Studies on the Dormancy of Calluna vulgaris (L.) Hull, During Winter: Classification of Dormancy

Experiments are described in which plants of Calluna vulgariswere transferred from a high altitudinal blanket bog to controlledenvironment rooms under a range of conditions, to determinethe type of dormancy mechanism which occurs in this speciesduring the winter months. It was found that in autumn, Callunaexhibits semi-dormancy, bud break not occurring under shortphotoperiods. From December onwards, bud break will occur under short photoperiodsif warm temperatures are applied; at this time the plants arein a state of post-dormancy. Calluna lacks a period of truedormancy since it can always be forced to resume growth by longphotoperiods. Although in autumn, plants will not break dormancy under shortdays, after experiencing the natural outdoor conditions of winter,they will break dormancy equally readily under long and shortdays when brought into warm controlled environment rooms inthe spring. Calluna vulgaris (L.) Hull., heather, dormancy, pre-dormancy, post-dormancy, semi-dormancy     

Influence of Nitrogen and Winter Climate Stresses on Calluna vulgaris(L.) Hull

Four nitrogen (N) levels, spanning those found naturally, wereestablished in Calluna vulgaris grown in containers in an oceanicclimate for 4 years on the southwest coast of Norway. EstablishedN contents of the plants were 0.7, 0.8, 1.3 and 2.5% of dryweight. Experiments comprising 5000 plants were performed inthe open and in growth chambers. In the open, dry matter accumulationand date of bud burst were recorded at the different N levels.Breaking of winter dormancy was studied under controlled conditions.In October/November, plants needed 1474 day-degrees for budburst at the lowest N level, whereas at the highest level only582 day-degrees were required. The natural start of growth inthe field was on 14 April at the highest N level and on 26 Aprilat the lowest. Freeze testing of the shoots (with the rootsprotected from frost) revealed that Calluna tolerates temperaturesas low as -15 °C as late as the beginning of April. However,plants at the two medium N levels were most tolerant of frost.During the last winter of the experiments, nearly 50% of plantsin the open died at the highest N level, whereas plants at thelower levels survived without damage. The effects of N are discussedin relation to the influence on physiological ageing of theplants, the change in nutrient balance of the plant tissue andthe subsequent increase in susceptibility to biotic and abioticstresses. Copyright 2001 Annals of Botany Company Calluna vulgaris, climate stress, growth initiation, nitrogen     

A reappraisal of cyclical processes in Calluna heath

Small-scale, short-term cycles constitute an important aspect of vegetation dynamics. It has been widely accepted that such cycles are characteristic of unmanaged heath communities where the dominant, Calluna vulgaris, undergoes an age-related series of changes in its growth and morphology. This is thought to give rise to a repetitive sequence of changes in the occupancy of gaps which form in the Calluna canopy and are first colonized by other species, with Calluna re-establishing later. Recently, doubts have been cast as to whether the changes are in fact often repetitive. A new study has shown much greater variability than was previously suspected in the transitions occurring in gaps. There is evidence that, in some instances though by no means all, Calluna can re-establish and thereby initiate a genuine cycle. It seems, however, that such re-entry is seldom by means of seedling establishment, but more frequently by development of adventitious shoots and roots on stems which have come to lie across the gap and have been covered by moss, moist litter or humus.     

The Effects of Temperature and Photoperiod on the Onset of Summer Dormancy in Poa bulbosa L.

Poa bulbosa L. plants became dormant in long days (16 h), whilein short days (8 h) they remained non-dormant for extended periods.Morphologically, the onset of dormancy was expressed by theformation of a true bulb at the base of every tiller, by thecessation of tillering and leaf emergence and, finally, by thedrying-up of the leaves. Low temperature delayed the onset ofdormancy but did not prevent it. This effect of temperaturemay explain the delayed onset of dormancy observed in naturalpopulations under a cool climate at a hilly habitat, comparedto plants growing under a warmer climate, at a lower, coastal-plainhabitat. Dormancy could be induced under short days by pre-exposureof the plants to a limited number of long days. The responsewas proportional to the number of long days given. The adaptivesignificance of the results for plant survival in a Mediterraneanclimate is discussed. Poa bulbosa L., summer dormancy, photoperiod, temperature, leaf emergence, bulbs, tillers     

Effects of browsing and grazing on cyclic succession in nutrient‐limited ecosystems

Abstract. This paper deals with browsing and grazing as forces driving cyclic succession. Between 1989 and 1994 reciprocal transitions between the dwarf shrub Calluna vulgaris and the grass Deschampsia flexuosa were monitored in permanent plots in a cattle grazed grass‐rich Dutch heathland on podsolic soils in which tree encroachment was prevented. Heather beetles killed Calluna in four of the nine plots during 1991/1992. The monitoring revealed reciprocal transitions and cycles between Calluna and Deschampsia on a subplot scale. Beetles and cattle had additional and complementary effects on the two competing species. Defoliation by beetles and trampling by cattle‐killed Calluna and favoured grass invasion. Grazing and gap creation by cattle in Deschampsia favoured the establishment and recovery of Calluna. Analysis of the causal mechanisms suggests that indirect, resource‐mediated herbivory effects may be as important for the replacement processes as direct effects of defoliation and trampling. Herbivory created differential light and nutrient levels in Calluna and Deschampsia gaps. Grazing and browsing improved the resource‐capturing abilities of Calluna and its resistance to herbivory and abiotic disturbances. The emerged Calluna‐Deschampsia cycle and its driving forces are summarized in a conceptual triangular resource‐mediated successional grazing cycle (RSGC) model, a limit cycle involving herbivore‐plant‐plant resource interactions. It offers a deterministic equilibrium model as alternative for stochastic transitions between the meta‐stable states with dominance of Calluna and Deschampsia respectively. The validity range of the RSGC model and its management implications are briefly discussed.     

The Mechanism of Low Temperature Dormancy in Mature Seeds of Syringa Species

The mechanism of seed dormancy at low temperatures (15-9°C) was investigated in the seeds of Syringa josikaea, S. reflexa and S. vulgaris. Low temperature dormancy in Syringa species was mainly imposed by endosperm embedding the radicle. Different degrees of embryo dormancy may occur in S. reflexa seeds. In most cases the low temperature dormancy was broken completely by removing the endosperm around the radicle. The endosperm did not seem to contain significant quantities of germination inhibitors, and the results indicate that it prevents germination mainly due to its mechanical resistance. The mechanical resistance of endosperm did not change during chilling or during induction of dormancy by high temperature incubation. The strength of the endosperm decreased rapidly in non-dormant seeds before visible germination. Similar changes were not observed in dormant seeds. Generally, the strength of the endosperm was lower in the non- (or less) dormant species S. josikaea and S. vulgaris than in the more dormant S. reflexa seeds. The growth potential of the embryos, measured as their ability to germinate in osmotic solutions (mannitol or polyethylen glycol 4000), was increased by chilling and by GA₃-treatment. The growth potential of untreated S. josikaea and S. vulgaris embryos was generally higher than that of S. reflexa embryos. Acid ethyl acetate fractions of methanol extracts from embryos of all three species contained substances with GA³-like activity in the lettuce hypocotyl test. The activity was found at Rf 0.9–1.0 on paper chromatograms run in distilled water. No significant changes in the activity were detected during chilling or prior to visible germination.     

Effects of fertilization and cutting on the chemical composition of vegetation and soils of mountain heathlands in Spain

Abstract. In the mountains of northern Spain, patches dominated by Calluna vulgaris are scarce and they may disappear or change as a result of continued lack of management and possibly increasing nutrient availability through atmospheric deposition. The effects in the soil properties and in the composition of Calluna vulgaris and Erica tetralix shoots on heathlands dominated by Calluna and Erica subjected to fertilization and experimental cutting were studied in three mountain passes in northern Spain. A total of 90 1‐m² plots received different combinations of cutting and twice the estimated atmospheric deposition of nitrogen (5.6 g.m^?2.yr^?1) as ammonium nitrate. One of the dominant ericaceous species (Calluna and Erica) was selectively cut by hand at ground level and their nitrogen shoot content were compared in the presence or absence of the other. Treatments were carried out in April 1998. In each plot one soil sample was taken in the original situation and 12, 24 and 36 months after the treatments. Soil properties such as organic matter, total nitrogen, available phosphorus and pH were determined. In every plot five shoots of Calluna and Erica were also taken to analyse total nitrogen content in the original situation and 12, 24 and 36 mo after the treatments. Nitrogen addition does not necessarily lead to increased levels in the soil, and a clear pattern was not found in the three areas. A gradual decrease in available phosphorus content was detected in the three areas until two years after treatment, although values tend to recover in two of the areas in the third study year. An increase in organic matter content was observed in all areas. It is concluded that increased nutrients alone, at twice the rate of the estimated current atmospheric deposition for the area, which is relatively low, will not alter significantly the soil characteristics of the mountain heathland stands. A clear increase in plant N‐content is observed in the fertilized plots in comparison with the non‐fertilized ones and Calluna always has higher nitrogen content than Erica. This increase is most pronounced one year after the treatments started in one of the areas and after two years in the other two areas. In some cases the elimination of one species is seen to favour nitrogen increase in the other.     

Effects of litters with different concentrations of phenolics on the competition between Calluna vulgaris and Deschampsia flexuosa

We hypothesized that the outcome of competition between ericaceous plants and grasses is strongly affected by the concentrations of phenolics in the litter that they produce. To test the effect of phenolic-rich litter on soluble soil nitrogen concentrations, plant nitrogen uptake and inter-specific competition, we conducted a greenhouse experiment with the shrub Calluna vulgaris and the grass Deschampsia flexuosa and their leaf litters. Two litters of C. vulgaris were used, with equal nitrogen concentration but different (high and low) concentrations of total phenolics. The D. flexuosa leaf litter contained lower concentrations of phenolics, but higher concentrations of nitrogen than the C. vulgaris litters. The plants were grown in monocultures and in mixed cultures. Inorganic and dissolved organic nitrogen were measured monthly during the experiment. After four months, we measured above- and belowground biomass and the nutrient concentrations in above- and belowground plant parts. In monocultures, C. vulgaris produced more shoot and root biomass on its own litter than with no litter. Growth of Calluna was reduced on grass litter. D. flexuosa plants produced most biomass on their own litter type, whether in monocultures or in mixed cultures. Addition of Calluna litter stimulated the growth of D. flexuosa both in monoculture and in mixtures. The grass plants outcompeted Calluna both on shrub litter and on grass litter but not when grown without litter. The two C. vulgaris litter types that differed in their concentration of phenolics did not differ in their effects on the competition between the two species or on the production of inorganic and dissolved organic nitrogen. We conclude that the nitrogen content of the litter is more important as a plant feature driving competition between shrubs and grasses than the concentrations of phenolics.     

Long-day Induced Dormancy in Anemone coronaria L

In Anemone coronaria L. summer dormancy in Israel can be inducedby long photoperiods even in the middle of the winter. Cultivationin Europe has reduced but not eliminated this response to daylength.Daylength did not affect the time of flowering but dormancycaused early cessation of flowering under long days. Anothereffect of long-day treatment was to produce longer scapes. Anemone coronaria, summer dormancy, long-day induction, daylength response     

The Influence of Photoperiod on the Timing of Reproductive Cycles in Gammarus species (Crustacea, Amphipoda)

SYNOPSIS. Gammarus species time the release of their young byusing ovarian diapause and the duration of embryonic development.During diapause vitellogenesis is arrested and oostegite setaeare lost. The onset of diapause is influenced by photoperiodunder experimental conditions. Two Gammarus species were studied.Gammarus lawrencianus is typical of the majority of Gammarusspecies that produce sequential broods through the summer andhave a diapause in the autumn. Gammarus setosus is typical ofthose which have an obligatory diapause lasting most of theyear that follows the annual autumn brood. Experimental shortday photoperiods (4L to 12L) induce vitellogenesis in Gammarussetosus, and have the opposite effect on Gammarus lawrencianus.Long day photoperiods promote vitellogenesis in Gammarus lawrencianusand prolong the duration of diapause in Gammarus setosus. Itis concluded that these two Gammarus species have their ownendogenous rhythms and that the role of photoperiod is to timethe reproductive cycle. Short days near the time of the fallequinox ensure that this timing is synchronized over a rangeof latitudes. The effect of photoperiod is modulated by theinfluence of temperature on embryonic development over the broadrange of geographic latitude inhabited by the species. Shortphotoperiods inhibit but do not prevent sexual maturation ofthe young. Short days also result in a slower average growthrate and greater size at maturity.     

Changes in Dormancy and Sensitivity to Vernalization in Axillary Buds of Satsuma Mandarin Examined in vitro During the Annual Cycle

Effect of season and the presence of fruit on bud-endodormancyand the flowering response to low temperature treatments weredetermined using bud cultures of Owari satsuma mandarin (Citrusunshiu Marc). Bud dormancy was deeper in fruiting as comparedto defruited trees. In fruiting trees, the intensity of buddormancy was highest in spring, decreased to a low value byearly Jul. and then increased until early winter. This increasein dormancy during summer and early autumn did not occur innon-fruiting trees. No flowers formed in buds cultured betweenMay and Sep. Both in fruiting and defruited trees, buds becamecompetent to show a vernalization response to chilling by theend of Oct., at the time they also became capable of sproutingin vitro at low temperature (15/10 °C). There was a directeffect of fruit on the buds which persisted long after fruitremoval and resulted in a reduction of the flowering responseto chilling.Copyright 1995, 1999 Academic Press Citrus flowering, Citrus unshiu Marc., dormancy, flower induction, flowering, in vitro flowering, satsuma mandarin, vernalization     

An Examination of some Symbiotic Systems for Fixation of Nitrogen

By the use of ¹⁵N the occurrence of fixation of elemental nitrogenhas been demonstrated in the lichens Collema granosum and Leptogiumlichenoides (both of which contain Nostoc as a symbiont), andalso in the liverwort Blasia pusilla (which has Nostoc-containingcavities in the thallus). The circumstances indicate that thefixation in these instances should be attributed to the Nostoc. No evidence of fixation was obtained in similar isotopic testson the mycorrhizal roots of intact plants of Calluna vulgarisand of Pinus sylvestris, although these, more especially inthe case of Calluna, have been held by some previous authorsto be nitrogen-fixing.     

Are Seasonal Dormancy Patterns in Arabidopsis thaliana Regulated by Changes in Seed Sensitivity to Light, Nitrate and Gibberellin?

Imbibed seeds of Arabidopsis thaliana (L.) Heynh., passed annuallythrough a pattern of changes in dormancy. Dormancy was brokenin summer and re-induced in autumn-winter. A second small germinationflush occurred in early spring. The role of sensitivity to light,nitrate and gibberellins (GAs) in regulating annual dormancypatterns and germination was studied with the use of GA-deficient(gal-2) and wild-type seeds. Dark-incubated seeds were exposedto a natural temperature regime for periods up to 18 monthsand at regular intervals germination capacity of portions ofseeds was tested at laboratory conditions. Germination datafitted as logistic dose response curves showed that sensitivityto light varied with the seasons in both genotypes. From interpretationof curve parameters, it is proposed that the observed sensitivitychanges involve alterations in the number of receptors, in thebinding characteristics of the receptors and/or in the responsechain initiated by ligand-receptor interaction. In this responsechain GA biosynthesis is stimulated (wild type) and sensitivityto GAs is enhanced (wild type, gal -2). GA sensitivity is alsodirectly influenced by temperature, thus without the interferenceof light. However, the significance of direct regulation ofGA requirement seemed to diminish with prolonged incubationoutdoors, whereas reversible changes in light sensitivity remainedclear. Therefore, we propose that seasonal dormancy patternsare mainly regulated by changes in sensitivity to light. GAsensitivity contributes to this pattern but is not primarilycontrolling dormancy. The GA requirement for germination isobvious as gal-2 seeds did not germinate at any time of theyear when deprived of applied GAs. However, GA biosynthesisis not required for dormancy control, as a dormancy patternwas also observed in the absence of the capacity to synthesizeGAs. Nitrate or sensitivity to nitrate did not contribute tothe regulation of dormancy and germination of this species.Copyright1994, 1999 Academic Press Arabidopsis thaliana (L.) Heynh., curve fitting, dormancy, fluence response curve, germination, gibberellin, gibberellin dose response curve, hormone mutant, light, mouse-ear-cress, nitrate, phytochrome, receptor, seasonal dormancy pattern, sensitivity     

Competition for nutrients between Calluna vulgaris (L.) Hull and Molinia caerulea (L.) Moench

Summary A combination of a removal and fertilization experiment in the field and a replacement experiment in containers in a cold frame was carried out to investigate the role of nutrient supply in the interactions between Calluna vulgaris and Molinia caerulea. It is concluded that the growth of Calluna as well as that of Molinia increased with increasing nutrient availability. However, the increase in biomass of Molinia was much larger than that of Calluna. It is also concluded that increased nutrient availability favours the competitive vigour of Molinia in interaction with Calluna.     

Vegetation re‐establishment on land previously subject to control of Pteridium aquilinum by herbicide

Abstract. Vegetation development after conventional bracken control with herbicide in the uplands is often slow and rarely results in the development of the target community. Two experiments were established on contrasting sites to investigate cost‐effective methods for the re‐establishment of vegetation dominated by Calluna vulgaris (heather), which is the usual target community seen as the desired replacement for Pteridium aquilinum in the uplands by many land managers. At one site, where grazing intensity was low and shelter was sufficient to prevent rapid Pteridium litter loss, the necessary treatments to establish Calluna were to disturb the litter and add a source of Calluna seeds. A difference between continued Pteridium control or a relaxation of control was not observed over the time‐scale of the experiment. At the second site, where grazing intensity was high and shelter minimal, vegetation dominated by Rumex acetosella had developed on a thin but dense litter layer. To enhance Calluna regeneration it was necessary to prevent sheep grazing, disturb the remaining litter layer and add a source of Calluna propagules. A nurse crop hindered Calluna establishment due to its vigorous growth. At neither site was an initial application of fertilizer an aid to Calluna establishment. It is possible to devise methods for the re‐establishment of Calluna at sites where previous attempts at Pteridium control alone have not had the desired effect on vegetation development. However, the precise choice of methods used is dependent on the grazing management at the site. High levels of livestock grazing prevented Calluna establishment, but some trampling may aid litter breakdown. The ability to adjust stocking rates and access to the site by machinery are critical factors to take into account in producing an effective strategy for long‐term Pteridium control and its replacement by target vegetation.     

Expression of two heterologous promoters, Agrobacterium rhizogenes rolC and cauliflower mosaic virus 35S, in the stem of transgenic hybrid aspen plants during the annual cycle of growth and dormancy

We monitored, for the first time, the activity of two model heterologous promoters, the Agrobacterium rhizogenes rolC and the cauliflower mosaic virus (CaMV) 35S, throughout the annual cycle of growth and dormancy in a perennial species, hybrid aspen. Each promoter was fused to the uidA -glucuronidase (GUS) reporter gene and the constructs were introduced into the hybrid aspen genome by Agrobacterium-mediated transformation. Both wildtype and transgenic plants were cultivated under different regimes of photoperiod and temperature to induce passage through one growth-dormancy-reactivation cycle, and at intervals GUS staining was assessed in stem sections. In rolC::uidA transformants, GUS activity in rapidly growing current-year shoots was not only tissue-specific, being localized to the phloem, but also cell-specific at the shoot base, where it was present only in the companion cells. However, during the onset of dormancy induced by short photoperiod, GUS activity shifted laterally from the phloem to include the cortex and pith. After subsequent exposure to chilling temperatures to induce the transition between the dormancy stages of rest and quiescence, GUS activity almost disappeared from all stem tissues, but regained its original phloem specificity and intensity after the shoots were reactivated by exposing them to long photoperiod and high temperatures. In contrast, GUS activity in the stem of 35S::uidA transformants was strong in all tissues except for the vascular cambium and xylem, and did not vary in intensity during the growth-dormancy-reactivation cycle. The lateral shift and increased intensity of GUS activity in the stem of rolC::uidA transformants during dormancy induction was shown to be associated with the accumulation of starch, and to be mimicked by incubating stem sections in sucrose, as well as glucose and fructose, but not sorbitol, prior to the GUS assay. Our results demonstrate that the activities of the rolC and 35S promoters varied in very different, unpredictable ways during the annual cycle of growth and dormancy in a perennial species, and indicate that the spatial and temporal variation in rolC promoter activity that we observed in the stem of transgenic hybrid aspen plants is attributable to cellular and seasonal changes in sucrose content.     

桃芽休眠的自然诱导因子及钙在休眠诱导中的作用

以2年生低需冷量设施栽培适宜品种“春捷毛桃”为试材,研究桃植株进入休眠和发展抗冻性的自然诱导因子及Ca²⁺在此过程中的作用.结果表明：短日照和自然低温单个因子或其共同作用均能诱导桃植株停止生长,然后进入休眠状态和发展抗冻性,但短日照和自然低温的作用机制不同.短日照首先诱导桃植株进入休眠状态,然后诱导抗冻性发展;而自然低温则是首先诱导桃植株抗冻性发展,而后诱导其进入休眠状态;当短日照和自然低温共同作用即自然条件下时,短日照起主导作用,自然低温起辅助促进作用.短日照处理研究表明,短日照诱导桃植株停止生长、休眠诱导和抗冻性发展离不开Ca²⁺的作用,在此过程中Ca²⁺起着传递短日照信号的作用;在人工补光长日照条件下,随着温度的降低,Ca²⁺逐渐由细胞间隙、细胞壁和液泡流入细胞质和核内,同时植株生长速度变缓并最终完全停止,随后进入休眠状态并发展抗冻性.表明Ca²⁺作为传递自然低温信号的信使在自然低温诱导桃植株停止生长、抗冻性发展和休眠过程中起着重要作用.     

Growth and Dormancy in Lunularia cruciata (L.) Dum.: II. THE RESPONSE TO DAYLENGTH AND TEMPERATURE

In the Israel strain of Lunularia under investigation long-daytreatment induces dormancy while short-day breaks it. Daylengthis perceived by all parts of the thallus, and even by younggemmae while in the cup of the mother thallus. Since light-breaktreatment is equivalent to long-day the effect is truly photoperiodic,but the critical daylength is not well defined. Temperatureinteracts strongly with photoperiod. High temperature (24°C.) combined with continuous light rapidly induces dormancy(c. 6 days). Fewer short days are required to restart growth,and the number (3–4) is unaltered by the duration of thecontinuous light treatment preceding it. Rinsing experimentssuggest that at least some of the inhibitory factor producedin long photoperiods can be leached from the plant. Dormancy-inducingtreatment quantitatively confers the capacity to resist drought(dormant plants survive air-drying, while actively growing thalliare killed in 80 per cent. R.H.). The photoperiodic effects on growth and dormancy are discussedand an inhibitor hypothesis is suggested to explain the responsesfound.     

Dormancy in Dioscorea: Range, Duration and Timing of High-Temperature Treatment in Germination Inhibition of D. tokoro Seeds

The effects of temperature on induction and release of high-temperatureinhibition in seed germination of Dioscorea tokoro Makino, amonocotyledonous summer perennial of the temperate zone of EastAsia, were investigated. Germination was increasingly inhibitedwith elevation of temperature over 23°C and lengtheningof its duration. The low temperature limit for germination inhibitiondecreased with lengthening of the duration of high temperature.The most sensitive phase for high temperature was 1–2days after the start of imbibition at 20°C. The germination inhibition by high temperature was reversedby chilling at 5°C, which is the optimum temperature forbreaking the natural dormancy (primary dormancy) of this seed.This showed that the high-temperature inhibition of germinationdoes not cause mortal damage but only secondary dormancy (induceddormancy). Seeds from a cold climate (Miyagi Pref.) responded rather quicklyto both high temperature and chilling compared to seeds froma warm climate (Kagoshima Pref.). The responsiveness to hightemperature and chilling of D. tokoro seed may affect the germinationtime under natural conditions. (Received October 22, 1982; Accepted January 14, 1983)     

Dormancy and Flowering in Anemone coronaria L. as Affected by Photoperiod and Temperature

The effects of day-length and temperature on flowering and dormancyinduction were studied in Anemone coronaria L., with plantsraised either from corms or achenes. An Israeli hybrid sourcewas used (de Caen cv. Hollandia x Israeli wild type). Dormancy onset is characterized by the cessation of foliageleaf production, the appearance of leaf scales protecting theperennating bud, and leaf senescence. Dormancy was induced byhigh temperature and long days but increasing temperatures (from17/12 °C to 32/12 °C) induced earlier dormancy thanprolonging the photoperiod (range 8–16 h). A significant(P = 0.01) interaction was found between these factors, withsmaller photoperiodic effects the higher the temperature. At22/17 °C the critical day-length for dormancy inductionwas between 11 and 12 h. The transition from the vegetative to the reproductive stageappears to be an autonomous process that occurs with developmentin plants raised from either corms or achenes and does not requireenvironmental induction. Photo- and thermoperiodic effects onflowering were indirect, being mediated through their influenceon dormancy induction. Anemone coronaria L., dormancy, flowering, photoperiod, thermoperiod