Oral Paper

         Development and Structure

Interspecific variation in the resting bud morphologies of Juglandoideae—a winter’s tale?

Presenting Author
Kristel Schoonderwoerd
Description
In temperate environments, resting buds are frequently regarded as static units that require cataphylls, or bud scales, to protect preformed embryonic foliage leaves during winter dormancy. However, this winter-protective view is challenged by the existence of naked buds: buds that overwinter future foliage leaves without the protection of cataphylls. Could there be a connection between resting bud structure in winter and the dynamic patterns of shoot extension over the growing season?             We investigated this in Juglandoideae (the temperate subfamily of Juglandaceae), a group that contains remarkable variation in terminal bud structures. Some of its species bear scaly buds with cataphylls. Others bear naked buds and never initiate cataphylls. Still others develop cataphylls during the first phase of bud initiation that are subsequently lost before winter dormancy. Yet, many of these species have large ranges that experience very similar (winter) climate conditions. To understand shoot ontogeny from bud initiation to mature shoot, we characterized the complete seasonal heteroblastic sequences of leaf forms in six species of Juglandoideae. The origin and developmental timing of metamers (foliage leaves, cataphylls, and intermediates) within the resting bud were investigated using micro-CT scanning. In addition, the dimensions of 2249 individual metamers were tracked from first exposure to abscission along the shoots of saplings and mature trees in a common garden setting over a seven-month growing season. For the metamers with prominent photosynthetic areas (n=1581), chlorophyll fluorescence data was collected monthly. The occurrence of cataphylls is associated with a single cohort of foliage leaves that flush and abscise synchronously. This growing pattern is highly determinate, with next year’s terminal buds initiated even before leaf out in spring. In contrast, in sequences without cataphylls, slightly shorter-lived foliage leaves appear and abscise in a staggered fashion. Neoformation featured more prominently in species without cataphylls. Chlorophyll fluorescence data reveal that the effective light-capture area of species with determinate growth (and cataphylls) in most cases reaches a peak early in the growing season and slowly declines thereafter. An indeterminate growth pattern (without cataphylls), on the other hand, ultimately leads to a steady increase in effective light-capture area several months into the growing season with a peak in late summer. Therefore, the observed determinate and indeterminate modes of sequential leaf development in Juglandoideae, and their associated terminal resting bud types, may be reflective of distinct ecological strategies during the growing season. While the protective role of cataphylls during the dormant season has been the subject of much speculation, the developmental consequences of inserting cataphylls into a dynamic seasonal sequence of leaf forms potentially reach far beyond winter in Juglandaceae. In a wider context, bringing a leaf economic spectrum mindset to seasonal heteroblasty could increase our understanding of the ecological functioning of shoot development in temperate, deciduous, woody species and their seasonal patterns of carbon sequestration.