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iForest -Biogeosciences and Forestry
IntroductionOut of all factors worldwide that potentially limit geographical distribution ranges of plants, minimum temperatures are considered the most important (Woodward 1987, Dahl 1998, Holten 1998. In particular, frost is thought to be the cause of northern and eastern distribution range boundaries of plant species in Europe (Sakai & Larcher 1987, Pither 2003, Kreyling 2010. The coincidence between species distribution boundaries and freezing temperature thresholds has been already demonstrated for various plant species (Sakai & Larcher 1987, Huntley 1990, Woodward 1997. For example, Fagus sylvatica L. does not occur in regions with a mean temperature in January below -3 °C (Bolte et al. 2007). However, these observations are purely based on correlations and knowledge on which plant organs are affected by frost in which phenological state is virtually absent. Low temperatures affect individuals directly or indirectly by frost dehydration (Pearce 2001, Beck et al. 2004, whereby single cells up to the whole organism can be damaged (Weiser 1970, Pearce 2001. Thus, tree species in the northern hemisphere have evolved a certain frost hardiness despite the high energetic costs involved (Huner et al. 1998). Frost hardiness of European plant species shows a seasonal pattern and at the end of the growing season it is induced by decreasing temperatures (Xin & Browse 2000, Repo et al. 2001, Beck et al. 2004 and is brought about by accumulation of carbohydrates and dehydration of the cells, tissues and organs (Siminovitch & Briggs 1953, Sakai & Larcher 1987, Améglio et al. 2004, Thomas et al. 2004, Morin et al. 2007, Callister et al. 2008.Long-lived plant species go through various development stages with different environmental condition requirements. It is wellknown that frost hardiness varies along the plant life cycle (Sakai & Larcher 1987, Bigras et al. 2004, Taschler et al. 2004, with seedlings exhibiting a lower frost hardiness than adults. For example, Morin et al. (2007) showed for different European Quercus species that adults were significantly more frosttolerant than seedlings. However, despite such differences in frost tolerance between adults and juveniles, one would expect that frost tolerance mechanisms of different ontogenetic stages are similar for a given species. Moreover, seedlings and adults might not share the same micro-environment but are exposed to the same macro-climate at any growth location. In consequence, the survival and growth of seedlings of the tree species Acer pictum subsp. mono (Maximowicz) H. Ohashi and Fagus crenata Blume were found to be in accordance with the distribution patterns of the adult individuals of these species in Japan (Masaki et al. 2005).Periods of low temperatures are critical events in the development of tree seedlings. While seeds of temperate tree species are generally frost-tolerant down to -196 °C (Sakai & Larcher 1987), they lose frost hardiness with the onset of germination (Marcante et al. 2012). As most temperate an...