Weak Apical Control of Swiss Stone Pine (Pinus cembra L.) May Serve as a Protection Against Environmental Stress Above Treeline in the Central European Alps

At the treeline in the Central European Alps, adverse climate conditions impair tree growth and cause krummholz formation of Swiss stone pine (Pinus cembra L.). Multi-stemmed trees (tree clusters) are frequently found in the treeline ecotone and are generally thought to originate from seed caches (multiple genets) of the European nutcracker (N. caryocatactes) or due to repeated damage of the leader shoot by browsing or mechanical stress (single genet). Additionally, lack of apical control can lead to upward bending of lateral branches, which may obscure single-genet origin if the lower branching points are overgrown by vegetation and the humus layer. The multi-stemmed growth form may serve as a means of protection against extreme environmental stress during winter, especially at wind-exposed sites, because leeward shoots are protected from, e.g. ice particle abrasion and winter desiccation. The aims of this study therefore were to analyze in an extensive field survey: (i) whether weak apical control may serve as a protection against winter stress; and (ii) to what extent the multi-stemmed growth form of P. cembra in the krummholz zone is originating from a single genet or multiple genets. To accomplish this, the growth habit of P. cembra saplings was determined in areas showing extensive needle damage caused by winter stress. Multi-stemmed saplings were assigned to single and multiple genets based on determination of existing branching points below the soil surface. The findings revealed that upward bending of lateral branches could protect saplings against winter stress factors, and, although multi-stemmed P. cembra trees were primarily found to originate from multiple genets (most likely seed caches), about 38% of tree clusters originated from upward bending of (partially) buried branches. The results suggest that weak apical control of P. cembra in the sapling stage might be an adaptation to increase survival rate under severe climate conditions prevailing above treeline during winter.

Apical dominance and apical control in multiple flushing of temperate woody species

In young plants of many woody species, the first flush of growth in the spring may be followed by one or more flushes of the terminal shoot if growing conditions are favorable. The occurrence of these additional flushes may significantly affect crown form and structure. Apical dominance (AD) and apical control (AC) are thought to be important control mechanisms in this developmental response. A two-phase AD – AC hypothesis for the factors controlling a subsequent flush is presented and evaluated on the basis of currently known studies. The first, very early phase of this additional flush consists of budbreak and the very beginning of outgrowth of the newly formed current buds on the first flushing shoot. There is evidence that this response often involves the release of AD, which is significantly influenced by the auxin:cytokinin ratio as well as by other signals including nutrients and water. This first phase is immediately followed by a second phase, which consists of subsequent bud outgrowth under the influence of apical control. Although definitive data for hormone involvement in this latter process is sparse, there is some evidence suggesting nutritional mechanisms linked to possible hormone activity. Stem-form defects, a common occurrence in multiple-flushing shoots, are analyzed via the AD – AC hypothesis with suggestions of possible means of abatement.