THE ROTATED-LAMINA SYNDROME .1. ULMACEAE

In a number of plants, mostly woody, the components of the buds are arranged so that the laminae of the young leaves all face towards the same (upper) side of the bud, rather than towards the bud apex; in axillary buds they usually face towards the parent axis. This situation has been known for many years. For convenience, the general case is here called the rotated-lamina syndrome. There have been very few developmental investigations of how the laminae attain their unusual orientation, and these have come to different conclusions about cases in the Ulmaceae. This paper reports a detailed investigation of the syndrome in Ulmus glabra and Zelkova serrata, with comparative observations on other Ulmaceae, including cases in Celtis that do not exhibit the syndrome. The syndrome arises by different means in Ulmus arid Zelkova. In Ulmus the leaf primordium is asymmetrical from the outset, the leaf blade region is obliquely dorsiventral from an early stage, and further asymmetrical growth of the leaf buttress rotates die whole leaf blade region into its final orientation as it develops. Individual shoots show heteroblastic development in progressing from bud scale to foliage leaf initiation, in increasing accentuation of the rotated-lamina syndrome, and in an increasing degree of dorsiventrality. In Zelkova, as previously described, the leaf blade region appears first as a radially symmetrical upgrowth, and it acquires dorsiventral symmetry directly in the rotated position. In Celtis spp. the lamina arises in a quite normal orientation, but reorients as it emerges from the bud. The leaf primordia of all species studied show asymmetry in other aspects, particularly in respect of stipule development, and these seem to be general features of the organisation of dorsiventral shoots.