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.     

High autumn temperature delays spring bud burst in boreal trees, counterbalancing the effect of climatic warming

The effect of temperature during short-day (SD) dormancy induction was examined in three boreal tree species in a controlled environment. Saplings of Betula pendula Roth, B. pubescens Ehrh. and Alnus glutinosa (L.) Moench. were exposed to 5 weeks of 10-h SD induction at 9, 15 and 21 degrees C followed by chilling at 5 degrees C for 40, 70, 100 and 130 days and subsequent forcing at 15 degrees C in a 24-h photoperiod for 60 days. In all species and with all chilling periods, high temperature during SD dormancy induction significantly delayed bud burst during subsequent flushing at 15 degrees C. In A. glutinosa, high temperature during SD dormancy induction also significantly increased the chilling requirement for dormancy release. Field experiments at 60 degrees N with a range of latitudinal birch populations revealed a highly significant correlation between autumn temperature and days to bud burst in the subsequent spring. September temperature alone explained 20% of the variation between years in time of bud burst. In birch populations from 69 and 71 degrees N, which ceased growing and shed their leaves in August when the mean temperature was 15 degrees C, bud burst occurred later than expected compared with lower latitude populations (56 degrees N) in which dormancy induction took place more than 2 months later at a mean temperature of about 6 degrees C. It is concluded that this autumn temperature response may be important for counterbalancing the potentially adverse effects of higher winter temperatures on dormancy stability of boreal trees during climate warming.     

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.     

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.     

Modeling the effects of winter environment on dormancy release of Douglas-fir

Most temperate woody plants have a winter chilling requirement to prevent budburst during mid-winter periods of warm weather. The date of spring budburst is dependent on both chilling and forcing; modeling this date is an important part of predicting potential effects of global warming on trees. There is no clear evidence from the literature that the curves of chilling or forcing effectiveness differ by species so we combined our data and published information to develop new curves on the effectiveness of temperature for chilling and forcing. The new curves predict effectiveness over a wide range of temperatures and we suggest both functions may be operating at the same time. We present experimental data from 13 winter environments for 5 genotypes of Douglas-fir (Pseudotsuga menziesii var. menziesii) and use them to test various assumptions of starting and stopping dates for accumulating chilling and forcing units and the relationship between budburst and the accumulation of chilling and forcing units. Chilling started too early to be effective in one treatment but the other 12 environments resulted in budburst from many combinations of chilling and forcing. Previous reports have suggested benefits or cancellations of effects from alternating day/night or periodic temperatures. Our simple models do not include these effects but nevertheless were effective in predicting relationships between chilling and forcing for treatments with a wide range of conditions. Overall, the date of budburst changed only slightly (+1 to ?11 days) across a wide range of treatments in our colder test environment (Olympia, WA, USA) but was substantially later (+29 days) in the warmest treatment in our warmer environment (Corvallis, OR, USA). An analysis of historical climate data for both environments predicted a wide range in date to budburst could result from the same mean temperature due to the relative weightings of specific temperatures in the chilling and forcing functions. In the absence of improved understanding of the basic physiological mechanisms involved in dormancy induction and release, we suggest that simple, universal functions be considered for modeling the effectiveness of temperature for chilling and forcing. Future research should be designed to determine the exact shape of the curves; data are particularly lacking at the temperature extremes. We discuss the implications of our data and proposed functions for predicting effects of climate change. Both suggest that the trend toward earlier budburst will be reversed if winter temperatures rise substantially.     

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.     

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.     

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.     

Frost tolerance and bud dormancy of container-grown yellow birch, red oak and sugar maple seedlings

Container-grown seedlings of red oak (Quercus rubra L.), sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britton) in their first year of growth were overwintered outdoors. Tolerance of roots and stems to freezing was compared from late summer to the following spring. Mitotic activity in the apical bud was related more closely to air temperature than to bud dormancy as defined by days to bud break. In all species, stem hardening was observed before days to bud break reached a maximum. Dormancy release (days to bud break equal to zero) of yellow birch coincided with loss of stem hardening in the spring. Roots hardened more slowly, had a lower frost tolerance than stems in fall and winter, and dehardened earlier than stems in the spring. There were differences in stem and root hardiness among the species, with yellow birch being the most tolerant, followed by sugar maple and red oak. Primarily because of root sensitivity to frost, winter was a critical period for all three species, but particularly for red oak.     

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.     

Seasonal changes in intensity of bud dormancy in loblolly pine seedlings

The terminal buds of six-month-old loblolly pine (Pinus taeda L.) seedlings remained closed for approximately six months, although bud dormancy, as measured by rate of bud break in a standard greenhouse environment, was only exhibited for about one month. The peak of bud dormancy was in December for seedlings grown near Auburn, Alabama. However, the timing and intensity varied with seed source and may have been affected by the warm fall temperatures. Seedlings from the more northern provenances entered dormancy first and reached a deeper state of dormancy than seedlings from southern provenances. The rate of shoot elongation was not consistently related to the rate of bud break.     

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.     

Short-day photoperiods affect expression of genes related to dormancy and freezing tolerance in Norway spruce seedlings

Key Message

Gene expression analysis showed that prolonged short day (SD) treatment deepened dormancy and stimulated development of freezing tolerance of Picea abies seedlings. Prolonged SD treatment also caused later appearance of visible buds in autumn, reduced risks for reflushing, and promoted earlier spring bud break.

Context

Short day (SD) treatment of seedlings is a common practice in boreal forest tree nurseries to regulate shoot growth and prepare the seedlings for autumn planting or frozen storage.

Aims

The aim of this study was to examine responses of Norway spruce (Picea abies (L.) Karst.) to a range of SD treatments of different length and evaluate gene expression related to dormancy induction and development of freezing tolerance.

Methods

The seedlings were SD treated for 11 h a day during 7, 14, 21, or 28 days. Molecular tests were performed, and the expression profiles of dormancy and freezing tolerance-related genes were analyzed as well as determination of shoot growth, bud set, bud size, reflushing, dry matter content, and timing of spring bud break.

Results

The 7-day SD treatment was as effective as longer SD treatments in terminating apical shoot growth. However, short (7 days) SD treatment resulted in later activation of dormancy-related genes and of genes related to freezing tolerance compared to the longer treatments which had an impact on seedling phenology.

Conclusion

Gene expression analysis indicated an effective stimulus of dormancy-related genes when the SD treatment is prolonged for at least 1–2 weeks after shoot elongation has terminated and that seedlings thereafter are exposed to ambient outdoor climate conditions.

     

Stock quality assessment through an integrated approach

Forest regeneration can be improved by matching seedling stock with reforestation site environmental conditions. To achieve this improvement, one must characterize seedling performance potential with anticipated field site conditions. A theoretical performance potential index (PPI) has been developed to forecast seedling field performance during the first growing season on a reforestation site. PPI includes material and performance attribute tests that characterize seedling performance potential under optimum and limiting environmental conditions. Test results are standardized and then integrated resulting in a comprehensive assessment procedure. The PPI concept is demonstrated using containerized western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedlings subjected to a series of dormancy induction treatments and the results are compared with actual seedling performance during the first growing season on a reforestation site. Results indicated that PPI values for western hemlock seedlings treated with various dormancy induction treatments corresponded to field performance. Possible use of PPI in stock selection is discussed.     

Seed germination and seedling development in the mango (Mangifera indica L.)

Mango (Mangifera indica L., cv Ruby) seeds taken from ripe fruit showed no dormancy. They germinated at temperatures between 5 and 40 degrees C, but germination was most rapid near the upper end of this range (25-40 degrees C). The fresh seeds had a high moisture content (85%, dry weight basis) and quickly died on dehydration. The optimal temperature for growth of the seedlings was close to 30 degrees C. High temperatures (40 degrees C) and temperatures below 15 degrees C were lethal. Growth of the stem occurred in successive flushes separated by rest periods. When the leaves of the preceding flush finished growing, the axis lengthened beneath the apical bud.     

Relationships among bud dormancy status,cold hardiness,and stress resistance in 2+0 Douglas-fir

An experiment was conducted to ascertain what relationships might exist among dormancy status, cold hardiness and stress resistance in 2+0 Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), seedlings. Seedlings were lifted from a western Washington nursery on six dates spanning the 1980–81 lifting season. On each date samples of seedlings were subjected to the following treatment: (1) tumbling for 5 minutes, (2) desiccation of roots for 30 minutes at 30°C and 2.1 kPa vapor pressure deficit, (3) exposure of shoots to temperatures of –10°C, –15°C or –20°C for two hours and (4) unstressed control. On two lift dates sub-samples of seedlings were placed into –1°C storage and held for two months before the above stress treatments were administered. Bud dormancy status was determined, using a bud break test, on seedlings from each lift date before and after storage.After one growing season in the field percent survival, vigor, height growth and shoot and root weight were determined on stressed and unstressed seedlings. Survival and vigor were less affected by the stress treatments than were height and weight. Severity of stress was in the order –20°C > –15°C > desiccation > handling > –10°C. Degree of cold injury was directly related to seedling dormancy status whether dormancy status had been attained in the nursery from natural chilling or in frozen storage. Seedlings in a mid-range of dormancy release (between deep rest and quiescence) were most resistant to all imposed stresses.