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.     

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.     

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.     

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.     

Frost hardiness, bud phenology and growth of containerized Picea mariana seedlings grown at three nitrogen levels and three temperature regimes

We studied the influence of temperature and near- and sub- optimal mineral nutrition of black spruce seedlings (Picea mariana [Mill.] B.S.P.) during their second growing period on bud set, bud development, growth, mineral content and cold tolerance. Bud break and growth after bud break were also studied. Seedlings were grown for 106 d in growth chambers under three temperature regimes in combination with three concentrations of a fertilizer. They were then cold hardened for 56 d and dehardened for 66 d.Under these near- and sub-optimal N levels, bud formation occurred during the growing season. Bud formation was accelerated with decreasing fertilization, but was not affected by temperature treatments. Needles from seedlings with 0.64% N (dry mass basis) before hardening did not harden. Those with 0.87% N showed a lesser degree of hardiness than those with 1.28% N. Stem diameter increased at the beginning of the hardening period. During this acclimation period, shoot dry mass decreased with time at a constant rate and at the same rate over time for all treatments whereas root dry mass was more variable. Total number of needle primordia was low and no difference was observed among growing conditions. Bud break was similar in all treatments. Following bud break, shoot height and stem diameter increases were small but their magnitude varied with the nutritional regimes applied during the previous growing period. During hardening, nitrogen concentration of shoot tissues first increased and then decreased; phosphorus concentration first increased and then remained stable; potassium concentration remained stable. Concentration of these three elements generally decreased in the roots during this hardening.     

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.     

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.     

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 bud dormancy in relation to bud morphology, water and starch content, and abscisic acid concentration in adult trees of Betula pubescens

Annual cycles of change in bud morphology, bud burst ability, abscisic acid (ABA) concentration, and starch and water content were studied in mid-crown terminal buds of short shoots and underground basal buds of Betula pubescens Ehrh. In particular, we investigated the roles of ABA and bud water content in the regulation of bud growth. Basal buds differed morphologically from terminal buds of short shoots in that their leaf initials did not develop into embryonic foliage leaves and their total size did not increase significantly during summer. Bud burst ability, measured by forcing detached short shoots and stumps under controlled conditions, was maintained in the basal buds throughout the year, whereas the terminal buds of short shoots remained dormant until October, thereafter their bud burst ability increased gradually and reached a maximum in March-April. The ABA concentration of the basal buds was relatively constant throughout the sampling period (1-3 micro g g(DW) (-1)), whereas that of the terminal buds of short shoots, which was much higher (5-10 micro g g(DW) (-1)), showed a distinct seasonal cycle with a maximum from August to November. Bud ABA concentration decreased during the first 10 days of forcing, especially in basal buds. In both bud types, the amount of starch increased toward the autumn, declined in November, and was negligible in the terminal buds of short shoots between January and March, but in April, the amount was high again in both bud types. Water content varied characteristically in both bud types, although more distinctly in the terminal buds of short shoots, with an increase in spring before bud burst and a decrease during the summer until September. The significant morphological and physiological differences between the mid-crown terminal buds of short shoots and the underground basal buds may partly explain the characteristic growth habit of the basal buds and their development into coppice shoots after cutting the tree. The results also indicate a role for ABA in maintaining dormancy of the terminal buds of short shoots and emphasize the relationship between tissue water status and ABA concentration.     

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.

     

Seasonal alteration in amount of Ca(2+) in apical bud cells of mulberry (Morus bombciz Koidz): an electron microscopy-cytochemical study

Subcellular localization of calcium in apical bud cells of mulberry (Morus bombciz Koidz) was investigated by a combination of calcium antimonate precipitation and electron microscopy (EM), over a 10-month period extending from July to May. Calcium antimonate deposits, an indication of Ca(2+) localization, were found mainly in the intercellular spaces and vacuoles of tissues collected in summer (e.g., July 10). Few deposits were seen in the cytosol and the nucleus, indicating that when plants were actively growing, the cytoplasm had a low concentration of Ca(2+). As the day length became shorter (e.g., August 8), Ca(2+) deposits increased in the cytosol and the nucleus. When plant dormancy and cold hardiness were rapidly developing in September and October, a large number of Ca(2+) deposits were seen in the cytoplasm and the nucleus, where they remained for about 60 days. Tissue samples collected during midwinter (e.g., January 20) displayed few Ca(2+) deposits in the cytoplasm and the nucleus compared with the sample collected on November 1. The number of Ca(2+) deposits remained low even when day length increased in the spring (e.g., May 5). We conclude that the seasonal dynamics of intracellular Ca(2+) concentration bear a close relationship with growth cessation and the development of dormancy and cold hardiness in temperate woody perennials.     

平榛冷适应相关基因CBF的克隆及时空表达特性分析

低温是影响植物分布、进而影响生长和产量的关键因素之一。Weiser(1970)指出低温导致植物的基因表达发生变化;早在20世纪90年代研究者就发现了植物的冷适应现象(Guyetal.,1985)。虽然迄今为止植物抗寒的分子机制还没完全清楚,但研     

Cold acclimation induced accumulation of phenolic compounds and freezing tolerance in Ammopiptanthus mongolicus

Ammopiptanthus mongolicus, the only freezing tolerant evergreen broad-leaved shrub, local species of the Alashan desert, northwest sand area of China, can survive -30°C or even lower temperature in winter. In the present study, the secondary products phenolics in A. mongolicus cotyledons were determined to study the effects of phenolics on cold tolerance. Cytochemical localization of phenolics in cotyledon cells was observed by electron microscopy and the content of phenolic compounds was assayed by spec-trophotometric measurement. The results showed that the freezing tolerance of A. mongolicus seedlings increased after acclimation at 2-6°C for 14 days, which accompanied the increase of the content of phenolic compounds in cotyledons. Cytochemical observation showed that phenolic deposits were mainly localized in vacuoles and in close proximity to tonoplast, and also in the cytoplasm. The amount and the size of phenolics droplets increased obviously in cytoplasm and vacuoles after cold acclimation, predominantly ag-gregated along membranes of vacuoles and tonoplast. No phenolic deposits were found in cell walls. As hydrogen-or elec-tron-donating agents, phenolics may protect plant cells against reactive oxygen species formed during chilling or freezing stress and improve the freezing tolerance of cold-acclimated A. mongolicus seedlings.     

First frost: Effects of single and repeated freezing events on acclimation in Picea abies and other boreal and temperate conifers

In experiments with needles of Picea abies, we tested the specific hypothesis that a single night of freezing acts as a signal that triggers a rapid increase in low temperature (LT) tolerance, and the more general hypothesis that repeated or prolonged freezing stimulates increased LT acclimation. In three growth chamber experiments involving acclimation under early- to mid-autumn light and temperature conditions followed by one or more freezing treatments, we found no significant effect of a single night of freezing on LT tolerance, and only limited and inconsistent effects of repeated and prolonged freezing. We also tested the effect of prolonged storage at −5 °C on LT tolerance on samples of three boreal and three temperate conifer species during acclimation under field conditions, and again found no consistent enhancement of LT tolerance attributable to freezing in either group. In agreement with our own and others’ anecdotal observations that some species can attain nearly maximal LT tolerance in the absence of freezing under field conditions, we conclude that freezing is neither required nor a major influence in LT acclimation, at least in well-studied boreal conifer species, while the effects of freezing on temperate conifers are not as well-documented. We conclude that freezing treatment of conifer seedlings to ensure sufficient hardiness for late planting seems to offer little practical advantage.     

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).     

Frost hardening and dehardening in Abies procera and other conifers under differing temperature regimes and warm-spell treatments

Frequent bud frost damage in cultivation of Abies procera Rehderand pending climate changes are the background for this studyof cold hardiness under varying acclimation regime (in closed-topchambers) and experimental warm spells during the cold season.LT₅₀ values were established by freezing tests at differenttimes of year. Damage and deaths were assessed on leader buds,subapical lateral buds, needles and cambium. Minor parallelexperiments involved Abies nordmanniana, Picea abies and Piceasitchensis. Lower acclimation temperatures resulted in deeperfrost hardiness during late autumn but less during spring, comparedwith ambient temperature controls. Elevated temperatures resultedin less deep frost resistance. Apical buds generally developeddeeper frost hardiness than lateral buds but less deep thanthe cambium, varying with species, however. Frost damage inbuds ranged from death over partially destroyed bud contentsresulting in distorted shoots to buds seemingly remaining dormant.Responses to warm spells differed with duration, timing andspecies, ranging from dramatic decrease in frost hardiness withor without subsequent recovery to no reaction. Furthermore,the reactions did not show any clear relation to dormancy level.For A. procera, exposure to fluctuating temperatures appearedto be particularly problematic. This explains why this speciesdevelops best in coastal climates, and in sites sheltered fromtemperature extremes either by hedging, a winter snow cover,or topography. The Christmas tree production will suffer severelyon sites with harsh temperatures due to losses of lateral andterminal buds, which destroy the crown symmetry. Clipping ofgreenery is less influenced by frost damages, although the developmentof normal branch whorls is often disturbed.     

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.     

Changes in freezing tolerance, plasma membrane H+-ATPase activity and fatty acid composition in Pinus resinosa needles during cold acclimation and de-acclimation

It has previously been suggested that plasma membrane ATPase (PM H+-ATPase, EC 3.6.1.3.) is a site of incipient freezing injury because activity increases following cold acclimation and there are published data indicating that activity of PM H+-ATPase is modulated by changes in lipids associated with the enzyme. To test and extend these findings in a tree species, we analyzed PM H+-ATPase activity and the fatty acid (FA) composition of glycerolipids in purified plasma membranes (PMs) prepared by the two-phase partition method from current-year needles of adult red pine (Pinus resinosa Ait.) trees. Freezing tolerance of the needles decreased from -56 degrees C in March to -9 degrees C in May, and increased from -15 degrees C in September to -148 degrees C in January. Specific activity of vanadate-sensitive PM H+-ATPase increased more than two-fold following cold acclimation, despite a concurrent increase in protein concentration. During de-acclimation, decreases in PM H+-ATPase activity and freezing tolerance were accompanied by decreases in the proportions of oleic (18:1) and linoleic (18:2) acids and increases in the proportions of palmitic (16:0) and linolenic (18:3) acids in total glycerolipids extracted from the plasma membrane fraction. This pattern of changes in PM H+-ATPase activity and the 18:1, 18:2 and 18:3 fatty acids was reversed during cold acclimation. In the PM fractions, changes in FA unsaturation, expressed as the double bond index (1 x 18:1 + 2 x 18:2 + 3 x 18:3), were closely correlated with changes in H+-ATPase specific activity (r2 = 0.995). Changes in freezing tolerance were well correlated with DBI (r2 = 0.877) and ATPase specific activity (r2 = 0.833) in the PM fraction. Total ATPase activity in microsomal fractions also closely followed changes in freezing tolerance (r2 = 0.969). We conclude that, as in herbaceous plants, simultaneous seasonal changes in PM H+-ATPase activity and fatty acid composition occur during cold acclimation and de-acclimation in an extremely winter hardy tree species under natural conditions, lending support to the hypothesis that FA-regulated PM H+-ATPase activity is involved in the cellular response underlying cold acclimation and de-acclimation.