Effect of alternating day and night temperature on short day-induced bud set and subsequent bud burst in long days in Norway spruce.

Young seedlings of the conifer Norway spruce exhibit short day (SD)-induced cessation of apical growth and bud set. Although different, constant temperatures under SD are known to modulate timing of bud set and depth of dormancy with development of deeper dormancy under higher compared to lower temperature, systematic studies of effects of alternating day (DT) and night temperatures (NT) are limited. To shed light on this, seedlings of different provenances of Norway spruce were exposed to a wide range of DT-NT combinations during bud development, followed by transfer to forcing conditions of long days (LD) and 18°C, directly or after different periods of chilling. Although no specific effect of alternating DT/NT was found, the results demonstrate that the effects of DT under SD on bud set and subsequent bud break are significantly modified by NT in a complex way. The effects on bud break persisted after chilling. Since time to bud set correlated with the daily mean temperature under SD at DTs of 18 and 21°C, but not a DT of 15°C, time to bud set apparently also depend on the specific DT, implying that the effect of NT depends on the actual DT. Although higher temperature under SD generally results in later bud break after transfer to forcing conditions, the fastest bud flush was observed at intermediate NTs. This might be due to a bud break-hastening chilling effect of intermediate compared to higher temperatures, and delayed bud development to a stage where bud burst can occur, under lower temperatures. Also, time to bud burst in un-chilled seedlings decreased with increasing SD-duration, suggesting that bud development must reach a certain stage before the processes leading to bud burst are initiated. The present results also indicate that low temperature during bud development had a larger effect on the most southern compared to the most northern provenance studied. Decreasing time to bud burst was observed with increasing northern latitude of origin in un-chilled as well as chilled plants. In conclusion, being a highly temperature-dependent process, bud development is strongly delayed by low temperature, and the effects of DT is significantly modified by NT in a complex manner.

Effects of temperature and photoperiod on yield and chemical composition of northern and southern clones of bilberry (Vaccinium myrtillus L.).

After pollination outdoors, individual bilberry plants from two Northern and two Southern clones were studied for climatic effects on berry yield and quality in a controlled phytotrone experiment at 12 and 18 °C. At each temperature, the following light treatments were tested: (1) 12 h natural light, (2) 24 h natural light, and (3) 24 h natural light plus red light. The first experimental year there was no difference in yield between temperatures; however, the second experimental year the berry yields was significantly higher at 18 °C. Berry ripening was faster in the Northern than in the Southern clones at 12 °C. Northern clones also showed significantly higher contents of total anthocyanins, all measured anthocyanin derivatives, total phenolics, malic acid and sucrose. Metabolic profiling revealed higher levels of flavanols, hydroxycinnamic acids, quinic acid and carbohydrates at 12 °C.

The dynamic nature of bud dormancy in trees: environmental control and molecular mechanisms.

In tree species native to temperate and boreal regions, the activity-dormancy cycle is an important adaptive trait both for survival and growth. We discuss recent research on mechanisms controlling the overlapping developmental processes that define the activity-dormancy cycle, including cessation of apical growth, bud development, induction, maintenance and release of dormancy, and bud burst. The cycle involves an extensive reconfiguration of metabolism. Environmental control of the activity-dormancy cycle is based on perception of photoperiodic and temperature signals, reflecting adaptation to prevailing climatic conditions. Several molecular actors for control of growth cessation have been identified, with the CO/FT regulatory network and circadian clock having important coordinating roles in control of growth and dormancy. Other candidate regulators of bud set, dormancy and bud burst have been identified, such as dormancy-associated MADS-box factors, but their exact roles remain to be discovered. Epigenetic mechanisms also appear to factor in control of the activity-dormancy cycle. Despite evidence for gibberellins as negative regulators in growth cessation, and ABA and ethylene in bud formation, understanding of the roles that plant growth regulators play in controlling the activity-dormancy cycle is still very fragmentary. Finally, some of the challenges for further research in bud dormancy are discussed.

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.

Differential gene expression related to an epigenetic memory affecting climatic adaptation in Norway spruce.

In Norway spruce, the temperature during zygotic embryogenesis appears to adjust an adaptive epigenetic memory in the progeny that may regulate bud phenology and cold acclimation. Conditions colder than normal advance the timing whilst temperatures above normal delay the onset of these processes and altered performance is long lasting in progeny with identical genetic background. As a step toward unraveling the molecular mechanism behind an epigenetic memory, transcriptional analysis was performed on seedlings from seeds of six full-sib families produced at different embryogenesis temperature-cold (CE) vs warm (WE) under long and short day conditions. We prepared two suppressive subtracted cDNA libraries, forward and reverse, representing genes predominantly expressed in plants from seeds obtained after CE and WE embryogenesis following short day treatment (inducing bud set). Sequencing and annotation revealed considerable differences in the transcriptome of WE versus CE originated plants. By using qRT-PCR we studied the expression patterns of 32 selected candidate genes chosen from subtractive cDNA libraries analysis and nine siRNA pathways genes by a direct candidate approach. Eight genes, two transposons related genes, three with no match to Databases sequences and three genes from siRNA pathways (PaDCL1 and 2, PaSGS3) showed differential expression in progeny from CE and WE correlated with the family phenotypic differences. These findings may contribute to our understanding of the epigenetic mechanisms underlying adaptive changes acquired during embryogenesis.

Gene expression changes during short day induced terminal bud formation in Norway spruce.

The molecular basis for terminal bud formation in autumn is not well understood in conifers. By combining suppression subtractive hybridization and monitoring of gene expression by qRT-PCR analysis, we aimed to identify genes involved in photoperiodic control of growth cessation and bud set in Norway spruce. Close to 1400 ESTs were generated and their functional distribution differed between short day (SD-12 h photoperiod) and long day (LD-24 h photoperiod) libraries. Many genes with putative roles in protection against stress appeared differentially regulated under SD and LD, and also differed in transcript levels between 6 and 20 SDs. Of these, PaTFL1(TERMINAL FLOWER LIKE 1) showed strongly increased transcript levels at 6 SDs. PaCCCH(CCCH-TYPE ZINC FINGER) and PaCBF2&3(C-REPEAT BINDING FACTOR 2&3) showed a later response at 20 SDs, with increased and decreased transcript levels, respectively. For rhythmically expressed genes such as CBFs, such differences might represent a phase shift in peak expression, but might also suggest a putative role in response to SD. Multivariate analyses revealed strong differences in gene expression between LD, 6 SD and 20 SD. The robustness of the gene expression patterns was verified in 6 families differing in bud-set timing under natural light with gradually decreasing photoperiod.

Thermoperiodic growth control by gibberellin does not involve changes in photosynthetic or respiratory capacities in pea.

Active gibberellin (GA(1)) is an important mediator of thermoperiodic growth in pea. Plants grown under lower day than night temperature (negative DIF) elongate less and have reduced levels of GA(1) compared with plants grown at higher day than night temperature (positive DIF). By comparing the wild type (WT) and the elongated DELLA mutant la cry(s), this study has examined the effect of impaired GA signalling on thermoperiodic growth, photosynthesis, and respiration in pea. In the WT a negative DIF treatment reduced stem mass ratio and increased both root mass ratio and leaf mass ratio (dry weight of specific tissue related to total plant dry weight). Leaf, root and stem mass ratios of la cry(s) were not affected by DIF. Under negative DIF, specific leaf area (projected leaf area per unit leaf dry mass), biomass, and chlorophyll content of WT and la cry(s) plants were reduced. Young, expanding leaves of plants grown under negative DIF had reduced leaf area-based photosynthetic capacity. However, the highest photosynthetic electron transport rate was found in fully expanded leaves of WT plants grown under negative DIF. Negative DIF increased night respiration and was similar for both genotypes. It is concluded that GA signalling is not a major determinant of leaf area-based photosynthesis or respiration and that reduced dry weight of plants grown under negative DIF is caused by a GA-mediated reduction of photosynthetic stem and leaf tissue, reduced photosynthesis of young, expanding leaves, and reduced growth caused by low temperature in the photoperiod.

Gibberellin mediates daylength-controlled differentiation of vegetative meristems in strawberry (Fragaria x ananassa Duch).

BACKGROUND: Differentiation of long and short shoots is an important developmental trait in several species of the Rosaceae family. However, the physiological mechanisms controlling this differentiation are largely unknown. We have studied the role of gibberellin (GA) in regulation of shoot differentiation in strawberry (Fragaria x ananassa Duch.) cv. Korona. In strawberry, differentiation of axillary buds to runners (long shoot) or to crown branches (short shoot) is promoted by long-day and short-day conditions, respectively. Formation of crown branches is a prerequisite for satisfactory flowering because inflorescences are formed from the apical meristems of the crown. RESULTS: We found that both prohexadione-calcium and short photoperiod inhibited runner initiation and consequently led to induction of crown branching. In both cases, this correlated with a similar decline in GA1 level. Exogenous GA3 completely reversed the effect of prohexadione-calcium in a long photoperiod, but was only marginally effective in short-day grown plants. However, transfer of GA3-treated plants from short days to long days restored the normal runner formation. This did not occur in plants that were not treated with GA3. We also studied GA signalling homeostasis and found that the expression levels of several GA biosynthetic, signalling and target genes were similarly affected by prohexadione-calcium and short photoperiod in runner tips and axillary buds, respectively. CONCLUSION: GA is needed for runner initiation in strawberry, and the inhibition of GA biosynthesis leads to the formation of crown branches. Our findings of similar changes in GA levels and in GA signalling homeostasis after prohexadione-calcium and short-day treatments, and photoperiod-dependent responsiveness of the axillary buds to GA indicate that GA plays a role also in the photoperiod-regulated differentiation of axillary buds. We propose that tightly regulated GA activity may control induction of cell division in subapical tissues of axillary buds, being one of the signals determining bud fate.

Dehydrins expression related to timing of bud burst in Norway spruce.

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.

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.

Differential temperature regulation of GA metabolism in light and darkness in pea.

In greenhouse production of a number of flowering plant species, a short diurnal temperature drop in the morning is commonly used to reduce stem elongation. Earlier studies of pea (Pisum sativum) exposed to different combinations of day and night temperature, indicate that light, temperature, and gibberellin (GA) interact in the control of stem elongation. However, the mechanisms behind the effects of short-term temperature drops and differential sensitivity depending on the timing of the drop treatment have not been reported. Here, the involvement of GA metabolism in this has been investigated by exposing pea to short-term temperature drops in light or darkness. A 2 h temperature drop from 21 degrees C to 13 degrees C in the middle of the light period rapidly reduced the rate of stem elongation temporarily by 55% and increased mRNA levels of the GA-deactivation gene PsGA2ox2 by 2-fold within 30 min and up to 4-fold within 1.5 h. GA(1) levels were reduced by 36% after a 3-4 h time lag. A temperature drop in the night reduced stem elongation by 27%, but had no effect on transcript levels of PsGA2ox2. Instead, steady-state expression of the GA-biosynthesis genes NA, PsGA20ox1, and PsGA3ox1 was slightly stimulated, but there was no effect on GA(1) level. In conclusion, the effect of a temperature drop on GA metabolism in pea is qualitatively different in light and dark. Light is required for deactivation of GA(1) resulting from increased expression of PsGA2ox2. This suggests that GA-metabolism is a component in the short-term adaptation to changes in ambient temperature and putatively in low temperature-light stress responses.

Effects of red, far-red and blue light in maintaining growth in latitudinal populations of Norway spruce (Picea abies).

Seedlings of trees with a free growth pattern cease growth when night-lengths become shorter than a critical value, and this critical night-length (CNL) decreases with increasing latitude of origin. In northern populations, the light quality also appears to play an important role and a clinal variation in requirement for far-red (FR) light has been documented. In this study we dissected the light quality requirements for maintaining growth in different latitudinal populations of Norway spruce (Picea abies (L.) H. Karst.) using light emitting diodes for red (R), FR and blue (B) light, as 12 h day extension to provide 24 h photoperiod. At equal spectral photon flux, FR light was more effective than R light in maintaining growth, and the requirement of both R and FR increased with northern latitude of origin. One-to-one mixtures of R and FR light were more effective in maintaining growth than either FR or R light alone, indicating a possible interaction between R and FR light maintaining growth. Using the blue light as day extension could not prevent growth cessation in any of the populations, but delayed the bud set slightly in all populations. Our results suggest that phytochrome(s) are the primary photoreceptors in high irradiance responses maintaining growth in Norway spruce seedlings.

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.

Growth and physiological responses to supplemental UV-B radiation of two contrasting poplar species.

We compared the growth and physiological responses of cuttings of Populus kangdingensis C. Wang et Tung and P. cathayana Rehder originating from altitudes of 3500 m and 1500 m, respectively, when exposed to three ultraviolet-B (UV-B) radiation regimes: zero UV-B, ambient (4.5 kJ m(-2) day(-1)) and twice-ambient (9 kJ m(-2) day(-1)) biologically effective UV-B radiation for one growing season, to determine if Populus trees that are adapted to contrasting UV-B habitats exhibit different tolerances to enhanced UV-B radiation. Compared with cuttings grown without UV-B radiation, twice-ambient UV-B radiation significantly decreased total biomass, total leaf area and internode length in both species, whereas root/shoot ratio, leaf number, amount of photosynthetic pigments and ascorbate peroxidase activity were unaffected. Differences in responses were also observed between the study species. Cutting height increment, total biomass, total leaf area, free proline concentration and membrane damage assessed by electrolyte leakage were significantly more affected by the twice-ambient UV-B radiation in P. cathayana than in P. kangdingensis. However, specific leaf mass, amount of UV-B absorbing compounds and superoxide dismutase and guaiacol peroxidase activities increased more with increasing UV-B radiation in P. kangdingensis than in P. cathayana, perhaps reflecting important characteristics of species with low to moderate tolerance to UV-B radiation. Overall, the results indicated that P. kangdingensis, which originates from altitudes of 3500 m, has greater tolerance to enhanced UV-B radiation than P. cathayana originating from altitudes of 1500 m.

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.

Gibberellin structure and function: biological activity and competitive inhibition of gibberellin 2- and 3-oxidases.

Some gibberellin (GA) analogues, especially with C-16,17 modifications of GA(5), can inhibit growth of plants apparently by acting as competitors with the endogenous substrate of GA biosynthetic enzymes. Here, we directly confirm the competitive action of GA derivatives but also show that some analogues may retain significant bioactivity. A recombinant 3-oxidase from pea, which converts GA(20) to bioactive GA(1), was inhibited by GA(5), and 16,17-dihydro-GA(5) derivatives, especially if the C-17 alkyl chain length was increased by up to three carbons or if the C-13 hydroxyl was acetylated. Genetic confirmation that GA(5) analogues target 3-oxidases in vivo was provided by comparing the growth response of a WT (LE) pea with a 3-oxidase mutant (le-1). Two pea 2-oxidases that inactivate bioactive GAs, were inhibited by GA(1) and GA(3) but were generally insensitive to GA(5) analogues. alpha-Amylase production by barley half-seeds in response to GA analogues provided a method to study their action when effects on GA biosynthesis were excluded. This bioactivity assay showed that 16,17-dihydro GA(5) analogues have some inherent activity but mostly less than for GA(5) (5-50-fold), which in turn was 100-fold less active than GA(1) and GA(3). However, although C-17 alkyl derivatives with one or two added carbons showed little bioactivity and were purely 3-oxidase inhibitors, adding a third carbon (the 17-n-propyl-16,17-dihydro GA(5) analogue) restored bioactivity to that of GA(5). Furthermore, this analogue has lost its capacity to inhibit stem elongation of Lolium temulentum (Mander et al., Phytochemistry 49:1509-1515, 1998a), although it strongly inhibits the 3-oxidase. Thus, the effectiveness of a GA derivative as a growth retardant will reflect the balance between its bioactivity and its capacity to inhibit the terminal enzyme of GA biosynthesis. The weaker growth inhibition in dicots including pea (approximately 10%) than in monocots such as L. temulentum (>35%) is suggestive of taxonomic differences in the bioactivity of GAs and/or their effects on GA biosynthesis.

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.

Independent activation of cold acclimation by low temperature and short photoperiod in hybrid aspen.

Temperate zone woody plants cold acclimate in response to both short daylength (SD) and low temperature (LT). We were able to show that these two environmental cues induce cold acclimation independently by comparing the wild type (WT) and the transgenic hybrid aspen (Populus tremula x Populus tremuloides Michx.) line 22 overexpressing the oat (Avena sativa) PHYTOCHROME A gene. Line 22 was not able to detect the SD and, consequently, did not stop growing in SD conditions. This resulted in an impaired freezing tolerance development under SD. In contrast, exposure to LT resulted in cold acclimation of line 22 to a degree comparable with the WT. In contrast to the WT, line 22 could not dehydrate the overwintering tissues or induce the production of dehydrins (DHN) under SD conditions. Furthermore, abscisic acid (ABA) content of the buds of line 22 were the same under SD and long daylength, whereas prolonged SD exposure decreased the ABA level in the WT. LT exposure resulted in a rapid accumulation of DHN in both the WT and line 22. Similarly, ABA content increased transiently in both the WT and line 22. Our results indicate that phytochrome A is involved in photoperiodic regulation of ABA and DHN levels, but at LT they are regulated by a different mechanism. Although SD and LT induce cold acclimation independently, ABA and DHN may play important roles in both modes of acclimation.

Far red end-of-day treatment restores wild type-like plant length in hybrid aspen overexpressing phytochrome A.

Shoot elongation in woody plants is modulated by a multitude of light signals, including irradiance, photoperiod and spectral composition, for which the phytochrome system is the probable photoreceptor. In hybrid aspen (Populus tremula x tremuloides) overexpression of the oat phytochrome A (PHYA) prevents growth cessation in response to short photoperiod, and plants exhibit dwarf growth that is related to reduced cell numbers and reduced gibberellin contents. End-of-day far-red treatment significantly enhances internode elongation in PHYA overexpressors as well as in the wild type, and this was found here to be caused by stimulation of cell division and cell extension. In PHYA overexpressors the effects were substantially larger than in the wild type, and resulted in complete restoration of wild type-like plant length as well as cell numbers, and gibberellin content was greatly increased. No clear effect of far-red end-of-day treatment on gibberellin levels could be detected in the wild type. It thus appears that the far-red end-of-day treatment might modify the responsiveness of the tissue to GA rather than the GA levels. The observed effects were completely reversed by a subsequent irradiation with red light. The present data show that dwarfism due to PHYA overexpression can be completely overcome by far red end-of-day treatment, and the observations indicate that effects of far red end-of-day treatments appear to be mediated by phytochrome(s) other than phytochrome A.