Centric Allometry: Studying Growth Using Landmark Data

Abstract

The geometric morphometric (GMM) construction of Procrustes shape coordinates from a data set of homologous landmark configurations puts exact algebraic constraints on position, orientation, and geometric scale. While position as digitized is not ordinarily a biologically meaningful quantity, and orientation is relevant mainly when some organismal function interacts with a Cartesian positional gradient such as horizontality, size per se is a crucially important biometric concept, especially in contexts like growth, biomechanics, or bioenergetics. “Normalizing” or “standardizing” size (usually by dividing the square root of the summed squared distances from the centroid out of all the Cartesian coordinates specimen by specimen), while associated with the elegant symmetries of the Mardia–Dryden distribution in shape space, nevertheless can substantially impeach the validity of any organismal inferences that ensue. This paper adapts two variants of standard morphometric least-squares, principal components and uniform strains, to circumvent size standardization while still accommodating an analytic toolkit for studies of differential growth that supports landmark-by-landmark graphics and thin-plate splines. Standardization of position and orientation but not size yields the coordinates Franz Boas first discussed in 1905. In studies of growth, a first principal component of these coordinates often appears to involve most landmarks shifting almost directly away from their centroid, hence the proposed model’s name, “centric allometry.” There is also a joint standardization of shear and dilation resulting in a variant of standard GMM’s “nonaffine shape coordinates” where scale information is subsumed in the affine term. Studies of growth allometry should go better in the Boas system than in the Procrustes shape space that is the current conventional workbench for GMM analyses. I demonstrate two examples of this revised approach (one developmental, one phylogenetic) that retrieve all the findings of a conventional shape-space-based approach while focusing much more closely on the phenomenon of allometric growth per se. A three-part Appendix provides an overview of the algebra, highlighting both similarities to the Procrustes approach and contrasts with it.