Modelling and Measuring Transpiration from Scots Pine with Increased Temperature and Carbon Dioxide Enrichment

Starting in 1996, individual trees of Scots pine (Pinus sylvestrisL.) aged 30 years, were grown in closed-top chambers and exposedto either normal ambient conditions (CON), elevated CO₂(approx.700 µmol mol^-1; Elev. C), elevated temperature (approx.2 °C and approx. 6 °C above the outside ambient temperatureduring the ‘growing season’ and ‘off season’,respectively; Elev. T) or a combination of elevated CO₂and warmertemperature (Elev. CT). Sap flow was monitored simultaneouslyby the constant-power heat balance method in a total of 16 trees,four for each treatment, over a 32 d period in summer 1998 (afterthe completion of needle expansion and branch elongation). Toquantify the contributions of crown and physical environmentalvariables to total crown transpiration, a ‘sun/shade model’was developed and used to partition the changes in transpirationto different sources. The results of the sap flow measurementsindicate that (1) total daily sap flow (E_tree.d) varied from0.15–3.41 kg per tree; (2) the treatment effect on E_tree.ddependedgreatly on the weather conditions; (3) the cumulative E_tree.dforthe 32 d dropped significantly by 22% relative to CON (P =0.038)under Elev. C and increased significantly by 21% (P =0.043)and 16% (P =0.048) under Elev. T and Elev. CT, respectively.In general, the modelled transpiration gave good agreement withthe sap flow results. The model computations showed that, ona typical sunny day in summer, the effect of treatment on crownstomatal conductance was responsible for approx. 80% of thechange inE_tree.d , while the increase in needle area and theeffect on total radiation absorption contributed only a smallpercentage. Furthermore, sunlit needles were responsible forover 60% of change in transpiration. The effect of the treatmentson E_tree.dwas larger at high temperature and vapour pressuredeficit but was not sensitive to incident daily radiation. Copyright2000 Annals of Botany Company Transpiration model, sap flow, CO₂and temperature elevation, environment-controlled chamber, Pinus sylvestris L.     

Changing Environmental Effects on Frost Hardiness of Scots Pine During Dehardening

The effects of raised temperature and extended photoperiod onthe dehardening of quiescent and winter-hardy Scots pine saplingswere examined in an open-top-chamber experiment. The saplingswere exposed during winter to natural, square-curve fluctuating(between 1 and 11 °C with a 14 d interval), and constant(6 °C) temperatures with a natural and an extended (17 h)photoperiod. Frost hardiness of needles was determined by controlledfreezing tests and visual damage scoring. The constant 6 °Ctemperature treatment caused a gradual dehardening of needleswhereas under fluctuating temperatures the level of frost hardinessfluctuated. Trees exposed to extended photoperiods were lesshardy than under natural photoperiods after the initiation ofshoot elongation, but before this there were no clear differencesin frost hardiness between different photoperiodic treatments.The results indicate that the frost hardening competence ofScots pine changes during quiescence. Climate change; frost hardiness; hardening competence; photoperiod; Pinus sylvestris, Scots pine; temperature     

Movement of Nutrients in Radiata Pine Needles in Relation to the Growth of Shoots

The pattern of needle growth and the movement of mineral nutrients(nitrogen, phosphorus, potassium and calcium) in needles ofradiata pine (Pinus radiata D. Don) were studied from needleinitiation to 2 years of age. During this period, very prominentcyclic patterns of nutrient accumulation, retranslocation andreplenishment were observed for nitrogen, phosphorus and potassium,which highlighted the potential role of needles as a nutrientreserve for growth. Significant retranslocation occurred from very young needlesabout 3 months after initiation. The phases of retranslocationcoincided with new flushes of shoot growth, and the growth ofnew shoots on a branch resulted in nutrient withdrawal frompre-existing needles, regardless of needle age and season. Suchwithdrawal occurred even in fertilized and irrigated trees onhigh quality sites and under environmental conditions conducivefor nutrient uptake. At all times, except for the short periodafter initiation when needles were actively growing, the nutrientsin the needle were readily available for retranslocation. Contraryto the general view, retranslocation of nutrients was not necessarilyrelated to senescence and ageing of needles. Because new shoots are the primary ‘sinks’ for retranslocatednutrients, an ongoing competition between different parts ofa branch for internal nutrients can be envisaged, preferencebeing for the youngest shoot in the hierarchy. The relevanceof these results to our understanding of ‘optimum nutrition’of pine trees is discussed. Pinus radiata D. Don, radiata pine, mineral nutrition, retranslocation, phosphorus, nitrogen, shoot growth     

Developmental control of xylem hydraulic resistances and vulnerability to embolism in Fraxinus excelsior L.: impacts on water relations

The freezing tolerance of many plants, such as pea (Pisum sativum),is increased by exposure to low temperature or abscisic acidtreatment, although the physiological basis of this phenomenonis poorly understood. The freezing tolerance of pea shoot tips,root tips, and epicotyl tissue was tested after cold acclimationat 2C, dehydration/rehydration, applications of 10^–4M abscisic acid (ABA), and deacclimation at 25C. Tests wereconducted using the cultivar ‘Alaska’, an ABA-deficientmutant ‘wil’, and its ‘wildtype’. Freezinginjury was determined graphically as the temperature that caused50% injury (T₅₀) from electrical conductivity. Endogenous ABAwas measured using an indirect enzyme-linked immunosorbant assay,and novel proteins were detected using 2-dimensional polyacrylamidegel electrophoresis. The maximum decrease in T₅₀ for root tissuewas 1C for all genotypes, regardless of treatment. For ‘Alaska’shoot tips and epicotyl tissue, exogenous ABA increased thefreezing tolerance by –1.5 to –4.0C, while coldtreatment increased the freezing tolerance by –7.5 to–14.8C. Cold treatment increased the freezing toleranceof shoot tips by –9 and –15C for ‘wil’and ‘wild-type’, respectively. Cold acclimationincreased endogenous ABA concentrations in ‘Alaska’shoot tips and epicotyls 3- to 4-fold. Immunogold labeling increasednoticeably in the nucleus and cytoplasm of the epicotyl after7 d at 2C and was greatest after 30 d at the time of maximumfreezing tolerance and soluble ABA concentration. Cold treatmentinduced the production of seven, three, and two proteins inshoot, epicotyl, and root tissue of ‘Alaska’, respectively.In ‘Alaska’ shoot tissue, five out of seven novelproteins accumulated in response to both ABA and cold treatment.However, only a 24 kDa protein was produced in ‘wil’and ‘wild-type’ shoot and epicotyl tissues aftercold treatment. Abscisic acid and cold treatment additivelyincreased the freezing tolerance of pea epicotyl and shoot tissuesthrough apparently independent mechanisms that both resultedin the production of a 24 kDa protein. Key words: Pisum sativum, cold acclimation, immuno-localization     

Freezing tolerance, protein composition, and abscisic acid localization and content of pea epicotyl, shoot, and root tissue 3

The freezing tolerance of many plants, such as pea (Pisum sativum),is increased by exposure to low temperature or abscisic acidtreatment, although the physiological basis of this phenomenonis poorly understood. The freezing tolerance of pea shoot tips,root tips, and epicotyl tissue was tested after cold acclimationat 2C, dehydration/rehydration, applications of 10^–4M abscisic acid (ABA), and deacclimation at 25C. Tests wereconducted using the cultivar ‘Alaska’, an ABA-deficientmutant ‘wil’, and its ‘wildtype’. Freezinginjury was determined graphically as the temperature that caused50% injury (T₅₀) from electrical conductivity. Endogenous ABAwas measured using an indirect enzyme-linked immunosorbant assay,and novel proteins were detected using 2-dimensional polyacrylamidegel electrophoresis. The maximum decrease in T₅₀ for root tissuewas 1C for all genotypes, regardless of treatment. For ‘Alaska’shoot tips and epicotyl tissue, exogenous ABA increased thefreezing tolerance by –1.5 to –4.0C, while coldtreatment increased the freezing tolerance by –7.5 to–14.8C. Cold treatment increased the freezing toleranceof shoot tips by –9 and –15C for ‘wil’and ‘wild-type’, respectively. Cold acclimationincreased endogenous ABA concentrations in ‘Alaska’shoot tips and epicotyls 3- to 4-fold. Immunogold labeling increasednoticeably in the nucleus and cytoplasm of the epicotyl after7 d at 2C and was greatest after 30 d at the time of maximumfreezing tolerance and soluble ABA concentration. Cold treatmentinduced the production of seven, three, and two proteins inshoot, epicotyl, and root tissue of ‘Alaska’, respectively.In ‘Alaska’ shoot tissue, five out of seven novelproteins accumulated in response to both ABA and cold treatment.However, only a 24 kDa protein was produced in ‘wil’and ‘wild-type’ shoot and epicotyl tissues aftercold treatment. Abscisic acid and cold treatment additivelyincreased the freezing tolerance of pea epicotyl and shoot tissuesthrough apparently independent mechanisms that both resultedin the production of a 24 kDa protein. Key words: Pisum sativum, cold acclimation, immuno-localization     

A Simulation Model for the Annual Frost Hardiness and Freeze Damage of Scots Pine

The changes in the frost hardiness of Scots pine were modelledby a dynamic model where the input variables were temperatureand photoperiod and the phase of annual development. The damagecaused by freezing was described by the sigmoidal relationshipbetween the relative needle damage and freezing temperature.The model simulations were carried out using temperature datafrom two sites in central Finland—Suonenjoki and Tampere.The validity of the frost hardiness model was tested with measuredfrost hardiness data from Suonenjoki. The effects of climaticwarming were also simulated by increasing temperature of thelong-term climatic data. Genotypic differences in chilling requirement,which determines the timing of the reduction of hardening competence,were included in the simulations. The simulated needle damageincreased as a result of climatic warming, and the differencesin the chilling requirement had a stronger effect on the amountof damage in the warmed climate than in the present climate.A large variation between years was found in the level of damage. Annual development; climatic change; dynamic model; freeze damage; frost hardiness,Pinus sylvestris ; Scots pine     

Optimal Control of Gas Exchange during Drought: Empirical Evidence

The optimal regulation model by Mäkelä, Berningerand Hari (Annals of Botany 77: 461–467, 1996) was appliedto data for photosynthesis and transpiration of Scots pine duringa 22-d drought period. There was a clear decrease in photosynthesisand transpiration during that period. The agreement betweenmodel and photosynthesis data was good. The residuals of photosynthesiswere not systematic with respect to temperature, irradianceor water vapour deficit. However, the model initially overestimatedtranspiration by 50%, although there was a clear linear relationshipbetween measured and estimated values. The results suggest thatthere was no decrease in photosynthetic capacity during theperiod, but a decrease in stomatal conductance was responsiblefor the changes in photosynthesis and transpiration. The observationsare similar to results in the literature. Transpiration; photosynthesis; stomatal conductance; drought; Pinus sylvestris     

Spatial Distribution of Ethylene Production by Individual Needles Along a Shoot ofPinus sylvestrisL.: Relationship with Peroxidase Activity

Spatial distribution of ethylene production by individual currentyear needles and the relationship with peroxidase activity wasstudied along a shoot of Scots pine (Pinus sylvestrisL.). Muchvariation in ethylene production between and within individualseedlings was found. There was a significant difference in meanintensity of ethylene production between orthostichies withinindividual seedlings. The capacity of individual needles toconvert 1-aminocyclopropane-1-carboxylic acid (ACC) to ethylenedid not correlate with the basal rate of ethylene production.Mean ethylene production intensity by orthostichies correlatedpositively with mean peroxidase activity. A positive correlationbetween the intensity of ethylene production by individual needlesand detachment-induced peroxidase activity was found. The datasuggests that the rate of ethylene production by individualneedles may be directly related to their peroxidase activityand at least in part to their antioxidative capacity.Copyright1998 Annals of Botany Company Pinus sylvestris,pine, ethylene production, spatial distribution, individual needles, peroxidase, seasonal changes.     

Physiology of Flowering in Saccharum: I. DAYLENGTH CONTROL OF FLORAL INITIATION AND DEVELOPMENT IN S. SPONTANEUM L.

Flowering in two clones of Saccharum spontaneum L. is controlledby photoperiod. The earliest stages of development, ‘induction’and ‘initiation of the inflorescence axis primordium’(IAP) were optimally promoted under intermediate days of 12h 30 min, while the subsequent stage ‘initiation of inflorescencebranch primordia’ (IBP) was inhibited by days longer than13 h. The following stage ‘initiation of spikelet primordia’(ISP) showed a quantitatively intermediate response with anoptimum photoperiod of 9 h to 11 h. The elongation of the differentiatedinflorescence was found to be only slightly sensitive to photoperiodsof 13 h or longer in one of the clones. Unfavourable photoperiodsat stages following induction resulted in the arrest or delayof inflorescence development and when these were given duringthe IAP and IBP stages, reversion to the vegetative conditioncommonly occurred.     

The uptake of and structural changes induced by trichloroacetic acid in the needles of Scots pine seedlings

The uptake of trichloroacetic acid (TCA) and structural changesinduced in the needles of Scots pine (Pinus sylvestris L.) seedlingswere studied. Two exposure set-ups, a root route and an atmosphericroute through the surfaces simulating the wet deposition offog, were used. Both set-ups included two dose levels and correspondingcontrol treatments. The temperature and the relative humidityin the climate chambers were adjusted to represent the conditionsof June–July in a subarctic area in central Finland. Theseedlings were exposed three times a week for two months. Theresults showed that the uptake of TCA in needles occurred bothvia roots and via needle surface. However, most of the TCA viathe atmospheric route was absorbed on the surface of the needles.The structural responses in pine needles depended partly onthe treatment method: TCA applied via the atmospheric routedisintegrated the structures of the epicuticular waxes and thatof the stomatal cells, which was not seen in the exposures viaroots. A common feature was the decrease in size of the chloroplastsin concert with the increasing TCA concentrations inside theneedles. Key words: Pinus sylvestris L., climate chamber, effects, microscopic structure, secondary pollutant, TCA, trichloroacetic acid     

Effect of exogenous gibberellins on flowering in Pinus sylvestris grafts

Ethanolic sprays of gibberellins were applied to developing shoots of about 12-year-old Scots pine (Pinus sylvestris L.) grafts during the shoot elongation period in two consecutive years. Both male and female flowering was increased by these treatments in both years. The effect was particularly distinct in male flowering. However, different clones showed varying responses to the treatment. This variation was associated with both the genotype and the environmentally determined year-to-year fluctuation in flowering. Differences among clones were analysed further by introducing a model earlier developed for comparisons of the growth rhythm in various woody and herbaceous species.     

Patterns of Root Respiration Associated with the Induction of Aluminium Tolerance in Phaseolus vulgaris L.

Aluminium (Al) tolerance in an Al-tolerant cultivar of Phaseolusvulgaris L. (‘Dade’) was found to be an inducibletrait. Upon exposure to 10 µM Al, the rate of root elongationwas inhibited in comparison to controls. During the following72 h, the rate of elongation returned to levels comparable tocontrols. In contrast, root elongation of an Al-sensitive cultivar(‘Romano’) did not recover after exposure to Al.In Dade, the resumption of root elongation following exposureto Al was accompanied by increased rates of root respiration,whereas respiration rates slowly declined over the 72 h treatmentperiod in Romano. When partitioned into growth and maintenanceexpenditures, a larger proportion of root respiration of Dadeexposed to Al was allocated to maintenance processes, potentiallyreflecting diversion of energy to metabolic pathways that offsetthe adverse effects of Al toxicity. Romano did not show sucha pattern and respiration associated with both growth and maintenancewas reduced after exposure to Al. Root and shoot growth of bothcultivars were also measured to determine the effects of long-term(21 d) exposure to 10 µM Al. Dade plants exposed to Alexhibited enhanced growth in comparison to controls, whereasRomano plants were characterized by reduced shoot and root growth.Modelling the time-course of root respiration and measuringthe long-term growth responses to Al is a valuable method ofelucidating respiratory costs of stress tolerance. Key words: Aluminium, differential tolerance, maintenance respiration, Phaseolus vulgaris, root respiration     

The Effect of Applications of a Synthetic Cytokinin on Shoot Dominance and Grain Yield in Spring Barley

Spring barley cv Ark Royal was grown in pots in the open, andsoil drenches containing 6-benzyl-aminopurine were repeatedlyapplied during the pre-heading phase starting either ‘early’when the main shoot apex had reached the glume primordium stage,during tillering, or ‘later’ at the anther primordiumstage, at the start of stem elongation Although grain yield was unaffected by the ‘late’cytokinin treatments, it was increased by up to 57 per centby the ‘early’ treatments. The increase was dueto greater grain yields of the smaller shoots of each plantthere was no effect on the yields of the two largest shootsor on the total number of fertile shoots The increased grain yield of these smaller shoots was attributableto increased weights of their individual kernels, especiallyin the basal and distal parts of the ears such that a greateruniformity of kernel size within the ear was achieved Thus the cytokinin treatment increased uniformity both betweenshoots and between spikelets within the ears Hordeum vulgare L, spring barley, cytokinin, 6-benzylaminopunne, shoot dominance, harvest index, grain yield     

Relative Sensitivity and Tolerance of some Gladiolus Cultivars to Sulphur Dioxide

Five Gladiolus cultivars, namely ‘Aldebaran’, ‘BrightEye’, ‘Illusion’, ‘Manisha’ and‘Manmohan’, were exposed to 1 and 2 µg l^–1sulphur dioxide to test their relative-sensitivity toleranceto the pollutant Plants were fumigated experimentally for 2h daily Foliar injury symptoms were observed first in ‘Manisha’followed by ‘Aldebaran’ and ‘Illusion’at the higher dose Photosynthetic pigments and leaf extractpH were significantly decreased, particularly in ‘Manisha’and ‘Illusion’ Overall disturbances in the plantmetabolism due to SO₂ treatment led to retarded growth of plants,as evident from decreased shoot length and phytomass valuesThe order of sensitivity of the five Gladiolus cultivars toSO₂ was as follows, with the greatest first Manisha, Illusion,Aldebaran, Bright Eye, Manmohan Cultivars, Gladiolus, sensitivity, sulphur dioxide, tolerance     

Effects of Gibberellin, Clone and Environment on Cone Initiation, Shoot Growth and Branching in Pinus contorta

Effects of photoperiod, temperature and gibberellin applicationwere clearly separated from clonal differences when small rootedcuttings, originating from two ‘floriferous’ treesof the same provenance of Pinus contorta Dougl., were treatedwith a simple micro-injection technique and grown in growthcabinets. Short days (10 h) for 11 weeks induced six times asmany female cones as long days (19.5 h), completely inhibitedfurther needle elongation after 5 weeks, and also led to enhancedbranching by year 3. Cool temperatures appeared to favour malecone initiation in year 1, and (in conjunction with long days)significantly promoted height in year 2. Injection with 1 mgGA_4/7 led to substantial and highly significant increases inbud growth in year 1, and in height, needle number and totalneedle length in year 2. Gibberellin treatment also increasedbranching but did not significantly affect internode elongationin year 2, nor cone initiation or needle elongation in year1. Highly significant clonal differences occurred, with onegenotype having longer leaves and buds in year 1, and up to80 per cent greater numbers and total lengths of needles inyear 2. Marked clonal differences in height growth, branchingand sex ratio were also found. The implications of both resultsand approach are discussed in relation to developmental physiology,genetic selection and advances in forestry research. leaf growth, height growth, branching, cone initiation, sex, photoperiod, temperature, controlled environments, gibberellin, vegetative propagation, clonal differences, Pinus contorta,, pine     

Ageing in Woody Plants

The shoot systems of trees undergo characteristic changes withtime, as they increase in size and complexity. Certain of thesechanges, such as reduced annual growth increment, loss of apicaldominance, and changes in the geotropic responses, are easilyreversible, by pruning or by grafting shoots from older treeson to young stocks. These changes are referred to as ‘ageing’. This paper describes the changes occurring in Pinus sylvestrisand Larix leptolepis during ageing and reports an investigationinto the possible mechanisms controlling these changes. Pruningexperiments suggest that ageing is due to increased competitionfor nutrients between the various constituent shoots as thebranch systems increase in complexity, but the distributionof available nutrients is modified by apical dominance. Theseconclusions were supported by the results of experiments involvingthe injection of radioactive phosphorus (²²P), in associationwith the pruning treatments.     

Primary and Secondary Induction Requirements for Flowering of Contrasting European Varieties of Lolium perenne

The flowering requirements of six European varieties of Loliumperenne L. were studied in controlled environments. In experimentson primary induction, flowering was recorded after transferto long days (LD) in a greenhouse at 12–24°C. In experimentson secondary induction, primary induction was first accomplishedat 6°C/10 h daylength for 12 weeks. When evaluated by the50% heading criterion, the requirement for duration of primaryinduction at 6°C/8 h daylength was <3 weeks in Mediterranean,5–6 weeks in Central European and 7–8 weeks in Scandinavianvarieties. While ‘Veyo’ (Italy) flowered profuselyregardless of temperature or daylength during primary induction,critical temperatures for primary induction in SD and LD were15 and 11°C in ‘Baca’ (Czech Republic) and 11and 7°C in ‘Falster’ (Denmark). The criticalphotoperiod for secondary induction at 15°C ranged from12 h in ‘Veyo’ and 14 h in ‘Baca’ to16.5 h in ‘Falster’ and 17.5 in ‘Kleppe’(Norway). The critical number of LD cycles varied correspondingly.While the Central and North European varieties required fewerLD cycles for 50% heading at 18 than at 12°C, ‘Veyo’showed the opposite response. It is concluded that the requirementsfor both primary and secondary induction of Lolium perenne increasewith increasing latitude of origin of the germplasm. In oneexperiment, 39–87% of the inflorescences came from tillersthat were not visible on transfer from primary to secondaryinduction, thus it is also concluded that there is no juvenilestage in tillers of Lolium perenne. Copyright 2000 Annals ofBotany Company Daylength, flowering, juvenility, perennial ryegrass (Lolium perenne L.), primary induction, secondary induction, temperature, varieties, vernalization     

Seasonal Changes in the Physiology of S24 Perennial Ryegrass (Lolium perenne L.): 3. Partition of Assimilates between Root and Shoot during the Transition from Vegetative to Reproductive Growth

The partition of ¹⁴C labelled current assimilates to root insimulated swards of Lolium perenne cv. S24 was measured duringthe transition from vegetative growth in autumn to reproductivegrowth in spring under close to natural conditions of lightand temperature. Assimilate partitioning was also measured in‘established’ swards cut three times during thegrowing season and in vegetative ‘seedling’ swardsgrowing in autumn and in spring. All measurements were madewhen the swards had achieved more than 90 per cent light interception,and all swards were abundantly supplied with water and mineralnutrients. During autumn there was a gradual decrease in the proportionof assimilates partitioned to the roots in both the ‘established’and the ‘seedling’ swards. In the established swards,partition to roots was low over winter, increased during earlyspring, but decreased dramatically, later in the spring, whenstem elongation began. In contrast, in the unvernalized vegatativeseedling swards in spring, partition to roots remained high. The seasonal pattern of assimilate partitioning is consideredin relation to changes in the natural environment and the rateat which the crop fixed carbon in photosynthesis. A decreasein the proportion of assimilates partitioned to roots duringlate spring was significant in increasing the production ofshoot at that time but seasonal differences in partition contributedvery little to the marked differences in shoot growth betweenthe spring and autumn crop. Lolium perenne L., perennial ryegrass, partition of assimilates, flowering     

An Agar-diffusion Technique Applied to the Study of Gibberellins in Tomato (Lycopersicon esculentumMill.) Plants

Chromatography of extracts of agar blocks on which shoot tipsof tomato had stood for 24 h revealed the presence of at leasttwo gibberellin-like substances. Gibberellin-like activity detectedby the dwarf pea bioassay was shown to be associated with activityin promoting elongation growth in tomato seedlings. The activesubstances were acidic; tests for the presence of ‘neutral’,‘basic’ and ‘bound’ gibberellins inextracts of the agar blocks gave negative results. Amounts ofgibberellin-like substances obtained from the shoot tips wereincreased by the incorporation of EDTA into the agar blocksbut were apparently unaffected by the temperature (within therange 5–25 °C) at which the diffusion was carriedout. It is suggested that the technique measured levels of diffusiblegibberellins in the tissues at the time of their excision ratherthan rates of gibberellin synthesis during the diffusion period.     

Plant resistance to cold stress: Mechanisms and environmental signals triggering frost hardening and dehardening

This introductory overview shows that cold, in particular frost, stresses a plant in manifold ways and that the plant’s response, being injurious or adaptive, must be considered a syndrome rather than a single reaction. In the course of the year perennial plants of the temperate climate zones undergo frost hardening in autumn and dehardening in spring. Using Scots pine (Pinus sylvestris L.) as a model plant the environmental signals inducing frost hardening and dehardening, respectively, were investigated. Over 2 years the changes in frost resistance of Scots pine needles were recorded together with the annual courses of day-length and ambient temperature. Both act as environmental signals for frost hardening and dehardening. Climate chamber experiments showed that short day-length as a signal triggering frost hardening could be replaced by irradiation with far red light, while red light inhibited hardening. The involvement of phytochrome as a signal receptor could be corroborated by respective night-break experiments. More rapid frost hardening than by short day or far red treatment was achieved by applying a short period (6 h) of mild frost which did not exceed the plant’s cold resistance. Both types of signals were independently effective but the rates of frost hardening were not additive. The maximal rate of hardening was − 0.93°C per day and frost tolerance of < − 72°C was achieved. For dehardening, temperature was an even more effective signal than day-length.