementary material, which is available to authorized users.
Scanning electron microscope (SEM) investigations on the holdfast elements, proboscis hooks, and trunk spines of Dentitruncus truttae (Acanthocephala, Palaeacanthocephala), an endoparasite of Salmo trutta (brown trout), provide more data about the surface of these taxonomic relevant structures. In both acanthocephalan sexes, the fully everted cylindrical proboscis possessed 18 longitudinal rows of hooks with 18 hooks per row (rarely 19-20). Hook length varied according to position on the proboscis; apical hooks were 40-52 microm long, middle hooks were 31.7-36.6 microm, and basal hooks were 38.1-40 microm. Starting from the anterior end of the metasoma, numerous cuticular spines (26.7-30 microm in length) were visible and their number progressively decreased posteriorly. SEM observations of D. truttae hooks and spines revealed the presence of many surface striations on each proboscis hook. These surface striations were absent from trunk spines. From the base of the hook, the striations ran parallel toward the point of convergence. Additionally, survey of longitudinal and transversal sections of the hook using transmission electron microscope confirmed that the hook surface was not smooth. SEM comparison with the hooks of several palaeacanthocephalan species, as well as with the hooks of species belonging to Eoacanthocephala and Polyacanthocephala, indicated that the striations are currently exclusive to D. truttae proboscis hooks.
Adults of the acanthocephalan Corynosoma cetaceum deeply attach to the stomach of dolphins using the proboscis and its spiny foretrunk as a disk while the spiny hindtrunk bends to also embed its ventral spines. During deep attachment, two ventral folds of tegument, anterior and posterior, are created. Spine growth is inhibited to a variable degree in folds, generating an extraordinary phenotypic variability, with most individuals, especially females, having folds partially or totally devoid of spines. Little is known on how this variability is generated and why it is not apparently found in other Corynosoma spp. In this paper, we examined the trunk armature of 77 and 388 cystacanth larvae of C. cetaceum and C. australe, respectively, from teleosts, and over 8800 adult specimens of C. australe, C. bullosum, C. cetaceum, C. strumosum, C. villosum and C. wegeneri from marine mammals. Cystacanths and adults of C. cetaceum exhibited the same range of fold spine reduction and variability, suggesting that they are generated prior to the adult stage (i.e., before spines are functional) and do not result from phenotypic plasticity. The other Corynosoma species analyzed created only the anterior fold during deep attachment, but it was always spined. Females of C. cetaceum had significantly larger foretrunk and hindtrunk spines than the other species and likely suffer stronger fold compression during deep attachment. The exceptional colonization of a harsh microhabitat, the stomach, could have generated a trade-off in C. cetaceum, which must bend the trunk to attach (as other Corynosoma spp.) but must also produce large spines that, in the folds, presumably are maladaptive and must be reduced.
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