Constructing characters to reflect homology accurately remains a critical problem in systematic analysis of vascular plants, especially where evidence consists primarily of heterogeneous position-related subunits making up an overall complex pattern, which we term "biological fabric." Ideally, accurate assessment of homology lies at the developmental level rather than at the level of mature structure. However, elucidating developmental patterns requires appropriate analytic methods. As an example of a useful approach to analysis of plant tissue fabrics, we present a model of ontogeny in the Calamopityaceae, a family of early seed plants. Members of this group have an unusually thick cortex with complex patterns of cell size and shape. Adopting the methodological assumption that physical stress is the primary causal agent in patterning these tissues, we utilize cell size, shape, and orientation to infer a likely sequence of developmental events. We have employed Fry analysis, a technique of strain analysis derived from structural geology, in order to measure and compare cell shape and orientation in various tissue regions, and from this to infer patterns of tissue strain and cell division. Our analysis indicates that tissue patterning in the mature calamopityacean cortex is a consequence of differential cell growth and proliferation of different cell types and tissue regions, which establish regions of local and regional tissue strain. Increased diameter of the stem takes place by a combination of vascular cambial activity and diffuse secondary growth in the cortex, especially around petiole bases. Whereas most regions of the cortex accommodate expansion of secondary tissues through cell division, inner cortex and tissues immediately surrounding outer fiber bundles retain a surprising degree of primary architecture. The inner cortex displays a complex pattern of strain as a consequence of low cell division rates and complex forces exerted by expansion of secondary vascular tissues and emission of leaf traces. The hypodermis of fiber strands, limited vascular cambial development, and a cortex that remains largely intact through development indicate that at least lateral growth of calamopityacean shoots was determinate. Preliminary observations indicate that the ontogenetic model derived here from Calamopitys is applicable to other members of the Calamopityaceae, and in part to other early seed plants as well. However, the massive and complex calamopityacean cortex is unique among early seed plants and is probably a consequence of the production of large compound leaves in these plants. In assessing functional, ecological, and evolutionary significance of these features, it is necessary to consider them as manifestations of the same developmental process in the plant as a whole rather than as isolated functional units.
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