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Studies addressing endodormancy release in adult trees are usually carried out using twigs detached from the trees in the experiments. Potential problems caused by cutting the root-shoot connection when detaching the twigs can be avoided by using grafts as the experimental material. We studied the effects of chilling on the endodormancy release in Norway spruce (Picea abies (L.) Karst.) grafts where twigs of 16-, 32- and 80-year-old trees were used as the scions. The grafts were first exposed to chilling in natural conditions and then samples of them were transferred at intervals to a regrowth test in forcing conditions in a greenhouse. The bud burst percentage, BB%, in the forcing conditions generally increased from zero to near 100% with increasing previous chilling accumulation from mid-October until mid-November, indicating that endodormancy was released in almost all of the grafts by mid-November. The days to bud burst, DBB, decreased in the forcing conditions with successively later transfers until the next spring. Neither BB% nor DBB was dependent on the age of the scion. However, in the early phase of ecodormancy release, the microscopic internal development of the buds was more advanced in the grafts representing the 16-year-old than in those representing the 32- or 80-year-old trees. In conclusion, our findings suggest that no major change in the environmental regulation of endodormancy release in Norway spruce takes place when the trees get older. Taken together with earlier findings with Norway spruce seedlings, our results suggest that regardless of the seedling or tree age, the chilling requirement of endodormancy release is met in late autumn. The implications of our findings for Norway spruce phenology under climatic warming and the limitations of our novel method of using grafts as a proxy of trees of different ages are discussed., (© The Author(s) 2020. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

The timing of bud development in ecodormancy is critical for trees in boreal and temperate regions with seasonally alternating climates. The development of vegetative buds and the growth of primordial shoots (the primordial shoot ratio) in Norway spruce were followed by the naked eye and at stereo and light microscopic levels in fresh-cut and fixed buds obtained by regular field samplings during the spring of 2007, 2008 and 2009. Buds were collected from 15 randomly selected trees (all 16 years old in 2007) of one southern Finnish half-sib family. The air temperature was recorded hourly throughout the observation period. In 2008 and 2009, initial events in the buds, seen as accumulation of lipid droplets in the cortex area, started in mid-March and were depleted in late April, simultaneously with the early development of vascular tissue and primordial needles. In mid-April 2007, however, the development of the buds was at least 10 days ahead as a result of warm spells in March and early April. Variation in the timing of different developmental phases within and among the sample trees was negligible. There was no clear one-to-one correspondence between the externally visible and the internal development of the buds. The dependence of the primordial shoot ratio on different types of temperature sum was studied by means of regression analysis. High coefficients of determination (R(2) ≈ 95%) were attained with several combinations of the starting time (beginning of the year/vernal equinox), the threshold value (from -3 to +5 °C), and the time step (hour/day) used in the temperature summation, i.e., the prediction power of the primordial shoot ratio models turned out to be high, but the parameter estimate values were not unambiguous. According to our results, temperature sums describe the growth of the primordial shoot inside the bud before bud burst. Thus, the results provide a realistic interpretation for the present phenological models of bud development that are based on temperature sums and external observations of bud burst only, and they also provide new tools for improving the models.

Cold deacclimation and preparation to flushing likely requires rehydration of meristems. Therefore, water stress related genes, such as dehydrins (DHN), might play an important role in providing protection during winter dormancy, deacclimation and bud burst timing processes. Here we report the sequence analysis of several Norway spruce DHN identified in late and early flushing suppressive subtraction hybridization cDNA libraries and in our Norway spruce EST database. We obtained 15 cDNAs, representing eight genes from three distinct types of DHN, and studied differential expression of these genes before and during bud burst in spring, using qRT-PCR. We found the visible reduction in transcript level of most DHN towards the bud burst, supported by a significant down-regulation of the DHN in needles during experimental induction of bud burst applied at three time points during autumn in Norway spruce grafts. For most of the DHN transcripts, their expression levels in late-flushing spruces were significantly higher than in the early flushing ones at the same calendar dates but were remarkably similar at the same bud developmental stage. From our results we may conclude that the difference between the early and the late families is in timing of the molecular processes leading to bud burst due to differences in their response to the increasing temperature in the spring. They are induced much earlier in the early flushing families.