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Redwood branches provide all the carbohydrates for the most carbon-heavy forests on Earth, and recent whole-tree measurements have quantified trunk growth rates associated with complete branch inventories. Providing all of a tree's photosynthetic capacity, branches represent an increasing proportion of total aboveground wood production as trees enlarge. To examine branch development and its effects on wood volume growth, we dissected 31 branches from eight  Sequoia sempervirens  (D. Don) Endl. and seven  Sequoiadendron giganteum  Lindl. trees. The cambium-area-to-leaf-area ratio was maintained with size and age but increased with light availability, whereas the heartwood-deposition-area-to-leaf-area ratio increased with size and age but was insensitive to light availability. The proportion of foliage mass arrayed in &lt;1-cm-diameter epicormic shoots increased with decreasing light and was higher in  Sequoia  (20–60%) than in  Sequoiadendron  (3–16%). Well-illuminated branches concentrated leaves higher and distally, while shaded branches distributed leaves lower and proximally. In similar light environments, older branches distributed leaves lower and more proximally than younger branches. Branch size, light, species, heartwood area, a heartwood-area–species interaction, and ovulate cone mass predicted 87.5% of the variability in wood volume growth of branches. After accounting for the positive effects of size and light, wood volume growth declined with heartwood area and age. The effect of age was trivial compared to the effect of heartwood area, suggesting that heartwood expansion caused the age-related decline in wood volume growth. Additionally,  Sequoiadendron  branches of similar size and light environment with more ovulate cones produced less wood, even though these cones were long-lived and photosynthetic, reflecting the energetic cost of seed production. These results contributed to a conceptual model of branch development in which light availability, injury, heartwood content, gravity, and time interact to produce the high degree of branch structural variation evident within redwood crowns.

Structural development of redwood branches and its effects on wood growth