Post-mortem oxygen isotope exchange within cultured diatom silica.

RATIONALE: Potential post-mortem alteration to the oxygen isotope composition of biogenic silica is critical to the validity of palaeoclimate reconstructions based on oxygen isotope ratios (δ18 O values) from sedimentary silica. We calculate the degree of oxygen isotope alteration within freshly cultured diatom biogenic silica in response to heating and storing in the laboratory. METHODS: The experiments used freshly cultured diatom silica. Silica samples were either stored in water or dried at temperatures between 20 °C and 80 °C. The mass of affected oxygen and the associated silica-water isotope fractionation during alteration were calculated by conducting parallel experiments using endmember waters with δ18 O values of -6.3 to -5.9 ‰ and -36.3 to -35.0 ‰. Dehydroxylation and subsequent oxygen liberation were achieved by stepwise fluorination with BrF5 . The 18 O/16 O ratios were measured using a ThermoFinnigan MAT 253 isotope ratio mass spectrometer. RESULTS: Significant alterations in silica δ18 O values were observed, most notably an increase in the δ18 O values following drying at 40-80 °C. Storage in water for 7 days between 20 and 80 °C also led to significant alteration in δ18 O values. Mass balance calculations suggest that the amount of affected oxygen is positively correlated with temperature. The estimated oxygen isotope fractionation during alteration is an inverse function of temperature, consistent with the extrapolation of models for high-temperature silica-water oxygen isotope fractionation. CONCLUSIONS: Routinely used preparatory methods may impart significant alterations to the δ18 O values of biogenic silica, particularly when dealing with modern cultured or field-collected material. The significance of such processes within natural aquatic environments is uncertain; however, there is potential that similar processes also affect sedimentary diatoms, with implications for the interpretation of biogenic silica-hosted δ18 O palaeoclimate records.

Diatoms can be an important exception to temperature-size rules at species and community levels of organization.

Climate warming has been linked to an apparent general decrease in body sizes of ectotherms, both across and within taxa, especially in aquatic systems. Smaller body size in warmer geographical regions has also been widely observed. Since body size is a fundamental determinant of many biological attributes, climate-warming-related changes in size could ripple across multiple levels of ecological organization. Some recent studies have questioned the ubiquity of temperature-size rules, however, and certain widespread and abundant taxa, such as diatoms, may be important exceptions. We tested the hypothesis that diatoms are smaller at warmer temperatures using a system of geothermally heated streams. There was no consistent relationship between size and temperature at either the population or community level. These field data provide important counterexamples to both James' and Bergmann's temperature-size rules, respectively, undermining the widely held assumption that warming favours the small. This study provides compelling new evidence that diatoms are an important exception to temperature-size rules for three reasons: (i) we use many more species than prior work; (ii) we examine both community and species levels of organization simultaneously; (iii) we work in a natural system with a wide temperature gradient but minimal variation in other factors, to achieve robust tests of hypotheses without relying on laboratory setups, which have limited realism. In addition, we show that interspecific effects were a bigger contributor to whole-community size differences, and are probably more ecologically important than more commonly studied intraspecific effects. These findings highlight the need for multispecies approaches in future studies of climate warming and body size.

A time-calibrated multi-gene phylogeny of the diatom genus Pinnularia.

Pinnularia is an ecologically important and species-rich genus of freshwater diatoms (Bacillariophyceae) showing considerable variation in frustule morphology. Interspecific evolutionary relationships were inferred for 36 Pinnularia taxa using a five-locus dataset. A range of fossil taxa, including newly discovered Middle Eocene forms of Pinnularia, was used to calibrate a relaxed molecular clock analysis and investigate temporal aspects of the genus' diversification. The multi-gene approach resulted in a well-resolved phylogeny of three major clades and several subclades that were frequently, but not universally, delimited by valve morphology. The genus Caloneis was not recovered as monophyletic, confirming that, as currently delimited, this genus is not evolutionarily meaningful and should be merged with Pinnularia. The Pinnularia-Caloneis complex is estimated to have diverged between the Upper Cretaceous and the early Eocene, implying a ghost range of at least 10 million year (Ma) in the fossil record.

Haslea silbo, A Novel Cosmopolitan Species of Blue Diatoms.

Specimens of a new species of blue diatoms from the genus Haslea Simonsen were discovered in geographically distant sampling sites, first in the Canary Archipelago, then North Carolina, Gulf of Naples, the Croatian South Adriatic Sea, and Turkish coast of the Eastern Mediterranean Sea. An exhaustive characterization of these specimens, using a combined morphological and genomic approach led to the conclusion that they belong to a single new to science cosmopolitan species, Haslea silbo sp. nov. A preliminary characterization of its blue pigment shows similarities to marennine produced by Haslea ostrearia, as evidenced by UV-visible spectrophotometry and Raman spectrometry. Life cycle stages including auxosporulation were also observed, providing data on the cardinal points of this species. For the two most geographically distant populations (North Carolina and East Mediterranean), complete mitochondrial and plastid genomes were sequenced. The mitogenomes of both strains share a rare atp6 pseudogene, but the number, nature, and positions of the group II introns inside its cox1 gene differ between the two populations. There are also two pairs of genes fused in single ORFs. The plastid genomes are characterized by large regions of recombination with plasmid DNA, which are in both cases located between the ycf35 and psbA genes, but whose content differs between the strains. The two sequenced strains hosts three plasmids coding for putative serine recombinase protein whose sequences are compared, and four out of six of these plasmids were highly conserved.

Nature's Batik: a computer evolution model of diatom valve morphogenesis.

This paper describes a novel computer simulation that uses evolution to design functioning raphid pennate diatom valves. The model of valve morphogenesis used is based on current theories that highlight the importance of cytoskeletal elements in valve development. An "organic" negative imprint is grown in a grid-based system, using both local and global rules to dictate grid cell states. Silica then diffuses out into all remaining grid cells. This model is shown to generate raphid pennate diatom valves capable of functioning as cell walls. At every stage of development the generated valves are consistent with observations of real diatom valve growth. This model of diatom valve morphogenesis is interestingly similar to the negative technique used by artists in batik painting.

ONTOGENY, HOMOLOGY, AND TERMINOLOGY-WALL MORPHOGENESIS AS AN AID TO CHARACTER RECOGNITION AND CHARACTER STATE DEFINITION FOR PENNATE DIATOM SYSTEMATICS(1).

This article reviews current knowledge of wall morphogenesis in pennate diatoms in relation to the way characters are defined and described for taxonomic and systematic analyses. It argues that an understanding of ontogeny is essential for the accurate identification of character homologies, which in turn must underpin all phylogenetic and systematic analyses. Terminology should reflect character homology, but most diatom terminology fails to do this, with concomitant confusion and potential taxonomic mistakes. Identifying where information is lacking or misinterpreted are first steps toward improving our understanding of diatom structure and relationships. After reviewing the current knowledge on pennate diatom structure and its development, this article briefly discusses the significance of morphological variation, character polarity, and the vital importance of applying diatom terminology correctly.

DIATOM COLONY FORMATION: A COMPUTATIONAL STUDY PREDICTS A SINGLE MECHANISM CAN PRODUCE BOTH LINKAGE AND SEPARATION VALVES DUE TO AN ENVIRONMENTAL SWITCH(1).

The morphological plasticity and adaptive behavior exhibited during diatom colony formation in Aulacoseira is explored through computer simulation to study how the interplay of mechanisms such as cytoskeletal-driven membrane protrusions, silica deposition, and environmental factors may contribute to the generation of two distinct spine morphologies on linkage and separation valves. A multiscale agent-based computational model was developed, which showed that a single cytoskeleton-driven, competitive growth mechanism could generate either of the two characteristic phenotypes, given only a single switch in the environment (as might be experienced by a change in light regime). Hypotheses are formulated from the model, and predictions made for potential follow-up experiments.

Integrated simulation with experimentation is a powerful tool for understanding diatom valve morphogenesis.

We present a number of promising examples of computational studies, which improve our understanding of the morphogenesis process of diatom cell walls. Each example considers a different physical scenario whereby computational and mathematical models are used to evaluate hypotheses pertaining to diatom valve formation; considering the roles of cytoskeletal elements, interactions between cell components that might generate patterned structures from the submicron (nanoscale) to cell level, and the effect of environmental variables. We propose that the complex, multiscale phenomenon of diatom valve morphogenesis requires better integration of computational/mathematical and experimental procedures if we are to untangle all the contributing processes. Finally we outline a plan for future directions, to achieve this integration and further the field of diatom morphogenesis research.