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What was that? Unraveling a 400-million-year-old mystery
Graham and her colleagues hypothesized that Prototaxites fossils may be composed of partially degraded wind-, gravity-, or water-rolled mats of mixotrophic (capable of deriving energy from multiple sources) liverworts that are associated with fungi and cyanobacteria. This situation resembles the mats produced by the modern liverwort genus Marchantia. The authors tested their hypothesis by treating Marchantia polymorpha in a manner to reflect the volcanically-influenced, warm environments typical of the Devonian period and compared the resulting remains to Prototaxites fossils. Graham and her colleagues investigated the mixotrophic ability of M. polymorpha by assessing whether M. polymorpha grown in a glucose-based medium is capable of acquiring carbon from its substrate.
"For our structural comparative work," Graham said, "we were extremely fortunate to have an amazing thin slice of the rocky fossil, made in 1954 by the eminent paleobotanist Chester A. Arnold."
Their structural and physiological studies showed that the fossil Prototaxites and the modern liverwort Marchantia have many similarities in their external structure, internal anatomy, and nutrition. Despite being subjected to conditions that would promote decomposition and desiccation, the rhizoids of M. polymorpha survived degradation, and with the mat rolled, created the appearance of concentric circles. The fungal hyphae associated with living liverworts also survived treatment, suggesting that the branched tubes in fossils may be fungal hyphae. The very narrow tubes in the fossils resemble filamentous cyanobacteria that the researchers found wrapped around the rhizoids of the decaying M. polymorpha.
"We were really excited when we saw how similar the ultrastructure of our liverwort rhizoid walls was to images of Prototaxites tubes published in 1976 by Rudy Schmid," Graham said.
In their investigations into the nutritional requirements of M. polymorpha, Graham and her colleagues found that the growth of M. polymorpha in a glucose-based medium was approximately 13 times that seen when the liverwort was grown in a medium without glucose. Stable carbon isotope analyses indicated that less than 20% of the carbon in the glucose-grown liverwort came from the atmosphere. The stable carbon isotope values obtained from M. polymorpha grown with varying amounts of cyanobacteria present span the range of values reported for Prototaxites fossils. Taken together, these results demonstrate that the liverworts have a capacity for mixotrophic nutrition when glucose is present and that mixotrophy and/or the presence of cyanobacteria could be responsible for the stable carbon isotope values obtained from Prototaxites.
Graham and her colleagues' results demonstrate that liverworts were important components of Devonian ecosystems. Their results support previous hypotheses that microbial associations and mixotrophy are ancient plant traits, rather than ones that have evolved recently.
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