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

Spatial representation for navigation in animats

This article considers the problem of spatial representation for animat navigation systems. It is proposed that the global navigation task, or "wayfinding, " is best supported by multiple interacting subsystems, each of which builds its own partial representation of relevant world knowledge. Evidence from the study of animal navigation is reviewed to demonstrate that similar principles underlie the wayfinding behavior of animals, including humans. A simulated wayfinding system is described that embodies and illustrates several of the themes identified with animat navigation. This system constructs a network of partial models of the quantitative spatial relations between groups of salient landmarks. Navigation tasks are solved by propagating egocentric view information through this network, using a simple but effective heuristic to arbitrate between multiple solutions