On the experimental silicification of microorganisms II. On the time of appearance of eukaryotic organisms in the fossil record

Abstract On the basis of ultrastructural, biochemical and genetic studies, bacteria and blue green algae (Kingdom Monera, all prokaryotes) differ unambiguously from the eukaryotic organisms (Fungi, plants sensu stricto ) and protists or protoctists, (Copeland, 1956). The gap between eukaryotes and prokaryotes is recognized as the most profound evolutionary discontinuity in the living world. This gap is reflected in the fossil record. Fossil remains of Archaean and Proterozoic Aeons primarily consist of prokaryotes and the Phanerozoic is overwhelmingly characterized by fossils of the megascopic eukaryotic groups, both metazoa and metaphyta. Based on the morphological interpretation of microscopic objects structurally preserved in Precambrian cherts, the time of appearance of remains of eukaryotic organisms in the fossil record has been claimed to be as early as 2.7 · 10 9 years ago, (Kaźmierczak, 1976). Others suggest chronologies varying between 1.7 to 1.3 · 10 9 (Schopf et al., 1973) or a time approaching 1.3 · 10 9 years (Cloud, 1974). There is general agreement that many of the Ediacaran faunas, which have been dated at about 680 m.y. are fossils of megascopic soft-bodied invertebrate animals. Since all invertebrates are eukaryotic, the ca. 680 m.y. date for deposition of these animal assemblages may represent the earliest appearance of eukaryotic organisms. But the question remains as to whether there is definitive evidence for eukaryotic cells before this “benchmark” of the late Precambrian. An excellent discussion of this particular problem as especially relating to acritarchs extending from rocks of Upper Riphean through Vendian and into the basal Cambrian is presented in recent studies by Vidal (1974, 1976) in Late Precambrian microfossils from the Visingso rocks of southern Sweden. Previous work on the laboratory silicification of wood and algal mat communities (Leo and Barghoorn, 1976) suggested that further analysis of “artificial fossils” might be of aid in the interpretation of fossil morphology toward the ultimate solution of this problem. Thus the procedure developed by one of us (ESB) for laboratory wood silicification was adapted to various smaller objects. By successive immersions of wet cellular aggregates, colonies of various organisms and abiotic organic microspheres in tetraethyl orthosilicate, silicified cells and structures are produced which bear an interesting resemblance to ancient chert-embedded microfossils. Our observation of these microorganisms and proteinoid microspheres silicified in the laboratory as well as of degrading microorganisms, both eukaryotic and prokaryotic, have led us to conclude that many, if not all, of the criteria for assessing fossil eukaryotic microorganisms are subject to serious criticism in interpretation. We studied a large variety of prokaryotic algae, some eukaryotic algae, fungi, protozoa, and abiotic organic microspheres stable at essentially neutral pH. In some cases, degradation and/or silicification systematically altered both size and appearances of microorganisms. By the use of monoalgal cultures of blue-green algae, features resembling nuclei, chloroplasts, tetrads, pyrenoids, and large cell size may be simulated. In many cases individual members of these cultures show so much variation that they may be mistaken as belonging to more than one species. The size ranges for silicified prokaryotic and eukaryotic algae overlap. Several prokaryotes routinely yielded spherical or filamentous structures that resembled large cells. Because of genuine large sizes (e.g., Prochloron ), shrinkage, systematic alteration or congregation of unicells to form other structures we find sizes to be of very limited use in determining whether an organism of simple morphology was prokaryotic or eukaryotic. Although some “prebiotic proteinoid microspheres” (of Fox and Harada, 1960) are impossible to silicify with our laboratory methods, those stable at neutral pH (Hsu and Fox, 1976) formed spherical objects that morphologically resemble silicified algae or fungal spores. Many had internal structure. We conclude that even careful morphometric studies of fossil microorganisms are subject to many sources of misinterpretation. Even though it is a logical deduction that eukaryotic microorganisms evolved before Ediacaran time there is no compelling evidence for fossil eukaryotes prior to the late Precambrian metazoans.