Response to stem bending in forest shrubs: stem or shoot reorientation and shoot release

Shrubs in the forest understory may be bent by their own weight or by overstory debris. To maintain height growth they must respond to bending by vertical growth of new shoots, reorientation of older axes, or by releasing preventitious buds to form epicormic shoots. I tested for these responses in Ilex verticillata L., Cornus amomum Mill., Gaylussacia baccata (Wang.) K. Koch, Viburnum cassinoides L., Hamamelis virginiana L., and Kalmia latifolia L. For each species, I removed potentially supporting vegetation adjacent to 20 stems, left 10 stems untreated to test for bending by self weight, and bent the remaining 10 stems to 45° to simulate effects of fallen debris. Stem angles and curvatures were measured from before leaf out until just before leaf fall to detect either sagging from self weight or upward bending from tension wood action. Control stems initially leaned out of vertical and five of six species sagged further into a cantilever form. Several control stems failed and bent to the ground. Stems of H. virginiana, I. verticillata, and C. amomum formed tension wood, but only the first two species bent upward. Viburnum cassinoides, G. baccata, and K. latifolia formed no tension wood and sagged further down after being bent. Epicormic shoots formed with varying frequencies in all species except K. latifolia. Epicormic shoots were the major response in C. amomum, V. cassinoides, and G. baccata. New terminal shoots on bent stems recovered toward vertical in I. verticillata and K. latifolia. Negative gravitropic response of shoots was the only recovery mechanism for K. latifolia. Key words: forest shrubs, tension wood, epicormic shoots, bending, stem architecture.