Dormancy release and chilling requirement of buds of latitudinal ecotypes of Betula pendula and B. pubescens

Bud burst and dormancy release of latitudinal ecotypes of Betula pendula Roth and B. pubescens Ehrh. from Denmark ( approximately 56 degrees N), mid-Norway ( approximately 64 degrees N) and northern Norway ( approximately 69 degrees N) were studied in controlled environments. Dormant seedlings were chilled at 0, 5 or 10 degrees C from October 4 onward and then, at monthly intervals from mid-November to February, batches of seedlings were held at 15 degrees C in an 8-h (SD) or 24-h (LD) photoperiod to permit flushing. A decline in days to bud burst occurred with increasing chilling time in all ecotypes. In November, after 44 chilling days, time to bud burst was least in plants chilled at 0 and 5 degrees C. The difference diminished with increasing chilling time, and in February, after 136 chilling days, bud burst was earliest in plants chilled at 10 degrees C. Long photoperiods during flushing significantly reduced thermal time after short chilling periods (44 and 74 days), but had no effect when the chilling requirement was fully met after 105 or more chilling days. No significant difference in these responses was found between the two species. In both species, chilling requirement decreased significantly with increasing latitude of origin. Bud burst was normal in seedlings overwintered at 12 degrees C, but was erratic and delayed in seedlings overwintered at 15 and especially at 21 degrees C, indicating that the critical overwintering temperature is between 12 and 15 degrees C. We conclude that there is little risk of a chilling deficit in birch under Scandinavian winter conditions even with a climatic warming of 7-8 degrees C. The likely effects of a climatic warming include earlier bud burst, a longer growing season and increased risk of spring frost injury, especially in high latitude ecotypes.     

Twilight far-red treatment advances leaf bud burst of silver birch (Betula pendula)

Bud development of boreal trees in spring, once initiated, is driven by ambient air temperature, but the mechanism triggering bud development remains unclear. We determined if some aspect of the diurnal or seasonal light regime influences initiation of bud burst once the chilling requirement is met. We grew 3-year-old birch plantlets cloned from a mature tree of boreal origin in light conditions realistically simulating the lengthening days of spring at 60 degrees N. To emulate the reduction in red to far-red light (R:FR) ratio between daylight and twilight, one group of plantlets was subjected to reduced R:FR ratio in the morning and evening in addition to progressively lengthening days, whereas the other group was subjected to the same R:FR ratio throughout the day. The reduced R:FR ratio of twilight advanced bud burst by 4 days compared with the reference group (P = 0.04). To assess the interplay between the fulfillment of the chilling requirement and the subsequent response to warming, we fitted a thermal time model to the data with separate parameterizations for the starting dates of heat sum accumulation in each treatment. Least-squares fitting suggested that bud development started in light regimes corresponding to late March, almost two months after the chilling requirement for dormancy release was satisfied. Therefore, shortening night length or increasing day length, or both, appears to be the cue enabling bud development in spring, with twilight quality having an effect on the photoperiodic response. If twilight alone were the cue, the difference in bud burst dates between the experimental groups would have been greater than 4 days. The result gives experimental support for the use of thermal-time models in phenological modeling.     

Climatic control of bud burst in young seedlings of nine provenances of Norway spruce

Detailed knowledge of temperature effects on the timing of dormancy development and bud burst will help evaluate the impacts of climate change on forest trees. We tested the effects of temperature applied during short-day treatment, duration of short-day treatment, duration of chilling and light regime applied during forcing on the timing of bud burst in 1- and 2-year-old seedlings of nine provenances of Norway spruce (Picea abies (L.) Karst.). High temperature during dormancy induction, little or no chilling and low temperature during forcing all delayed dormancy release but did not prevent bud burst or growth onset provided the seedlings were forced under long-day conditions. Without chilling, bud burst occurred in about 20% of seedlings kept in short days at 12 degrees C, indicating that young Norway spruce seedlings do not exhibit true bud dormancy. Chilling hastened bud burst and removed the long photoperiod requirement, but the effect of high temperature applied during dormancy induction was observed even after prolonged chilling. Extension of the short-day treatment from 4 to 8 or 12 weeks hastened bud burst. The effect of treatments applied during dormancy development was larger than that of provenance; in some cases no provenance effect was detected, but in 1-year-old seedlings, time to bud burst decreased linearly with increasing latitude of origin. Differences among provenances were complicated by different responses of some origins to light conditions under long-day forcing. In conclusion, timing of bud burst in Norway spruce seedlings is significantly affected by temperature during bud set, and these effects are modified by chilling and environmental conditions during forcing.     

Effect of Temperature on the Induction of Bud Dormancy in Ecotypes of Betula pubescens and Betula pendula

The purpose of this study was to determine the influence of temperature applied during short day-induced budset on induction of dormancy in six ecotypes of Betula pubescens Ehrh. and two ecotypes of Betula pendula Roth. Seedlings were grown in a phytotron at constant temperatures of 9–21°C under a 12 h photoperiod (SD) during dormancy induction. Induction of dormancy was monitored by following bud flushing and shoot growth after transfer to long photoperiod conditions (24 h) at 18°C. Chilling requirement was studied in seedlings exposed to 10 weeks of SD. In both species induction of bud dormancy developed most rapidly at 15–18°C, and both 9–12°C and 21°C delayed the induction of dormancy. Raising the temperature (from 9 to 21°C) applied during induction of dormancy significantly increased the chilling requirement. These responses were noted for all ecotypes tested, but in general the northern ecotypes entered dormancy more quickly than the southern ones. No such trend was recorded for chilling requirement, although a B. pubescens ecotype from Iceland and another from the coast of northern Norway appeared to require a longer chilling treatment than the other ecotypes. In conclusion, induction and depth of bud dormancy in birch are significantly affected by temperature conditions and these effects may explain some of the annual variation in dormancy and chilling requirement observed in nature.     

Autumn temperature affects the induction of dormancy in first‐year seedlings of acer platanoides L.

First‐year seedlings of five latitudinal populations of Acer platanoides were subjected to decreasing photoperiod treatment under three different temperature regimes. The depth of the induced dormancy was quantified as the number of days to bud burst (DBB) under defined conditions favourable to growth. The results suggested a close relationship between autumn temperature and the strength of the induced dormancy, with high temperatures combined with short days leading to a deeper stage of dormancy. Northern and continental populations generally had bud burst earlier than southern. The results are discussed in relation to hypotheses for dormancy induction and release.     

Different responses of northern and southern ecotypes of Betula pendula to exogenous ABA application

We investigated responses of northern and southern ecotypes of silver birch (Betula pendula Roth) to exogenous abscisic acid (ABA) under controlled environmental conditions to determine the role of ABA in cold acclimation and dormancy development. Abscisic acid was sprayed on the leaves and changes in freezing tolerance, determined by the electrolyte leakage test, and bud dormancy were monitored. Applied ABA induced cold acclimation but had no effect on growth cessation in seedlings grown in long day conditions (LD, 24-h photoperiod at 18 degrees C). It enhanced freezing tolerance and accelerated growth cessation in seedlings grown in short day conditions (SD, 12-h photoperiod at 18 degrees C), and slightly enhanced freezing tolerance in seedlings grown at low temperature (LT, 24-h photoperiod at 4 degrees C) in both ecotypes. There were distinct ecotypic differences in ABA-induced cold acclimation and dormancy development. The northern ecotype was more responsive to applied ABA than the southern ecotype, resulting in more rapid development of freezing tolerance in all treatments, and earlier dormancy development in SD. When plants were grown in a photoperiod just above the critical photoperiod for the ecotype (defined as the longest photoperiod that induces growth cessation), applied ABA caused growth cessation and dormancy development. Compared with ABA-treated seedlings grown in SD, dormancy development was delayed in ABA-treated seedlings exposed to a near-critical photoperiod, but even in this treatment dormancy developed faster in the northern ecotype than in the southern ecotype.     

Effects of elevated CO(2) and temperature on cold hardiness and spring bud burst and growth in Douglas-fir (Pseudotsuga menziesii)

We examined effects of elevated CO(2) and temperature on cold hardiness and bud burst of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings. Two-year-old seedlings were grown for 2.5 years in semi-closed, sunlit chambers at either ambient or elevated (ambient + ~ 4 degrees C) air temperature in the presence of an ambient or elevated (ambient + ~ 200 ppm) CO(2) concentration. The elevated temperature treatment delayed needle cold hardening in the autumn and slowed dehardening in the spring. At maximum hardiness, trees in the elevated temperature treatment were less hardy by about 7 degrees C than trees in the ambient temperature treatment. In general, trees exposed to elevated CO(2) were slightly less hardy during hardening and dehardening than trees exposed to ambient CO(2). For trees in the elevated temperature treatments, date to 30% burst of branch terminal buds was advanced by about 6 and 15 days in the presence of elevated CO(2) and ambient CO(2), respectively. After bud burst started, however, the rate of increase in % bud burst was slower in the elevated temperature treatments than in the ambient temperature treatments. Time of bud burst was more synchronous and bud burst was completed within a shorter period in trees at ambient temperature (with and without elevated CO(2)) than in trees at elevated temperature. Exposure to elevated temperature reduced final % bud burst of both leader and branch terminal buds and reduced growth of the leader shoot. We conclude that climatic warming will influence the physiological processes of dormancy and cold hardiness development in Douglas-fir growing in the relatively mild temperate region of western Oregon, reducing bud burst and shoot growth.     

Differential responses of silver birch (Betula pendula) ecotypes to short-day photoperiod and low temperature

We investigated interrelations of dormancy and freezing tolerance and the role of endogenous abscisic acid (ABA) in the development of silver birch (Betula pendula Roth) ecotypes in controlled environments. Short-day treatment induced growth cessation, bud set and dormancy development, as well as initiation of cold acclimation and an increase in freezing tolerance. Subsequent low temperature and short days (12-h photoperiod) resulted in a significant increase in freezing tolerance, whereas bud dormancy was gradually released. The concentration of ABA increased in response to short days and then remained high, but ABA concentrations fluctuated irregularly when the dormant plants were subsequently exposed to low temperature during short days. Although there was a parallel development of freezing tolerance and bud dormancy in response to short days, subsequent exposure to low temperature had opposite effects on these processes, enhancing freezing tolerance and releasing dormancy. Compared with the southern ecotype, the northern ecotype was more responsive to short days and low temperature, exhibiting earlier initiation of cold acclimation, growth cessation and an increase in ABA concentrations in short days, and higher freezing tolerance, faster dormancy release and greater alteration in ABA concentrations when subsequently exposed to low temperature during short days. The rates and extent of the increases in ABA concentration may be related to increases in freezing tolerance and dormancy development during short days, whereas the extent of the fluctuations in ABA concentration may play an important role in enhancing freezing tolerance and releasing dormancy during a subsequent exposure to low temperature during short days.     

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.     

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.     

Temperature sum accumulation effects on within-population variation and long-term trends in date of bud burst of European white birch (Betula pendula)

Within-population variation in phenology of boreal trees indicates their adaptability to climatic variations. Although interannual variations in date of bud burst have been widely discussed, little is known about within-population variation, the key determinants for this variation and the effects of this variation on estimates of trends in bud burst date. Over a period of nine years, we monitored timing of bud burst daily in 30 mature white birch (Betula pendula Roth) trees in a naturally regenerated stand. Our results revealed not only large interannual variation but also considerable intraannual variation among individual trees in date of bud burst, the maximum within-population variation being four weeks. Bud burst can be accurately predicted by the date when a threshold value of temperature sum in spring is reached (base temperature +5 degrees C). Based on this temperature sum and past temperature records, we estimated the trend in date of bud burst. The linear trend estimate based on the years 1926-2005 is an advancement of 1.2 days per decade (95% confidence interval, +/- 0.7 days), which is much less than that predicted by time series based on coarser time intervals. We conclude that, because of large interannual differences, and large annual within-population variations in bud burst, estimates of bud burst date based on measurements made over a period of only a few decades are unreliable.     

Low temperature, but not photoperiod, controls growth cessation and dormancy induction and release in apple and pear

In contrast to most temperate woody species, apple and pear and some other woody species of the Rosaceae family are insensitive to photoperiod, and no alternative environmental seasonal signal is known to control their dormancy. We studied growth and dormancy induction in micropropagated plants of four apple (Malus pumila Mill.) and one pear (Pyrus communis L.) commercial rootstock cultivars in controlled environments. The results confirm that growth cessation and dormancy induction in apple and pear are not influenced by photoperiod, and demonstrate that low temperature (< 12 degrees C) consistently induces both processes, regardless of photoperiodic conditions. Successive stages of the autumn syndrome (growth cessation, formation of bud scales and winter buds, leaf senescence and abscission, and dormancy induction) occurred in response to low temperature. Long days increased internode length at higher temperatures, but had no significant effect on leaf production in any of the cultivars. Chilling at 6 or 9 degrees C for at least 6 weeks (about 1000 h) was required for dormancy release and growth resumption, whereas treatment at 12 degrees C was marginally effective, even after 14 weeks of exposure. We are thus faced with the paradox that the same low temperature conditions that induce dormancy are also required for dormancy release in these species.     

Relationships among cold hardiness, root growth potential and bud dormancy in three conifers

Greenhouse-cultured, container-grown ponderosa pine (Pinus ponderosa var. scopulorum Engelm.), interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) and Engelmann spruce (Picea engelmannii (Parry) Engelm.) were cold acclimated and deacclimated in growth chambers over 19 weeks. Stem cold hardiness, total new root length at 14 days and days to bud break were measured weekly. Relationships among cold hardiness, root growth potential (RGP) and bud dormancy suggest that cold hardiness, which can be measured quickly, could provide a useful basis for estimating the two other parameters. During cold acclimation, there was a lag period in which stem cold hardiness remained at -15 degrees C and RGP was at a minimum, in all three species. Douglas-fir and Engelmann spruce buds remained fully dormant during this lag period. Ponderosa pine buds had no chilling requirement for the loss of dormancy, and reached quiescence during the lag period. Immediately following the lag period, as stem cold hardiness progressed to -22 degrees C, RGP increased to a high plateau in all three species, and Douglas-fir and Engelmann spruce buds approached quiescence. Cold deacclimation and bud development began immediately on exposure to warm, long days, but RGP remained high until stem cold hardiness returned to approximately -15 degrees C. At bud break, cold hardiness and RGP were at the minimum.     

Effects of "near-lethal" stress on bud dormancy and stem cold hardiness in red-osier dogwood

We studied the effects of "near-lethal" (NL, 47 degrees C for 1 h) heat stress, applied to intact shoots of red-osier dogwood (Cornus sericea L.) during early (October), deep (November) or late (December) dormancy, on bud dormancy release and development of stem tissue cold hardiness under natural conditions and at a constant temperature of 0 or 23 degrees C in the dark. The NL heat-stress treatment overcame bud dormancy when applied during the early and late stages of dormancy. During October and December, all plants in the 23 degrees C + dark post-stress environment broke bud within 35 and 12 days, respectively, whereas the corresponding values for days to bud break in the control plants were more than 150 and 110 days, respectively. Application of NL heat stress during deep dormancy caused only slightly earlier bud break compared to the control plants. In the 0 degrees C + dark post-stress environment, all NL heat-treated plants died within 9 weeks. Under natural post-stress conditions, bud break in plants receiving NL heat stress during early and deep dormancy occurred at the same time as in control plants, whereas bud break of plants receiving NL heat stress during late dormancy occurred 55 days earlier than in control plants. Under both natural and 23 degrees C + dark post-stress conditions, cold hardiness of plants receiving NL heat stress during early dormancy was similar to that of controls. Application of NL heat stress during deep dormancy hastened the rate of deacclimation under the 23 degrees C + dark post-stress conditions but had no effect on deacclimation under natural post-stress conditions. Application of NL heat stress during late dormancy enhanced deacclimation of plants in both the 23 degrees C + dark and natural post-stress environments.     

Temperature regulation of bud-burst phenology within and among years in a young Douglas-fir (Pseudotsuga menziesii) plantation in western Washington, USA

Past research has established that terminal buds of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings from many seed sources have a chilling requirement of about 1200 h at 0-5 degrees C; once chilled, temperatures > 5 degrees C force bud burst via accumulation of heat units. We tested this sequential bud-burst model in the field to determine whether terminal buds of trees in cooler microsites, which receive less heat forcing, develop more slowly than those in warmer microsites. For three years we monitored terminal bud development in young saplings as well as soil and air temperatures on large, replicated plots in a harvest unit; plots differed in microclimate based on amount of harvest residue and shade from neighboring stands. In two of three years, trees on cooler microsites broke bud 2 to 4 days earlier than those on warmer microsites, despite receiving less heat forcing from March to May each year. A simple sequential model did not predict cooler sites having earlier bud burst nor did it correctly predict the order of bud burst across the three years. We modified the basic heat-forcing model to initialize, or reset to zero, the accumulation of heat units whenever significant freezing temperature events (> or = 3 degree-hours day(-1) < 0 degrees C) occurred; this modified model correctly predicted the sequence of bud burst across years. Soil temperature alone or in combination with air temperature did not improve our predictions of bud burst. Past models of bud burst have relied heavily on data from controlled experiments with simple temperature patterns; analysis of more variable temperature patterns from our 3-year field trial, however, indicated that simple models of bud burst are inaccurate. More complex models that incorporate chilling hours, heat forcing, photoperiod and the occurrence of freeze events in the spring may be needed to predict effects of future silvicultural treatments as well to interpret the implications of climate-change scenarios. Developing and testing new models will require data from both field and controlled-environment experiments.     

Slash pine bud dormancy as affected by lifting date and root wrenching in the nursery

Bud dormancy of root wrenched and unwrenched slash pine (Pinus elliottii Engelm.) seedlings growing in a forest nursery was measured on five lifting dates. Determination of bud dormancy was based on days to budbreak (DBB) under optimal growing conditions, mitotic activity in the apical meristem, chilling hours accumulated, and bud morphology. Based on DBB, seedlings were most dormant at Lift 2 on November 24 after exposure to 189 hours below 10 degrees C and 93 hours below 6.7 degrees C. Mitotic activity in the apical meristem was at its lowest 23 days later at Lift 3, possibly indicating the period when seedlings are most resistant to transplanting stresses. Multiple wrenching resulted in a slight shift in the dormancy cycle as wrenched seedlings set bud sooner in the nursery and broke bud sooner at the planting site in the spring than control seedlings. This implies that wrenched seedlings can be successfully lifted from the nursery earlier and will initiate spring shoot growth earlier than control seedlings.     

The minimum temperature for budburst in Betula depends on the state of dormancy

Vegis has put forward the theory that the range of growth-promoting temperatures changes during the induction and the release of dormancy. We have tested the response of buds of Betula pubescens Ehrh. and B. pendula Roth. on temperature during the induction and release of dormancy. Betula seedlings were exposed to dormancy-inducing high-temperature and short-day conditions and subsequently to dormancy-releasing chilling conditions in darkness. To monitor the dormancy status of the seedlings, subsets of them were transferred to five forcing temperatures and their budburst was observed. The results show that the expression of dormancy was temperature dependent, so that the minimum temperature for 100% budburst rose during the induction and dropped during the release of dormancy. These responses may explain previous contradictions between experimental and modelling studies, but that needs to be verified with more extensive experiments, some of which are identified in this study. The results provide further evidence for the concept of gradual change in bud dormancy. They also suggest that global change studies modelling budburst phenology should address the changing expression of bud dormancy.     

Soil temperature and plant growth stage influence nitrogen uptake and amino acid concentration of apple during early spring growth

In spring, nitrogen (N) uptake by apple roots begins about 3 weeks after bud break. We used 1-year-old 'Fuji' Malus domestica Borkh on M26 bare-root apple trees to determine whether the onset of N uptake in spring is dependent solely on the growth stage of the plant or is a function of soil temperature. Five times during early season growth, N uptake and total amino acid concentration were measured in trees growing at aboveground day/night temperatures of 23/15 degrees C and belowground temperatures of 8, 12, 16 or 20 degrees C. We used (15NH4)(15NO3) to measure total N uptake and rate of uptake and found that both were significantly influenced by both soil temperature and plant growth stage. Rate of uptake of 15N increased with increasing soil temperature and changed with plant growth stage. Before bud break, 15N was not detected in trees growing in the 8 degrees C soil treatment, whereas 15N uptake increased with increasing soil temperatures between 12 and 20 degrees C. Ten days after bud break, 15N was still not detected in trees growing in the 8 degrees C soil treatment, although total 15N uptake and uptake rate continued to increase with increasing soil temperatures between 12 and 20 degrees C. Twenty-one days after bud break, trees in all temperature treatments were able to acquire 15N from the soil, although the amount of uptake increased with increasing soil temperature. Distribution of 15N in trees changed as plants grew. Most of the 15N absorbed by trees before bud break (approximately 5% of 15N supplied per tree) remained in the roots. Forty-six days after bud break, approximately one-third of the 15N absorbed by the trees in the 12-20 degrees C soil temperature treatments remained in the roots, whereas the shank, stem and new growth contained about two-thirds of the 15N taken up by the roots. Total amino acid concentration and distribution of amino acids in trees changed with plant growth stage, but only the amino acid concentration in new growth and roots was affected by soil temperature. We conclude that a combination of low soil temperature and plant developmental stage influences the ability of apple trees to take up and use N from the soil in the spring. Thus, early fertilizer application in the spring when soil temperatures are low or when the aboveground portion of the tree is not actively growing may be ineffective in promoting N uptake.     

Variations in phenology and growth of European white birch (Betula pendula) clones

Phenology can have a profound effect on growth and climatic adaptability of northern tree species. Although the large interannual variations in dates of bud burst and growth termination have been widely discussed, little is known about the genotypic and spatial variations in phenology and how these sources of variation are related to temporal variation. We measured bud burst of eight white birch (Betula pendula Roth) clones in two field experiments daily over 6 years, and determined the termination of growth for the same clones over 2 years. We also measured yearly height growth. We found considerable genetic variation in phenological characteristics among the birch clones. There was large interannual variation in the date of bud burst and especially in the termination of growth, indicating that, in addition to genetic effects, environmental factors have a strong influence on both bud burst and growth termination. Height growth was correlated with timing of growth termination, length of growth period and bud burst, but the relationships were weak and varied among years. We accurately predicted the date of bud burst from the temperature accumulation after January 1, and base temperatures between +2 and -1 degrees C. There was large clonal variation in the duration of bud burst. Interannual variation in bud burst may have important consequences for insect herbivory of birches.