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.     

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 (Kittil?, 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.     

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.     

Effects of cold-hardening on compatible solutes and antioxidant enzyme activities related to freezing tolerance in Ammopiptanthus mongolicus seedlings

Cold acclimation is associated with many metabolic changes that lead to an increase of freezing tolerance. In order to investigate the biochemical process of cold acclimation in Ammopiptanthus mongolicus, seedlings were acclimated at 2℃ under 16-h photoperiod （150 μmol·m^-2·s^-1 photosynthetically active radiation） for 14 d. Freezing tolerance in seedlings increased after 14 d of cold-hardening. Contents of protein, proline and solute carbohydrate in cotyledon increased after cold acclimation. Patterns of isozymes of superoxide dismutase （SOD）, peroxidase, catalase and polyphenol oxidase （PPO） were investigated. The activities of SOD, peroxidase and PPO in cold acclimated plants were increased during cold-hardening. We deduced that compatible solutes and antioxidant enzymes play important roles in development of freezing tolerance during cold acclimation in this evergreen woody plant.     

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.     

A molecular marker associated with low-temperature induction of dormancy in red osier dogwood (Cornus sericea)

Dormancy induction in temperate deciduous plants is thought to be regulated by short photoperiods, but low temperature has been shown to eliminate the short photoperiod requirement in northern ecotypes. An F2 population (191 plants) red osier dogwood (Cornus sericea L.) derived from a polycross of an F1 population produced from reciprocal crosses of the parental clonal ecotypes, Northwest Territories (NWT, 62 degrees N) and Utah (42 degrees N), was examined to identify molecular markers of temperature-induced endodormancy. Dormancy induction curves were generated for each individual in the F2 population and a standard point prior to vegetative maturity (i-VM) was inferred from the change in slope of the dormancy acquisition curve. Under Saskatoon, Saskatchewan field conditions (52 degrees N), the NWT ecotype entered i-VM on average 5-6 weeks before the Utah ecotype. Two sub-populations of the F2 population were distinguishable based on VM acquisition on exposure to low temperature but not to short photoperiods. A sequence characterized amplified region (SCAR) marker was developed that correctly (> 92%) identified individual plants within the F2 subpopulation that were responsive to low-temperature induction of VM. Timing of bud break was strongly associated with the timing of VM in the geographical ecotypes but not in the F2 population, indicating that these are separate traits under genetic control.     

毛白杨抗冻性低温诱导过程中糖的效应(英文)

在低温锻炼和结合蔗糖处理的低温锻炼下 ,对毛白杨 (Populustomentosa)幼苗枝条和叶片的可溶性糖含量 ,以及幼苗存活率和抗冻性分别进行了测定 .结果表明 ,低温锻炼明显提高了幼苗枝条和叶片的可溶性糖含量 ,同时也提高了幼苗的存活率和抗冻性 .2 0 %蔗糖处理对上述低温锻炼的作用有加强的效果 .进一步分析发现 ,无论是单纯的低温锻炼还是结合蔗糖预处理的低温锻炼 ,幼苗枝条中可溶性糖含量的提高程度均较叶片明显 ,而且它们的含量增加与幼苗抗冻性的提高密切相关 .这表明低温锻炼所引起的可溶性糖积累可能参与了幼苗抗冻性的诱导     

Influence of ozone on cold acclimation in sugar maple seedlings

During summer 1994, sugar maple (Acer saccharum Marsh.) seedlings were grown in open-top chambers supplied with air containing near ambient ozone concentration (control, low O(3)) or three times the ambient ozone concentration (high O(3)). The rate of CO(2) assimilation was significantly reduced by chronic exposure to a high concentration of ozone during the summer. During fall, seedlings were removed from the open-top chambers and acclimated to cold under natural conditions. In both species during cold acclimation, the starch concentration decreased, whereas the sucrose concentration increased. There was no treatment effect on the freezing tolerance of roots, even though roots in the high-O(3) treatment accumulated higher concentrations of the cryoprotective oligosaccharides raffinose and stachyose than control roots. Cold acclimation occurred earlier and stachyose concentration of stems was higher in high-O(3)-treated seedlings than in low-O(3)-treated seedlings. Cold acclimation was associated with an earlier accumulation of ABA in the xylem sap of high-O(3)-treated seedlings compared with low-O(3)-treated seedlings.     

Atmospheric carbon dioxide enrichment reduces carbohydrate and nitrogen reserves in overwintering Picea mariana

Abstract

Elevated levels of atmospheric carbon dioxide (CO₂) can directly affect the cold hardening process in evergreens through their effect on the accumulation of carbon and nitrogen reserves. This study investigated the biochemical responses of black spruce [Picea mariana (Mill.) B.S.P.] seedlings to CO₂ enrichment during growth, cold hardening and dehardening. Seedlings were grown under 350 (ambient) or 710 (elevated) ppm of CO₂ for 12 months in eight mini-greenhouses. Photoperiod and temperature were gradually lowered in autumn to induce cold hardening, and the conditions were reversed in spring to promote dehardening. At regular intervals, cold tolerance was assessed and sugars, starch and amino acid concentrations were measured. The freezing tolerance differed between the two treatments only in early autumn, with seedlings growing under high CO₂ being more tolerant. The northern ecotype was more cold tolerant with concomitant higher concentrations of sucrose, fructose, pinitol, glucose and total soluble sugars. The concentration of soluble sugars increased in needles and roots of black spruce along with cold hardening, and the concentrations of the cryoprotective sugars sucrose and raffinose were lower under elevated CO₂. Amino acid concentrations were also lower under elevated than under ambient CO_2. The lower level of reserve did not translate into a lower level of freezing tolerance.     

Bud dormancy in beech (Fagus sylvatica L.). Effect of chilling and photoperiod on dormancy release of beech seedlings

Two-year-old Fagus sylvatica L. seedlings were subjected to natural winter chilling or were overwintered in a heated greenhouse. Plants were then grown in controlled environment chambers with photoperiods of 9 or 13 h. Renewal of bud growth was found to be mainly determined by winter chilling. There was a slight interaction between chilling and photoperiod. Sprouting of apical buds took two to three times as long in unchilled plants as in chilled plants. Shoot elongation was influenced by chilling and was also greater in the 13-h photoperiod than in the 9-h photoperiod, but this may have been due at least in part to the higher irradiance. Chilling resulted in rapid dormancy loss and changed the growth pattern from basitonal to acrotonal.     

Growth and reproduction in <Emphasis Type="Italic">Pinus sibirica</Emphasis> ecotypes from Western Siberia in a common garden experiment

Experimental studies of forest trees’ intraspecies variations in reproduction and reproductive allocation are scarce. Such studies are relevant for the Siberian stone pine (Pinus sibirica Du Tour) because of the high value of its seeds and wood. To reveal growth and reproductive variations among Siberian stone pine ecotypes in relation to the climate of the provenances, we examined their growth height, crown diameter, stem volume, number of female cones, and male shoots, at different ages, in the clonal common garden experiment established in 1996 in Russia. Scions were obtained from 11 to 19 randomly selected 100–170-year old trees in each of four natural stands located on the latitudinal transect on the West Siberian plain. They were grafted onto local 4 year old Siberian stone pine seedlings and grown in a common garden experiment comprising nine plots near the southern margin of the species’ natural range. Growth traits were affected by climatic conditions in the provenances: generally, a warmer climate was associated with more rapid growth. Variation among the ecotypes at the graft age of 12 years was not large; however, it significantly increased by the age of 19. The relationship was ambiguous between reproductive traits and the climate of the provenances. The proportion of female, male and bisexual clones varied among years in all ecotypes. Most of the clones had only female cones; male clones were less common. The southern ecotype showed rapid growth and formed the lowest number of male shoots and female cones. The northern ecotype showed the weakest growth and moderate cone bearing, along with abundant male flowering. The intermediate ecotypes had a moderate growth and the highest numbers of cones, along with moderate numbers of male shoots. These results demonstrate the fundamental differences in the patterns and nature of variation in growth and reproductive traits.     

Influence of photoperiod and temperature on frost hardiness and free amino acid concentrations in black spruce seedlings

We investigated the effects of a 4-week exposure to an 8-h or 18-h photoperiod at 5 or 25 degrees C on the development of hardiness to -20 degrees C and the accumulation of proline (Pro), arginine (Arg) and tryptophan (Trp) in shoots of black spruce (Picea mariana (Mill.) B.S.P.) seedlings. The greatest degree of hardening to -20 degrees C occurred in seedlings exposed to an 8-h photoperiod at 25 degrees C, and some hardening occurred in seedlings exposed to 5 degrees C in either an 8-h or 18-h photoperiod. Proline accumulated in shoots in response to 5 degrees C and either an 8-h or 18-h photoperiod, whereas Trp accumulated in response to an 8-h photoperiod at either temperature, and Arg only accumulated in shoots in the 5 degrees C + 8-h photoperiod treatment. Only the accumulation of Trp was significantly related to the degree of hardiness to -20 degrees C.     

植物抗寒性的生理生态学机制研究进展

综述植物在冷驯化过程中发生的一系列生理生化变化.环境对植物抗寒性的影响主要与光诱导、温湿度以及气候的变化有关.植物表面形成冰层会引起植物的无氧呼吸,导致植物受害;光抑制诱导活性氧的产生,从而导致植物光合系统的退化,抗寒能力下降,而短日照诱导植物休眠,有利于植物抗寒.光敏色素则被认为是启动冷驯化的光受体;植物通过冷驯化增加碳水化合物的积累及病原体相关蛋白的合成,以增强对低温病原体的抵抗能力;气候的变化使植物遭受了更大的冷伤害风险.微管最初遇冷时部分的解体可以有效诱导植物抗寒性;抗氧化酶活性增强,植物体内糖、脯氨酸、多胺等内含物含量上升.植物休眠状态中的生理变化(种子的休眠、芽的休眠)与ABA敏感性的差异有关.对植物抗寒性分子机制的研究表明:COR基因的表达对于植物抗寒性和冷驯化是十分关键的;与气候梯度有关的基因梯度的分布说明寒冷地区的树种更为抗寒;多表型性状的数量性状分析,为重要的农艺性状标记辅助选择(MAS)提供基础.对植物抗寒过程中的信号转导进行研究发现,Ca2 是低温下参与调节冷驯化应答机制中信号转导途径的重要的第二信使.未来植物抗寒领域的研究热点为信号转导和基因调节,低温抗性的遗传学和遗传应用及代谢组学,气候变化对于植物抗寒的影响等方面.     

Use of short-day treatment in the production of Norway spruce mini-plug seedlings under plant factory conditions

Mini-plug transplant seedlings of Norway spruce have been cultivated in closed growth systems, so-called plant factories, for few years. The aim of the experiment was to define a short-day treatment (SD) that harden seedlings to sustain 3 months of cold storage, but does not have adverse effects on growth, morphology, and vitality. The seedlings were subjected to one of the following treatments: (1) 12?h photoperiod + 3 weeks duration; (2) 8?h photoperiod + 3?wk duration; (3) 12?h photoperiod + 5?wk duration; and (4) 8?h photoperiod + 5?wk SD. All the SD treatments yielded healthy seedlings that grew well after the cold storage. The frost hardiness of the seedlings improved when the photoperiod was reduced from 12 to 8?h, and when the SD duration was increased from 3 weeks to 5 weeks, but reducing the photoperiod from 12?h to 8?h caused growth reductions. The root and shoot regrowth after cold storage was highest in seedlings that had received 12?h photoperiod and 5?wk duration. However, 12?h photoperiod and 3?wk duration may be an adequate practice for nurseries that treat multiple crops in their SD facilities.     

Characterizing the frost sensitivity of black spruce photosynthesis during cold acclimation

We used photosynthetic light response curves to measure and model the responses of two provenances of 3-year-old black spruce (Picea mariana (Mill.) BSP) seedlings to severe artificial frost treatments applied at 2-week intervals during cold acclimation. Black spruce seedlings responded to cold acclimation with long-term suppression of photosynthetic capacity (Amax) and apparent quantum-use efficiency (alpha'). Short-term reductions in both photosynthetic parameters following frost treatments were dependent on the extent of cold acclimation of the seedlings and the severity of the frost treatments. Large reductions in Amax in response to the frost treatments were observed in seedlings that had undergone little cold acclimation and these reductions were associated with an irreversible reduction in alpha'. Such seedlings recovered only partially during the subsequent 23 days, whereas seedlings in most other treatments showed complete recovery of Amax after 13 days. The impact of frost treatments on Amax and alpha' did not vary with seedling provenance. We propose an algorithm that predicts the combined effects of cold acclimation and severe freezing temperatures on the extent of the suppression of A(max) during autumn. The algorithm is based on (1) the maximum Amax observed during the growing season, (2) the accumulation of cold degree-days, based on a minimum nocturnal temperature < 5 degrees C, and (3) the severity of freezing temperatures during autumn. The parameters developed in the algorithm showed that cold acclimation of black spruce seedlings had a greater impact on the reduction of Amax in autumn than did the severe frost treatments. Mean Amax of seedlings subjected to artificial frosts showed a strong correlation with values predicted by the algorithm (r2 = 0.91).     

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.

     

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.     

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.     

低温锻炼与CaCl2处理对毛白杨幼苗可溶性总蛋白、CaM及抗冻性的影响

低温锻炼提高了毛白杨幼苗叶片和枝条中钙调素 (CaM)和可溶性总蛋白含量 ,同时也提高了幼苗的抗冻性 .CaCl2 处理对上述低温锻炼的作用有加强的效果 ,体现在明显提高了CaM和可溶性总蛋白含量以及幼苗抗冻性 ;钙离子螯合剂EGTA、钙离子通道阻断剂LaCl3 或钙调素拮抗剂CPZ处理则减弱了低温锻炼的效应 ,表现出CaM和可溶性总蛋白含量以及幼苗抗冻性明显降低 .进一步研究发现 ,CaCl2 处理的低温锻炼不仅能减轻低温胁迫引起的CaM和可溶性总蛋白含量的下降程度 ,而且有利于幼苗在恢复过程中CaM和可溶性总蛋白水平的迅速回升 .这表明Ca2 + CaM可能参与了CaM和可溶性总蛋白的合成及抗冻性的诱导