In pursuit of the whole hypha

We know organisms first of all by their forms. Rabbit and carrot, Neurospora, and Paramecium represent particular shapes and structures, patterns in space and time. Each pattern integrates innumerable molecules into a coherent whole, reproduces itself from one generation to the next, and may persist in this manner for millions of years. In this lecture, I shall discuss efforts to render a dynamic and causal account of biological morphogenesis, using fungal hyphae as a concrete exemplar. Molecular structures and interactions are necessary but not sufficient to specify patterns on a scale three to five orders of magnitude larger. The search for alternatives leads to the importation of the concept of dynamic fields, exemplified by the proposal of Bartnicki-Garcia and Gierz that apical growth and morphogenesis report the operation of a mobile vesicle-supply center. Application of field theories to biological morphogenesis is still at an early stage, but is necessary in order to resolve the paradoxical relationship between genes and form.