Some of the most remarkable fossils preserve cellular details of soft tissues. In many of these, the tissues have been replaced by calcium phosphate. This process has been assumed to require elevated concentrations of phosphate in sediment pore waters. In decay experiments modern shrimps became partially mineralized in amorphous calcium phosphate, preserving cellular details of muscle tissue, particularly in a system closed to oxygen. The source for the formation of calcium phosphate was the shrimp itself. Mineralization, which was accompanied by a drop in pH, commenced within 2 weeks and increased in extent for at least 4 to 8 weeks. This mechanism halts the normal loss of detail of soft-tissue morphology before fossilization. Similar closed conditions would prevail where organisms are rapidly overgrown by microbial mats.
A series of experiments was carried out to investigate the nature and controls (oxygen, microbial populations, agitation) on the degradation of soft tissues. Decay was monitored in terms of morphological change, weight loss, and change in chemical composition in the polychaete Nereis virens. Polychaetes include a range of tissue types of differing chemical composition and preservation potential: muscle, cuticle, setae, and jaws. Regardless of conditions, all the muscle had broken down and fluid loss through the ruptured cuticle had reduced the carcass to two dimensions within 8 days at 20°C. In most cases some cuticle, in addition to the jaws and setae, remained after 30 days. Where oxygen was completely eliminated, the rate of decay of the more volatile issues was significantly reduced. The degree of both osmotic uptake of water by the carcass and changes in water pH differed depending on whether the system was open or closed to oxygen diffusion. Autolytic and chemical processes are not sufficient to fully degrade the carcass in the absence of bacteria. Where internal bacteria are present, the presence or absence of water column bacteria made little difference to decay rate. Initial degradation (in the first 3 days) affects mainly the lipid fraction and the collagen of the cuticle. Later decay reduces the nonsoluble protein and increases the relative proportion of refractory structural components (tanned chitin and collagen) to more than 95% by day 30. Thus, only the sclerotized tissues are likely to survive beyond 30 days in the absence of early diagenetic mineralization. The sequence of degradation predicted from the relative decay resistance of macromolecules in the sedimentary record (protein → carbohydrate → lipid) is not, therefore, a consistent indicator of the preservation potential of structural tissues which incorporate them.The experiments reveal five stages in the decay of polychaete carcasses; whole/shriveled, flaccid, unsupported gut, cuticle sac, jaws and setae. All are represented in the fossil record. This allows an estimation of how far decay proceeded before it was halted by the fossilization process. The most complete preservations occur in the Cambrian where the Burgess Shale preserves evidence of muscle tissues. Traces of the gut and cuticle are more widely preserved, as at Mazon Creek, Grès à Voltzia, Solnhofen, and Hakel. Preservation varies within Konservat-Lagerstätten. The most common whole body preservation includes only the more recalcitrant tissues, jaws (where present) and setae, with an impression of the body outline. The stage of decay can be used as a taphonomic threshold, to provide an indication of how significantly the diversity of an exceptionally preserved biota is likely to have been reduced by taphonomic loss.
Although the graptolites lacked biomineralised tissue, their skeletons are abundantly preserved in deeper-water mudstones. Decay experiments and observations on the closely related living hemichordate Rhabdopleura demonstrate that the periderm and stolon are highly resistant to decay, remaining intact for mouths, whereas the zooids are unrecognizable within days. Curie-point-gas-chromatography (Py-GC) and Curie-point-gas-chromatography-mass spectrometry (Pt-GC-MS) of the periderm of Rhabdopleura confirms that proteinaceous organic matter is a major constituent. Ultrastructurally preserved graptolite periderm, on the other hand, is a highly altered kerogen-like substance rich in aliphatic biomacromolecules. The composition of the preserved graptolite periderm reflects diagenetic replacement by components probably mainly derived from algal cell walls. -from Author
Decay experiments on the cephalochordate Branchiostoma lanceolatum (‘amphioxus’) demonstrate that the most decay resistant structures are the notochord sheath and the cartilaginous rods which support the gill bars. However, even more labile soft parts, such as the muscles and skin may survive for at least 124 days under totally anoxic conditions. As the chevron‐shaped muscles of the myomeres shrink and collapse, those on opposite sides of the trunk maybe displaced, resulting in pronounced offsetting. Only 1.42% of the initial dry weight of Branchiostoma is resistant to alkali and acid hydrolysis, compared to 46% in the polychaete Nereis virens. Branchiostoma is only likely to be fossilized as a result of decay inhibition and replication by early diagenetic minerals. The results of these experiments cast light on the interpretation of a number of primitive fossil chordates. There is no reason to infer extracellular decay‐resistant cuticle in the Burgess Shale Pikaia. The axial lies preserved in the conodont animal specimens from the Carboniferous of Edinburgh, Scotland, represent the notochord. The displacement of the elements to one side of the head reflects the true position of the apparatus ‐ the surrounding tissue has been lost through decay. The chevron‐shaped structures in the Carboniferous chordate Conopiscius are the muscles of the myomeres, not external scales. The lines delineating the segments in the Silurian Jamoytius most likely represent the myosepta. There is some doubt about the nature of the only specimen interpreted as a fossil cephalochordate, Palaeobranchiostoma hamatotergum from the Permian of South Africa. □Taphonomy, decay, softparts, Cephalochordata, Branchiostoma, lancelet, Chordata, Pikaia, conodont, Conopiscius, Jamoytius, Palaeobranchiostoma.
The functional morphology of the buccal mass of 23 species of cephalopod (Octopoda, 4 species; Teuthoidea, 17; Sepioidea, 2) was investigated by gross dissection, histology and observations on fresh preparations. Cephalopod beaks lack a joint or articulation point. The jaws slide and rotate around an area rather than a fixed point. During closing the superior mandibular muscle (SMM) provides the force of a bite and the largest movement vector, whilst the inferior mandibular muscle (IMM) acts to retract the upper beak, causing shearing action. Dorsal portions of the lateral mandibular muscles (LMM) flex the upper beak walls outwards, probably to accommodate the backwards sweep of the radula and buccal palps during closing. To open the beaks, the ventral portions of the lateral mandibular muscles pull the rear lateral walls of the two beaks towards each other, moving the lower beak back relative to the upper.
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