Endodormancy release in Norway spruce grafts representing trees of different ages.

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.

Reliability of temperature signal in various climate indicators from northern Europe.

We collected relevant observational and measured annual-resolution time series dealing with climate in northern Europe, focusing in Finland. We analysed these series for the reliability of their temperature signal at annual and seasonal resolutions. Importantly, we analysed all of the indicators within the same statistical framework, which allows for their meaningful comparison. In this framework, we employed a cross-validation procedure designed to reduce the adverse effects of estimation bias that may inflate the reliability of various temperature indicators, especially when several indicators are used in a multiple regression model. In our data sets, timing of phenological observations and ice break-up were connected with spring, tree ring characteristics (width, density, carbon isotopic composition) with summer and ice formation with autumn temperatures. Baltic Sea ice extent and the duration of ice cover in different watercourses were good indicators of winter temperatures. Using combinations of various temperature indicator series resulted in reliable temperature signals for each of the four seasons, as well as a reliable annual temperature signal. The results hence demonstrated that we can obtain reliable temperature information over different seasons, using a careful selection of indicators, combining the results with regression analysis, and by determining the reliability of the obtained indicator.

Internal development of vegetative buds of Norway spruce trees in relation to accumulated chilling and forcing temperatures.

The timing of budburst of temperate trees is known to be controlled by complicated interactions of temperature and photoperiod. To improve the phenological models of budburst, better knowledge of the internal bud development preceding budburst in relation to environmental cues is needed. We studied the effect of accumulated chilling and forcing temperatures on the internal development of vegetative buds preceding budburst in Norway spruce [Picea abies (L.) Karst.]. Branches from 17-year-old trees of southern Finnish origin were transferred eight times at 1- to 2-week intervals from October to December 2007 from the field at Punkaharju (61°48'N, 29°20'E) to the greenhouse with forcing conditions (day length 12 h, +20 °C). After seven different durations of forcing, the developmental phase and primordial shoot growth of the buds were analysed at the stereomicroscopic level. Air temperature was recorded hourly throughout the study period. The accumulated chilling unit sum had a significant effect on the temperature sum that was required to attain a certain developmental phase; a higher amount of chilling required a lower amount of forcing. The variation in the rate of development of different buds within each sample branch in relation to the chilling unit and forcing temperature sum was low. Regarding primordial shoot growth, there was also an inverse relation between accumulated chilling and forcing, i.e., a higher accumulated chilling unit sum before forcing required a lower temperature sum to initiate primordial shoot growth and resulted in a stronger effect of accumulated forcing. A second-order regression model with an interaction of chilling and forcing explained the variation of primordial shoot growth with high precision (R(2) = 0.88). However, further studies are required to determine the final parameter values to be used in phenological modelling.

Development and growth of primordial shoots in Norway spruce buds before visible bud burst in relation to time and temperature in the field.

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.

Anatomy and morphology in developing vegetative buds on detached Norway spruce branches in controlled conditions before bud burst.

We studied the light and stereomicroscopic structure of developing vegetative buds from a 16-year-old Norway spruce [Picea abies (L.) Karst.] of southern Finnish origin in relation to temperature sum and to externally visible changes in the buds before and during bud burst in forcing conditions. Branches were collected on 17 January and transferred to the greenhouse where they were first subjected to preforcing conditions (darkness, +4 degrees C) for 7 days and then to the forcing conditions (day length 12 h, +20 degrees C). Buds were sampled 20 times between 17 January and 13 February. Air temperature was recorded hourly throughout the study period. The first microscopic change was a temporary increase in the size and number of lipid droplets before the onset of temperature sum (T > or = +5 degrees C) accumulation. From the 4th to the 9th day under the forcing conditions, tracheids started to develop from the base up to the top of the bud. This was closely synchronized with an observed morphological change in the shape of needle tip from rounded to pointed ones. Development from the first visible change in the bud scales on the 12th forcing day to bud burst took 9 days when the temperature sum was 313 d.d. The temperature sums in our experiment overestimated the requirements of temperature sum for bud development phases measured in the field. Bud development could be divided into four structural phases. The first two phases, i.e., morphological changes in the primary needles, occurred without any externally visible changes in the buds. Thus, these phases have a potential for testing and improving the phenological models, which, up to now, have mainly been based on the bud burst observation by the naked eye.

Models of the spring phenology of boreal and temperate trees: Is there something missing?

According to prevailing theory, air temperature is the main environmental factor regulating the timing of bud burst of boreal and temperate trees. Air temperature has a dual role in this regulation. First, after the cessation of growth in autumn, prolonged exposure to chilling causes rest completion, i.e., removes the physiological growth-arresting conditions inside the bud. After rest completion, prolonged exposure to warm conditions causes ontogenetic development leading to bud burst or flowering. During the past three decades, several simulation models based on chilling and forcing have been developed and tested. In recent modeling studies of the timing of bud burst in mature trees, the simpler thermal-time models that assume forcing starts on a fixed date in the spring have outperformed the chilling-forcing models. We hypothesize that this discrepancy may be due to some element missing from the chilling-forcing models. We tested two new model formulations by introducing reversing, temperature-driven elements that precede forcing and by fitting the models to seven historical time series of data of flowering and leaf bud burst of common boreal tree species. In these tests, both of the new models were generally more accurate in predicting the timing of bud burst than a classical chilling-forcing model, but less accurate than the simple thermal-time model. We therefore conclude that besides chilling, other environmental factors are involved in the regulation of the timing of bud burst. Further work is needed to determine if the regulatory factors derive from air temperature or from some other environmental condition such as changes in light conditions, like day length or night length.

Critical night length for bud set and its variation in two photoperiodic ecotypes of Betula pendula.

We studied the variation in critical night length for bud set in two photoperiodic ecotypes (two latitudinally distant stands) of silver birch (Betula pendula Roth) in three phytotron experiments. Seeds from 21 open-pollinated mother trees in a southern (Tuusula, 60 degrees N) and a northern (Kittilä, 67 degrees N) Finnish stand were germinated and grown for 4 weeks in a 24-h photoperiod in a greenhouse and then moved to different night length treatments at 18 degrees C for 4 to 6 weeks. Night lengths from 5 to 8.5 h were used for southern origin seedlings and from 1 to 4.5 h for northern origin seedlings. At the end of the treatments, apical bud set was observed and the percentage of seedlings with bud set calculated for each treatment and tree progeny. The critical night lengths (CNL) for 50% bud set were determined separately for seedlings from each mother tree by regression analysis. In both ecotypes, the mean percentage of seedlings with bud set was lowest for the shortest night lengths and increased rapidly as night lengths increased. Mean CNL with its 95% confidence interval for the southern and northern ecotypes was 6.3 +/- 0.2 and 3.1 +/- 0.3 h, respectively. The CNL of the two ecotypes differed significantly in three experiments. Within-ecotype variance of the CNL was significantly higher in the northern ecotype (0.484) than in the southern ecotype (0.150). Significant differences in CNL were detected between individual mother trees of the southern ecotype, but not between mother trees of the northern ecotype. The ranking of individual mother trees, based on CNL, differed in the three experiments.

Effects of seed origin and sowing time on timing of height growth cessation of Betula pendula seedlings.

We studied the effects of seed origin and sowing time on height development and timing of height growth cessation of first-year silver birch (Betula pendula Roth) seedlings in a greenhouse experiment. Seeds of seven origins ranging in latitudes from 58 degrees to 67 degrees N were sown at 1-2-week intervals eight times from May 21 to July 30, 2001. The day/night temperature in the greenhouse was set at 20/10 degrees C, but lighting was natural and day length varied accordingly. Seedling height was measured twice a week. The interaction term between seed origin and sowing date was significant, but the pattern of height development and timing of growth cessation depended systematically on latitude of seed origin and sowing date. As seed origin became increasingly northern, growth cessation began earlier and resulted in shorter growth periods. Later sowing dates delayed growth cessation but also shortened the growth period. Final seedling height systematically decreased with increasingly northern origins and with later sowings. Linear regression analysis predicted timing of growth cessation, night length at growth cessation, length of growth period and final seedling height with high precision when the latitude of seed origin and sowing time were predictor variables. The timing of height growth cessation was determined by the seed origin, night length and developmental stage of the seedlings.

Predicting spring phenology and frost damage risk of Betula spp. under climatic warming: a comparison of two models.

Timing of bud burst and frost damage risk for leaves of Betula spp. in response to climatic warming in Finland was examined with two models. In the first model, ontogenetic development in spring was triggered by an accumulation of chilling temperatures. The second model assumed an additional signal from the light climate. The two models gave radically different estimates of frost damage risk in response to climate warming. The chilling-triggered model forecast a significant and increasing risk with increased warming, whereas the light-climate-triggered model predicted little or no risk. The chilling-triggered model is widely applied in phenological research; however, there is increasing experimental evidence that light conditions play a role in the timing of spring phenology. Although it is not clear if the light response mechanisms are appropriately represented in our model, the results imply that reliance on a light signal for spring development would afford a degree of protection against possible frost damage under climate warming that would not be present if chilling were the sole determinant. Further experimental tests are required to ascertain the light-related mechanisms controlling phenological timing, so that credible model extrapolations can be undertaken.

Statistical evaluation of bud development theories: application to bud burst of Betula pendula leaves.

Resampling methods were used to evaluate models based on alternative bud development theories applied to Betula pendula Roth data. Statistical testing based on the bootstrap method showed that the mean square errors (MSE) of the predicted bud-burst dates of two models, in which the start of ontogenetic development depended on dormancy development only, did not differ significantly. However, the MSE of the model in which the start of ontogenesis depended on a signal from light climate, indicated by using a fixed calendar date, was significantly smaller than that of the models depending on dormancy development. Model parameters were highly multi-collinear; i.e., sensitive to changes in the data. The cross-validation method was used to determine the prediction error of the models. The predictive ability of the models was not much less for an independent data set than for the original data.

Effects of dormancy and environmental factors on timing of bud burst in Betula pendula.

We tested three theories predicting the timing of bud burst in mature birch (Betula pendula Roth) trees utilizing a 60-year phenological time series together with meteorological temperature observations. Predictions of the timing of bud burst based on light conditions in addition to temperature were more accurate than predictions based on dormancy development and temperature (prediction standard error of 2.4 days versus 4.3 days). The signal from light conditions, represented by fixed calendar date, determined the start of bud ontogenesis rather than dormancy release. We suggest that models developed to predict the timing of bud burst be utilized in the analysis of plant responses to climate change and of climate change itself.