The Functional and Evolutionary Significance of the Ultrastructure of the Ophryoscolecidae (Order Entodiniomorphida)1

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Abstract
Ophryoscolecid ciliates from the order Entodiniomorphida show a series of morphological types which have been interpreted as an evolutionary lineage. In this study, ultrastructural information from three species—the ancestral Entodinium, intermediate Eudiplodinium maggii, and the advanced Epidinium—has been evaluated in terms of ophryoscolecid evolution. The infraciliature, nuclei, contractile vacuoles, cortex, and cytoplasm are all very similar and sometimes indistinguishable among the three species, suggesting a close relationship. The cytoalimentary system, however, shows considerable interspecific variation. The cytopharynx differs in position and extent of microtubules and microfilaments, varying in appearance from a microtubular to a microfilament‐based mechanism while retaining similar component structures. The esophagus, a zone of cytoplasm extending from the cytopharynx and delimited by a microtubular/fibrillar wall, is rudimentary in Entodinium, sac‐like in Eudiplodinium, and tube‐like in Epidinium, where it also has convoluted walls and a sheath of fibrous material that suggest an expansible‐contractile structure. These variations have been related to type of food particle ingested. The capability of the cytoalimentary systems seems to be increased so that the more advanced forms can exploit a food resource, in the form of large plant fragments in the ruminal fluid, not available to the simpler, ancestral forms, which tend to ingest small particles such as bacteria and starch grains. The original evolutionary lineage based on morphological studies using light microscopy is supported by our observations, in these three forms, of ultrastructural variations in the cytopharynx and in their relationship with diet via possible ingestion mechanisms. Additional support for this evolutionary analysis comes from preliminary studies of other ophryoscolecids in which the cytoalimentary organization is consistent with their positions relative to one another in the evolutionary scheme.