Possible origins of macroscopic left-right asymmetry in organisms

I consider the microscopic mechanisms by which a particular left-right (L/R) asymmetry is generated at the organism level from the microscopic handedness of cytoskeletal molecules. In light of a fundamental symmetry principle, the typical pattern-formation mechanisms of diffusion plus regulation cannot implement the "right-hand rule"; at the microscopic level, the cell's cytoskeleton of chiral filaments seems always to be involved, usually in collective states driven by polymerization forces or molecular motors. It seems particularly easy for handedness to emerge in a shear or rotation in the background of an effectively two-dimensional system, such as the cell membrane or a layer of cells, as this requires no pre-existing axis apart from the layer normal. I detail a scenario involving actin/myosin layers in snails and in C. elegans, and also one about the microtubule layer in plant cells. I also survey the other examples that I am aware of, such as the emergence of handedness such as the emergence of handedness in neurons, in eukaryote cell motility, and in non-flagellated bacteria.Comment: 42 pages, 6 figures, resubmitted to J. Stat. Phys. special issue. Major rewrite, rearranged sections/subsections, new Fig 3 + 6, new physics in Sec 2.4 and 3.4.1, added Sec 5 and subsections of Sec

Carbon stable isotopes as a palaeoclimate proxy in vascular plant dominated peatlands

Carbon stable isotope (δ¹³C) records from vascular plant dominated peatlands have been used as a palaeoclimate proxy, but a better empirical understanding of fractionation processes in these ecosystems is required. Here, we test the potential of δ¹³C analysis of ombrotrophic restiad peatlands in New Zealand, dominated by the wire rush (Empodisma spp.), to provide a methodology for developing palaeoclimatic records. We took surface plant samples alongside measurements of water table depth and (micro)climate over spatial (six sites spanning > 10 latitude) and temporal (monthly measurements over 1 year) gradients and analysed the relationships between cellulose δ¹³C values and environmental parameters. We found strong, significant negative correlations between δ¹³C and temperature, photosynthetically active radiation and growing degree days above 0 C. No significant relationships were observed between δ¹³C and precipitation, relative humidity, soil moisture or water table depth, suggesting no growing season water limitation and a decoupling of the expected link between δ¹³C in vascular plants and hydrological variables. δ¹³C of Empodisma spp. roots may therefore provide a valuable temperature proxy in a climatically sensitive region, but further physiological and sub-fossil calibration studies are required to fully understand the observed signal

Size Doesn't Matter: Towards a More Inclusive Philosophy of Biology

notes: As the primary author, O’Malley drafted the paper, and gathered and analysed data (scientific papers and talks). Conceptual analysis was conducted by both authors.publication-status: Publishedtypes: ArticlePhilosophers of biology, along with everyone else, generally perceive life to fall into two broad categories, the microbes and macrobes, and then pay most of their attention to the latter. ‘Macrobe’ is the word we propose for larger life forms, and we use it as part of an argument for microbial equality. We suggest that taking more notice of microbes – the dominant life form on the planet, both now and throughout evolutionary history – will transform some of the philosophy of biology’s standard ideas on ontology, evolution, taxonomy and biodiversity. We set out a number of recent developments in microbiology – including biofilm formation, chemotaxis, quorum sensing and gene transfer – that highlight microbial capacities for cooperation and communication and break down conventional thinking that microbes are solely or primarily single-celled organisms. These insights also bring new perspectives to the levels of selection debate, as well as to discussions of the evolution and nature of multicellularity, and to neo-Darwinian understandings of evolutionary mechanisms. We show how these revisions lead to further complications for microbial classification and the philosophies of systematics and biodiversity. Incorporating microbial insights into the philosophy of biology will challenge many of its assumptions, but also give greater scope and depth to its investigations