Branches versus stems in woody plants: control of branch diameter growth and angle
The results of three studies at different stages of branch development demonstrated the importance of apical control of diameter growth in both stem formation and branch angle. Diameter growth is controlled by competition between branches and the stem for branch-produced photosynthate. Apical control of branch angle occurs only in species that can produce differential growth stresses. In those species, upward bending is largely regulated by the amount of branch diameter growth. The first study followed stem formation from current shoots in Kalmia latifolia L., a shrub without terminal buds or apical control of branch angle. When several current or older shoots were competing, the longest, most distal lateral shoot usually became the stem. Shoot angle was poorly correlated with eventual dominance. A more proximal lateral shoot on the underside of a leaning parent became the longest, dominant lateral in 24% of the parent shoots. The second study used stem girdles to test the hypothesis that the subjacent stem competes with the branch for branch-produced photosynthate. Results from Pinus strobus L. supported the hypothesis, but results from Betula lenta L. and Acer rubrum L. did not. The third study removed apical control from branches of six forest-shrub species by cutting off the stem above the branch. Branches of all species increased diameter growth after cutting the stem, but only branches of Ilex verticillata (L.) Gray, Hamamelis virginiana L., and Cornus amomum Mill. developed differential growth stress and bent upward. Treated branches of Gaylusaccia baccata (Wang.) K. Koch, Viburnum cassinoides L., and K. latifolia sagged as much as controls.Key words: apical control, diameter growth, branch angle, growth stress, reaction wood.