New perspectives on radial profiles of specific gravity in North American conifers

North American conifers exhibit three radial specific gravity (SG) patterns (Type 1, 2, and 3), which balance hydraulic and mechanical requirements. Type 1 and 2 patterns (Pinaceae) have low SG and high microfibril angle (MFA) corewood ensuring compliance, whereas in outerwood high SG/low MFA provide stiffness and strength resisting bending. Hydraulically, corewood, especially in Type 2 species, is resistant to embolism, whereas outerwood has higher specific conductivity. Cupressaceae (Type 3) have hydraulically very efficient, low SG outerwood, facilitating rapid growth. Corewood is flexible, whereas outerwood is mechanically weak and compensated for by more conical stems and durable heartwood (which prevents Brazier buckling). Radially earlywood (EW) decreases, and latewood (LW) increases for all types, whereas %latewood (%LW) increases (Type 1), decreases, then increases (Type 2) and decreases (Type 3). Ring SG increases when increasing LW SG and %LW are sufficient to counteract decreasing EW SG. Shade tolerance, crown recession, hormone gradients, and environmental variation affect patterns. Auxin concentration decreases with increasing distance from juvenile foliage slowing cell division, concomitantly gibberellin concentration (lignification) and carbohydrates (cell wall thickening) increase, producing higher %LW. Across a species range regions receiving relatively high summer rainfall have trees with higher %LW (by ring).