Functional Implications of Ubiquitous Semicircular Canal Non-Orthogonality in Mammals

The ‘canonical model’ of semicircular canal orientation in mammals assumes that 1) the three ipsilateral canals of an inner ear exist in orthogonal planes (i.e., orthogonality), 2) corresponding left and right canal pairs have equivalent angles (i.e., angle symmetry), and 3) contralateral synergistic canals occupy parallel planes (i.e., coplanarity). However, descriptions of vestibular anatomy that quantify semicircular canal orientation in single species often diverge substantially from this model. Data for primates further suggest that semicircular canal orthogonality varies predictably with the angular head velocities encountered in locomotion. These observations raise the possibility that orthogonality, symmetry, and coplanarity are misleading descriptors of semicircular canal orientation in mammals, and that deviations from these norms could have significant functional consequences. Here we critically assess the canonical model of semicircular canal orientation using high-resolution X-ray computed tomography scans of 39 mammal species. We find that substantial deviations from orthogonality, angle symmetry, and coplanarity are the rule for the mammals in our comparative sample. Furthermore, the degree to which the semicircular canals of a given species deviate from orthogonality is negatively correlated with estimated vestibular sensitivity. We conclude that the available comparative morphometric data do not support the canonical model and that its overemphasis as a heuristic generalization obscures a large amount of functionally relevant variation in semicircular canal orientation between species.Funding for this research was provided by grants NSFIIS-0208675 (http://www.nsf.gov/cise/iis/hcc_pgm.jsp), and EAR-0948842 (http://www.nsf.gov/awards/award_visualiz​ation.jsp?org=EAR). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.Geological SciencesAnthropologyEmail: [email protected]

Nitrogen enrichment modifies plant community structure via changes to plant-soil feedback.

We tested the hypothesis that N enrichment modifies plant–soil feedback relationships, resulting in changes to plant community composition. This was done in a two-phase glasshouse experiment. In the first phase, we grew eight annual plant species in monoculture at two levels of N addition. Plants were harvested at senescence and the effect of each species on a range of soil properties was measured. In the second phase, the eight plant species were grown in multi-species mixtures in the eight soils conditioned by the species in the first phase, at both levels of N addition. At senescence, species performance was measured as aboveground biomass. We found that in the first phase, plant species identity strongly influenced several soil properties, including microbial and protist biomass, soil moisture content and the availability of several soil nutrients. Species effects on the soil were mostly independent of N addition and several were strongly correlated with plant biomass. In the second phase, both the performance of individual species and overall community structure were influenced by the interacting effects of the species identity of the previous soil occupant and the rate of N addition. This indicates that N enrichment modified plant–soil feedback. The performance of two species correlated with differences in soil N availability that were generated by the species formerly occupying the soil. However, negative feedback (poorer performance on the soil of conspecifics relative to that of heterospecifics) was only observed for one species. In conclusion, we provide evidence that N enrichment modifies plant–soil feedback relationships and that these modifications may affect plant community composition. Field testing and further investigations into which mechanisms dominate feedback are required before we fully understand how and when feedback processes determine plant community responses to N enrichment. Electronic supplementary material The online version of this article (doi:10.1007/s00442-008-1104-0) contains supplementary material, which is available to authorized users