Hydrostatic constraints on morphological exploitation of light in tall Sequoia sempervirens trees.

We studied changes in morphological and physiological characteristics of leaves and shoots along a height gradient in Sequoia sempervirens, the tallest tree species on Earth, to investigate whether morphological and physiological acclimation to the vertical light gradient was constrained by hydrostatic limitation in the upper crown. Bulk leaf water potential (Psi) decreased linearly and light availability increased exponentially with increasing height in the crown. During the wet season, Psi was lower in the outer than inner crown. C isotope composition of leaves (delta(13)C) increased with increasing height indicating greater photosynthetic water use efficiency in the upper crown. Leaf and shoot morphology changed continuously with height. In contrast, their relationships with light availability were discontinuous: morphological characteristics did not correspond to increasing light availability above 55-85 m. Mass-based chlorophyll concentration (chl) decreased with increasing height and increasing light availability. In contrast, area-based chl remained constant or increased with increasing height. Mass-based maximum rate of net photosynthesis (P (max)) decreased with increasing height, whereas area-based P (max) reached maximum at 78.4 m and decreased with increasing height thereafter. Mass-based P (max) increased with increasing shoot mass per area (SMA), whereas area-based P (max) was not correlated with SMA in the upper crown. Our results suggest that hydrostatic limitation of morphological development constrains exploitation of light in the upper crown and contributes to reduced photosynthetic rates and, ultimately, reduced height growth at the tops of tall S. sempervirens trees.

Physiological and ecological implications of adaptive reiteration as a mechanism for crown maintenance and longevity.

Reiteration is the process whereby architectural units are replicated within a tree. Both immediate (from apical buds) and delayed (from suppressed or adventitious buds) reiteration can be seen in many tree species where architectural units ranging from clusters of shoots to entire branches and stems are replicated. In large old trees and suppressed trees, delayed reiteration occurs without an obvious external stimulus such as defoliation or traumatic loss of the branch apex. This suggests that, in trees that are growth-limited, reiteration is an adaptive mechanism for crown maintenance. We discuss theories about the aging process and how delayed adaptive reiteration may help maintain crown productivity and increase longevity. These include: (1) reducing the respiration/photosynthesis ratio; (2) increasing hydraulic conductance to newly developing foliage; (3) reducing nutrient loss from the tree; and (4) rejuvenating the apical meristem. The ability to reiterate various architectural units may contribute to increasing lifetime reproductive output by prolonging tree longevity. Further studies on the physiological and ecological implications of reiteration are needed to understand its adaptive significance in the life history of trees.