RESUMEN
In rhythmically growing woody species such as common oak (Quercus robur L.), stem growth is discontinuous and a bud forms at regular intervals at the shoot apex. These buds are composed of different types of leaves: laminate, aborted lamina and scale. The change in heteroblastic leaf shape from laminate to aborted lamina leaves is regarded as one of the events marking shoot growth arrest. To better understand the determinism of heteroblastic leaf shape change and thus, of rhythmicity, we studied morphogenetic events during the early days of the second flush of growth in oak, as well as changes in sucrose metabolism and abscisic acid (ABA) concentrations in control plants expressing the heteroblastic leaf shape change and in defoliated plants showing no heteroblastic leaf shape change and producing only laminate leaves. In control plants, the leaf shape change was underway on Day 5 of the second flush with the differentiation of the first two aborted lamina leaves. Sucrose concentration in the apices of control plants decreased between Days 3 and 5 during differentiation of the aborted lamina leaves. An inverse pattern was observed in defoliated plants, suggesting that sucrose acts as a signal triggering heteroblastic leaf shape changes. During the same period, acid cell wall invertase activity was high in young stem and laminate leaves of control plants, whereas the activity remained constant and low in the apices. If the laminate leaves were removed, the increase in apical sucrose concentration was proportionally higher than the decrease in apical acid vacuolar invertase activity, suggesting that, in the absence of young leaves, sucrose is imported to the apex. The sucrose concentration in the apex is therefore likely to be affected by trophic competition with the expanding laminate leaves. The decrease in apex sucrose concentration may be one of the mechanisms driving heteroblastic leaf shape change. Differentiation of aborted lamina leaves was followed by a decrease in the organogenic activity of the shoot apical meristem (SAM) between Days 7 and 9. High concentrations of ABA are associated with differentiation of aborted lamina and scale leaves and with low SAM organogenic activity. Shoot apical meristem organogenic activity remained high and ABA concentration in the apex remained low in defoliated plants producing only photosynthetic leaves. These results suggest that (1) ABA is involved in the gradual conversion of embryonic leaves to abnormal leaves, thereby regulating heteroblastic leaf shape changes and (2) changes in ABA concentration influence the intensity of SAM organogenic activity. Heteroblastic development and therefore rhythmic growth could be the result of competition between apices and laminate leaves, with competition first involving sucrose and thereafter ABA.