Photoperiodic control of flowering in Dactylis glomerata, a true short-long-day plant

Flowering requirements of three Scandinavian cultivars of Dactylis glomerata L. have been studied in controlled environments. At temperatures ranging from 9 to 21°C optimal flowering required 10 weeks of exposure to short days (SD) followed by exposure to long days (LD). Only a few plants flowered in continuous LD and no primary induction took place in any daylength at 24 or 27°C. However, at a temperature of 3°C primary induction occurred also in 24 h LD, but more than 20 weeks of treatment were required for 100% flowering. The critical photoperiod for secondary induction was about 12–13 h, depending on the latitude of origin of the cultivar. A critical number of 12 to 16 LD cycles was required for 100% flowering, although some plants flowered after only 4 LD. A high proportion of viviparous proliferation resulted from marginal LD induction. Initiation of floral primordia did not take place in SD but required a transition from SD to LD. These results demonstrate that D. glomerata is a true short-long-day plant.     

Dual induction control of flowering in Leucanthemum vulgare

The perennial herb Leucanthemum vulgare (oxeye daisy) has a dual induction requirement for flowering. The primary induction is a typical low temperature vemalization response. Temperatures up to 15°C are effective, and the optimum is 6–9°C. Short days (SD) during low temperature exposure enhanced primary induction, but SD could not fully substitute for low temperature in primary induction. At optimum temperatures about 6 weeks exposure were required for 100% flowering, but the flowering response increased with increasing exposure up to 12 weeks, especially at higher temperatures. Seedling have a short juvenile phase of about 4 weeks. Populations with origin ranging from 59 to 69°N in Norway did not vary in their primary induction requirements. Long days (LD) were required for inflorescence initiation and stem elongation at 9°C. At 21 and 15°C some plants initiated and developed inflorescences in SD, but the inflorescences were sessile and their development strongly delayed. More than 16 LD cycles were required for normal stem elongation (bolting).     

Flowering requirements in Bromus inermis, a short-long-day plant

Smooth bromegrass plants ( Bromus inermis Leyss.) have a dual photoperiodic requirement for flowering. At temperatures ranging from 6 to 24°C, short days (SD) are necessary for primary induction while a transition to long days (LD) is required for initiation of flower primordia, culm elongation and flower development (secondary induction). Critical photoperiods for primary induction (50% flowering) were 13.5 h (15°C) and 12 h (24°C) in the American cv. Manchar and 14.5 and 13 h, respectively, in the Norwegian cv. Löfar. For the secondary induction the respective critical photoperiods were 14 and 15 h in 'Manchar' and 16 and 17.5 h in 'Löar', which also appeared to be better adapted to low temperatures. Low temperature vernalization in LD for up to 16 weeks at 3°C was unable to cause primary induction and temperatures below 12°C also strongly reduced the SD effect. At optimum temperature (15-2TC) 4 to 6 weeks of 8-10 h SD treatments were needed for optimal primary induction effect. A minimum of 8 LD cycles of 24 h were required for complete secondary induction in 'Manchar', while more than 16 cycles were needed in 'Löfar'. Seedlings grown in SD developed a rosette type of growth with shoots growing in a decumbent position, while those in LD grew upright and formed elongated vegetative culms. Rate of leaf initiation was enhanced by about 60% by LD while tillering was promoted by SD.     

Interactions of temperature and photoperiod in the control of flowering of latitudinal and altitudinal populations of wild strawberry (Fragaria vesca)

Floral induction and development requirements of a range of latitudinal and altitudinal Norwegian populations of the wild strawberry Fragaria vesca L. have been studied in controlled environments. Rooted runner plants were exposed to a range of photoperiods and temperatures for 5 weeks for floral induction and then transferred to long day (LD) at 20°C for flower development. A pronounced interaction of temperature and photoperiod was shown in the control of flowering. At 9°C, flowers were initiated in both short day (SD) and LD conditions, at 15 and 18°C in SD only, whereas no initiation took place at 21°C regardless of daylength conditions. The critical photoperiod for SD floral induction was about 16 h and 14 h at 15 and 18°C, respectively, the induction being incomplete at 18°C. The optimal condition for floral induction was SD at 15°C. A minimum of 4 weeks of exposure to such optimal conditions was required. Although the populations varied significantly in their flowering performance, no clinal relationship was present between latitude of origin and critical photoperiod. Flower development of SD-induced plants was only marginally advanced by LD conditions, while inflorescence elongation and runnering were strongly enhanced by LD at this stage. The main shift in these responses took place at photoperiods between 16 and 17 h. Unlike all other populations studied, a high-latitude population from 70°N ('Alta') had an obligatory vernalization requirement. Although flowering and fruiting in its native Subarctic environment and after overwintering in the field in south Norway, this population did not flower in the laboratory in the absence of vernalization, even with 10 or 15 weeks of exposure to SD at 9°C. Flowering performance in the field likewise indicated a vernalization requirement of this high-latitude population.     

Flowering strategies of the high-arctic and high-alpine snow bed grass species Phippsia algida

Flowering requirements of the high-arctic and high-alpine snow bed grass species Phippsia algida (Sol.) R. Br. have been studied in controlled environments. Seedlings flowered rapidly in continuous long days (LD) at temperatures ranging from 9 to 21°C. They also initiated inflorescence primordia at the same temperatures in continuous short days (SD), whereas LD were required for heading and anthesis. The plant thus has the characteristics of a regular long day plant, although the daylength requirement is associated with floral development only. The critical daylength for the LD response was about 17 h at 21°C and 19 h at 9°C. A single LD cycle was enough to trigger inflorescence development, while 5 cycles were required for the full response. Anthesis was reached within a week of LD treatment at 21°C in SD grown plants with preformed inflorescence primordia. The advantages of these versatile flowering responses are discussed in relation to the extreme climatic regime of late snow bed sites.     

Environmental control of flowering and sex expression in Carex flava

The environmental control of flowering and sex expression has been studied under controlled environment conditions in three populations of the sedge Carex flava L. A dual floral induction requirement was demonstrated in all populations. Low temperature (< 12°C) was obligatory for, and short photoperiods strongly enhanced, primary induction and inflorescence initiation. Stem elongation and inflorescence development were promoted by long photoperiods, although most plants developed stunted flower stems also under short day (SD) conditions. Growth vigour, abundance of flowering and primary induction requirements varied widely among the populations, with critical exposure times for full flowering varying from less than 9 to about 12 weeks in SD at 9°C, and from about 9 to more than 15 weeks in long days (LD). Sex expression in the normally male terminal spike was shifted towards femaleness by marginal or incomplete primary induction. Primary induction in LD resulted in a complete change to entirely female inflorescences, whereas marginal induction in SD resulted in a similar sex reversal in some plants. The results are discussed in relation to environmental and hormonal factors known to modify sex expression in flowering plants and the significance of the results to Carex systematics and classification.     

Inflorescence commitment and subsequent development differ in their responses to temperature and photoperiod in Osteospermum jucundum

Although Osteospermum is a species which is known to require a period of chilling to induce flowering, the precise form of the relationships between temperature and photoperiod on the phases of flowering has not been quantified. This study aimed to investigate the effects of temperature and photoperiod on time to inflorescence commitment and on the rate of subsequent floral development in Osteospermum jucundum cv. Zulu. To assess how temperature and photoperiod affected the number of days needed for inflorescence commitment, plants were transferred from a range of photothermal environments to a non‐inductive environment. The effect of temperature and photoperiod on subsequent inflorescence development was examined by transferring plants with initiated inflorescences to a range of photothermal environments. Inflorescence commitment occurred first in plants grown at a low average diurnal temperature (10.6°C), but no evidence was found to suggest that photoperiod affected the duration of this phase. Once initiated, high temperatures and long days hastened inflorescence development. The rate of progress to flowering from initiation increased linearly with photoperiod and temperature (up to an optimum of 23.5°C).     

Dual Floral Induction Requirements in Phleum alpinum

Flowering requirements of four Norwegian populations of Phleumalpinum were studied in controlled environments. A dual inductionrequirement was demonstrated in all populations. Inflorescenceinitiation had an obligatory requirement for short days (SD)and/or low temperature, while culm elongation and heading wereenhanced by long days (LD) and higher temperatures. At 3 and6 °C primary induction was almost independent of photoperiod,whereas SD was more effective than LD at higher temperatures.The critical temperature for primary induction was about 15°C in SD and 12 °C in LD. Saturation of induction required12 weeks of exposure to inductive conditions, although someheading and flowering took place with 6 weeks exposure to optimalconditions (9 °C/SD). Inflorescence development also tookplace in 8 h SD although it was delayed and culm elongationwas strongly inhibited compared with LD conditions. Only smalldifferences in flowering response were found between the populations. Phleum alpinum L., alpine timothy, dual floral induction, flowering, photoperiod, temperature     

Primary and secondary induction requirements for flowering in Alopecurus pratensis

The critical temperature for primary induction of flowering in two Norwegian eco-types of meadow foxtail ( Alopecurus pratensis L.) was about 21°C, with 6 weeks induction period. Inflorescence primordia were initiated with increasing effectiveness as the temperature was reduced below this level in both short days (SD) and long days (LD), although SD was the more effective photoperiod at all temperatures above 6°C. The degree of primary induction was closely negatively correlated with the length of basal leaves. Culm elongation and heading (secondary induction) were promoted by LD and high temperature and inhibited by the combination of SD and low temperature. The relationship between SD primary induction and vernalization is discussed in the light of these and other results.     

Photoperiodic control of dormancy in Sedum telephium and some other herbaceous perennial plants

The environmental control of dormancy and flowering of the herbaceous perennial Sedum telephium was studied in controlled environments. Short photoperiods induced growth cessation and the formation of resting buds in both seedlings and mature plants, whereas long photoperiods resulted in immediate growth activation of dormant buds. No chilling was required for dormancy release, even in plants induced to dormancy and maintained at high temperature (21°C) for more than 3 months. The critical photoperiod for dormancy release was about 15 h, a minimum of four long-day (LD) cycles (24 h) being required. The true photoperiodic nature of this response was ascertained by night interruption experiments. Flowering of S. telephium was found to have an obligatory LD requirement, with no requirement for vernalization. The critical photoperiod and minimum number of inductive cycles for floral induction were the same as for dormancy release. Dormancy release by long days was also obtained in preliminary experiments with three other herbaceous perennials. The eco-physiological significance of photoperiodic control of dormancy is discussed, and it is concluded that the mechanism ensures stability of winter dormancy, even under conditions of climatic warming.     

Environmental Control of Flowering in some Northern Carex Species

The environmental control of flowering in some arctic-alpineCarexspecies has been studied in controlled environments.Carex nigra,C. brunnescens, C. atrata, C. norwegica andC. serotina all hada dual induction requirement for flowering. In all exceptC.nigra either low temperature (12 °C or lower) or short days(SD) over a wider range of temperatures were needed for primaryfloral induction and inflorescence formation. InC. nigra primaryfloral induction took place in SD only (9–21 °C),8–10 weeks of exposure being required for a full response.In all these species long days (LD) were required for, or stronglypromoted, culm elongation and inflorescence development (secondaryinduction). Quantitative ecotype differences in both primaryand secondary induction were demonstrated. Unlike the otherspecies,C. bicolor proved to be a regular LD plant which requiredLD only for inflorescence initiation and development. In allspecies leaf growth was strongly promoted by LD, especiallyin the higher temperature range (15–21 °C). In SDand temperatures below 15 °C the leaves became senescentand the plants entered a semi-dormant condition which was immediatelyreversed by LD. The results are discussed in relation to growthform and life history of shoots. Carex ; dual induction; ecotypic diversity; flowering; growth; photoperiod; sedges; temperature     

Environmental control of flowering and morphology in the high-arctic Cerastium regelii, and the taxonomic status of C. jenisejense

Cerastium regelii has a distribution confined largely to regions north of 70° N but has retained a strong short-day (SD) response for primary flower induction despite the fact that it will hardly ever experience SD in a non-frozen condition in its natural environment. However, like many other high-latitude short-long-day plants it has also an alterntive long day (LD) pathway for floral initiation at low temperatures (<15°C). Floral primordia which are fully differentiated during SD have an absolute requirement for LD for flower development. The critical photoperiod for this LD response is about 16 h at 18°C and more than 20 h at 9°C. Plant morphology, including key characters for identification of the species, is greatly modified by environment and stage of plant development. At higher temperatures and LD C. regelii develops a striking resemblance to the sub-arctic C. jenisejense . Based on examination of authentic herbarium material it is concluded that the latter is merely a high-temperature morphotype of C. regelii .     

The sensitivity of larval Plodia interpunctella and Ephestia elutella (Lepidoptera) to light during the photoperiodic induction of diapause

ABSTRACT. The incidence of diapause in larvae of Plodia interpunctella and Ephestia elutella held under two light systems was examined. Both systems progressively shortened the photophase of 24-h cycles, one with a motorized dimming switch providing dawns and dusks about 1 h long, the other switching the lights instantaneously. The mean critical photoperiod for P. interpunctella was about 131/4 h and for E. elutella just over 14 h. In both species light intensities as low as 0.2 lx influenced the induction of diapause. In P. interpunctella the critical photoperiod and sensitivity to light were similar at 23.±.;5°C and 20.5±0.5°C. At 22.5°C the percentage of diapausing larvae of E. elutella increased from 2% in long photoperiods (> 15 h light), to 100% in short photo-periods (t 12.5h light). Fox P. interpunctella , at 22.5°C the percentage increased from zero in long photoperiods (> 14 h light) to about 98% in short photoperiods (< 11.5h light), and at 20°C from 12% to 100% over a similar photoperiodic range. Similar results were obtained under selected fixed photoperiods, switched on or off instantaneously.     

Climatic flowering requirements of bipolar sedges, Carex spp., and the feasibility of their trans-equatorial migration by mountain-hopping

A selection of northern and southern hemisphere populations of a range of bipolar Carex species was grown in controlled environments for comparison of their temperature and daylength requirements for flowering. All studied species have basically the same dual induction requirements for flowering but differ quantitatively in their respective critical temperatures and daylengths for primary and secondary floral induction. Usually, low temperature or short days (SD), over a wider range of temperatures, were required for initiation of inflorescence primordia, whereas long days (LD) advanced but were not obligatory for heading and flower development. It is concluded that no change or adaptation in the flowering requirements would have been necessary for these plants to migrate across the tropics and their penetration into the southern hemisphere by mountain-hopping. The flowering requirements of these and a range of other bipolar plant species are compatible with the hypothesis that absence of an obligatory long day floral induction requirement has facilitated or been a prerequisite for trans-equatorial migration of plants by mountain-hopping and the establishment of bipolar species distributions.     

Effects of temperature and photoperiod on flowering in Hyptis brevipes

Few tropical species have been tested for their flowering response under controlled conditions. Hyptis brevipes Poit, is an annual herb, commonly found in wet margins of streams and ponds, being considered a weed for some perennial plantations in Brazil. Under experimental glasshouse conditions, this species proved to be an obligate short-day plant. Flowering was delayed when photoperiods longer than 8 h were given, the critical photoperiod being between 12 and 13 h. When both temperature and photoperiod were controlled, at 20°C a longer protoperiod (by almost 1 h) is still inductive compared to 25 and 30°C. The number of short-day cycles required for full induction is relatively high and dependent upon temperature; at 20°C or above, 10 cycles are adequate, but at 15°C, more short-day cycles are needed. The number of inflorescences formed as well as the floral index vary according to daylength × temperature × inductive cycle number, allowing flowering to be assessed quantitatively. Long days are inhibitory to flowering, either suppressing it completely (when symmetrically intercalated among 24 inductive cycles) or preventing the floral index from increasing.     

Remobilization of fructans in Phippsia algida during rapid inflorescence development

Plants of Phippsia algida (Sol.) R. Br. were cultivated in short days (SD; 8 h summer daylight) and in long days (LD; 8 h summer daylight + 16 h low irradiance extension of 5 μmol m⁻² s⁻¹) at 9, 15, and 21°C. In this plant, inflorescence primordia are initiated in both LD and SD, but LD are required for heading and inflorescence development (Heide, O.M.; Physiol. Plant. 85: 606–610. 1992). Total dry matter production was slightly increased by LD over SD at 9°C, while it was little affected by daylength at 15 and 21°C. Phippsia algida contained mainly fructans with a low degree of polymerization, largely of the kestose series. After 29 to 42 days (depending on the temperatature) of photoperiodic treatment, fructans constituted 15–20 percent of dry mass of SD-grown plants compared with only 2–3 percent of dry mass for LD-grown flowering plants. There was no difference due to photoperiod in levels of mono- and disaccharides. Shifting the SD-grown plants to LD conditions resulted in rapid inflorescence development, accompanied by a parallel rapid decrease in the fructan level, while the level of mono- and disaccharides remained constant. The results show that fructans are important as storage carbohydrates in the late snow-bed species P. algida that normally requires several growing seasons for completing its life cycle. Exhaustion of this storage pool during the extremely fast flower and fruit development constitutes an essential part of the plants adaption to a very short growing season.     

Effects of Photoperiod, Nitrogen Nutrition and Temperature on Inflorescence Initiation and Development in Onion (Allium cepa L.)

The effects of photoperiod, nitrogen nutrition and temperatureon inflorescence initiation and development in onion cv. Rijnsburgerand cv. Senshyu Semi-globe Yellow were studied in controlledenvironments. Rates of inflorescence initiation were estimatedusing the data for leaf numbers formed prior to flower formationand the rates of leaf initiation. At 9 °C inflorescenceinitiation was accelerated by long photoperiods particularlyfor cv. Rijnsburger where the average time for initiation was86 days in 8 h and 38 days in 20 h photoperiods. Initiationwas as rapid at 12 °C as at 9 °C but was slower at 6°C. A reduction in the nitrate concentration in the nutrientsolution from 0.012 to 0.0018 M greatly accelerated inflorescenceinitiation particularly in photoperiods and temperatures notconducive to rapid initiation. Cv. Senshyu initiated more slowlythan cv. Rijnsburger and was less sensitive to photoperiod andnitrogen level. The development rate of inflorescences afterinitiation was accelerated by long photoperiods and increasesin temperature from 6 to 12 °C but was retarded by the lowernitrogen level. Allium cepa L., onion, flower initiation, inflorescence development, photoperiod, nitrogen nutrition, temperature, vernalization     

Studies in the Physiology of the Onion Plant: IV. THE INFLUENCE OF DAY-LENGTH AND TEMPERATURE ON THE FLOWERING OF THE ONION PLANT

At temperatures above about 17° C. inflorescence initiationin growing onion plants, as in stored sets, is suppressed whetherthe plants are kept in long or short days. Independently ofcurrent day-length and of previous day-length treatment, ifthe plants are sufficiently large initiation begins very shortlyafter the temperature falls below c. 15° C. Emerged infiorescencesappear some ten or so weeks later. Small plants are unable toinitiate inflores cences under any of the conditions tested,and actual size (perhaps leaf area) rather than leaf or nodenumber seems to be the important factor. Inflorescence emergenceis suppressed at high temperatures in short days or long days;in long days bulb formation also suppresses emergence at lowertemperatures. In long days at temperatures sufficiently lowfor bulbing to be delayed, however, emergence is accelerated.Plants which have produced bulbs in long days in the summershow a delay of inflorescence emergence in the following winter.     

Photothermal control of the imposition of summer dormancy in Poabulbosa, a perennial grass geophyte

Poa bulbosa L., like many other Mediterranean geophytes, grows in the winter and enters a phase of summer dormancy in the spring. Summer dormancy enables these plants to survive the hot and dry summer. Long days are the main environmental factor active in the induction of summer dormancy in P . bulbosa and elevated temperatures accelerate dormancy development. P . bulbosa becomes dormant earlier than most other species that grow actively in the winter. Previous studies suggested that pre-exposure of P . bulbosa to short days and low temperatures during the autumn and early winter increased its sensitivity to photoperiodic induction in late winter, and thus enabled the early imposition of dormancy. To study this hypothesis, experiments were carried out under controlled photothermal conditions in the phytotron, under natural daylight extended with artificial lighting. The critical photoperiod for induction of summer dormancy at an optimal temperature (22/17°C day/night) was between 11 and 12 h. Photoperiods shorter than 12 h were noninductive, while 14- and 16-h days were fully inductive. A night break of 1 h of light given at the middle of the dark period of an 8-h photoperiod also resulted in full induction of dormancy. Pre-exposure to either low temperature (chilling at 5°C) or to short days of 8 h (SD) enhanced the inductive effect of subsequent 16-h long days (LD). The enhancing effect of chilling and SD increased with longer duration, i.e. fewer LDs were required to impose dormancy. However, the day-length during the low-temperature pretreatment had no effect on the level of induction at the following LD. Chilling followed by SD did not induce dormancy. The relevance of these responses to the development and survival of P . bulbosa in its natural habitat is discussed.     

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