Migrating Cambial Domains and the Origin of Wavy Grain in Xylem of Broadleaved Trees

Abstract

Anatomical analysis of wavy-grained xylem samples of Fraxinus, Acer, and Betula reveals a strong underlying pattern of orientational domains within the cambium. Domains are local areas in the cambium within which the orientation of pseudotransverse divisions of fusiform initial cells is predominately unidirectional rather than random. The orientation of ray splitting and ray uniting also tends to be unidirectional within domains. Because their boundaries migrate with time, domains are not fixed in shape or position. A specific group of initial cells may first be located in the interior of a domain, later near a boundary, and still later in another domain of opposite orientational prevalence. Domain patterns and their movements were delineated by classifying and mapping ray splitting and uniting events in temporal series of tangential sections of terminal late xylem. The domain patterns move slowly upward, preceding the similarly outlined, but noncoincident, grain pattern. A wavy grain pattern can be interpreted as being the result of a domain pattern of similar geometry. The efficacy of a domain pattern in eliciting a grain pattern is probably indirectly dependent upon the frequency of occurrence of pseudotransverse divisions and of ray splitting and uniting. The frequency is low in cambium producing straight-grained wood. Evidence of 10-fold and greater increases in frequency of ray splitting and uniting was found in straight-grained to wavy-grained transitional xylem.