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.     

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.     

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).     

Developmental strategies of Koenigia islandica, a high-arctic annual plant

Seed germination, growth and flowering of the arctic-alpine annual Koenigia islandica were studied in controlled environment. Intact (unabraded) seeds germinated poorely at temperatures up to 18°C, with an optimum at 24°C (89% in 10 d). Scarified seeds germinated rapidly, and reached 100% germination in 3 d at 21°C, but no >40% germination occurred at 9 and 12°C, The seeds had no light requirement for germination, nor did fluctuating temperatures improve germination
Dry matter production was optimal at 12°C in both short day (SD) and long day (LD) conditions, but was markedly higher in LD than in SD at identical fluences at all temperatures except 21°C where the plants showed symptoms of severe heat stress. The temperature compensation point for net productivity was estimated to 24°C, and negative carbon balance at higher temperatures might be an important physiological mechanism limiting the distribution of K. islandica in Scandinavia.
Flowering was extremely rapid and independent of daylength, even in a high-arctic population from 79°N, In full summer daylight anthesis was reached 24 d after germination and seeds ripened after 36 d at 15°C, Days to anthesis varied little across the temperature range from 6 to 21°C, giving a linear decrease in the heat-sum requirement for the attainment of flowering with decreasing temperature.
It is concluded that conservative seed germination strategy, tininess and rapid development, low temperature optima for growth and reproduction, and daylength indifference of flowering are important adaptations for success of an annual plant in high-arctic and high-alpine environments, Daylength neutrality has facilitated the wide-latitudinal distribution of K. islandica. including the penetration of the species to the southern hemisphere.     

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.     

Photoperiodic and temperature control of diapause induction and colour change in the southern green stink bug Nezara viridula

Abstract. The effect of photoperiod and temperature on the duration of the nymphal period, diapause induction and colour change in adults of Nezara viridula (L.) (Heteroptera: Pentatomidae) from Japan was studied in the laboratory. At 20 °C, the developmental period for nymphs was significantly shorter under LD 10 : 14 h (short day) and LD 16 : 8 h (long day) than under intermediate photoperiods, whereas at 25 °C it was slightly shorter under intermediate than short- and long-day conditions. It is assumed that photoperiod-mediated acceleration of nymphal growth takes place in autumn when day-length is short and it is unlikely that nymphal development is affected by day-length under summer long-day and hot conditions. Nezara viridula has an adult diapause controlled by a long-day photoperiodic response. At 20 °C and 25 °C in both sexes, photoperiodic responses were similar and had thresholds close to 12.5 h, thus suggesting that the response is thermostable within this range of temperatures and day-length plays a leading role in diapause induction. Precopulation and preoviposition periods were significantly longer under near-critical regimes than under long-day ones. Short-day and near-critical photoperiods induced a gradual change of adult colour from green to brown/russet. The rate of colour change was significantly higher under LD 10 : 14 h than under LD 13 : 11 h, suggesting that the colour change is strongly associated with diapause induction. The incidences of diapause or dark colour did not vary among genetically determined colour morphs, indicating that these morphs have a similar tendency to enter diapause and change colour in response to short-day conditions.     

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.     

Effect of temperature on photoperiodic response in a selected 'non-diapause' strain of Pyrrhocoris apterus (Heteroptera)

Abstract. The artificially selected 'non-diapause' strain of Pyrrhocoris apterus (L.) (Heteroptera) showed no diapause response to photoperiod at 26°C (Socha & Hodkova, 1994). However, the diapause response to short-day photoperiod (LD 12:12 h) became apparent at lower temperatures of 17°C (70% diapause) or 20°C (41% diapause). Diapause was induced in 60% females by short-day photoperiod combined with thermoperiod of 26/16°C, whereas only 20% diapause was induced by the same thermoperiod under continuous darkness. Thus the time-measuring system was not removed by artificial selection but the diapause response was shifted to lower temperatures. The diapause response to short days seems to be favoured rather by low temperature during scotophase than by low temperature throughout the whole light/dark cycle. If the percentage of diapause at 26°C is compared in F₁ hybrids and in wild and selected parental strains the diapause appears to be dominant at LD 13:11 h but recessive at LD 11:13 h and LD 10:14 h. A hypothesis is proposed that the inheritance of the percentage of diapause in F1 hybrids is determined by interactions of genes controlling the temperature dependence of photoperiodic response.     

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.     

Diapause induction and coloration in the Senegalese grasshopper, Oedaleus senegalensis

Abstract. Embryonic diapause induction in the Senegalese grasshopper, Oedaleus senegalensis Krauss (Orthoptera: Acrididae), is influenced both by the photoperiod and the temperature experienced by females. High temperatures (40°C) and long photoperiods (LD 14:10h), which characterize the beginning of the rainy season in the Sahel, cause non-diapausing eggs to be laid. Lower temperatures (25°C) and shorter photoperiods (LD 12:12h), which occur at the end of the rains, result in the production of diapausing eggs. At 30°C and constant photoperiods, O. senegalensis exhibited a long-day-short-day response with critical photoperiods of c. 13 h and c. 20 h, only the former value being of ecological significance. The photoperiodically sensitive stages to diapause induction in females occurred from the fifth stadium onwards. Temperature also affected the coloration of both nymphs and adults. Dark-black and pale-white individuals were produced by low (25°C) and high (40°C) temperatures respectively, whereas an intermediate temperature (30°C) produced individuals which were greyish brown. These results are discussed in relation to the ecology of O. senegalensis.     

Seed Germination in Amaranthus retroflexus L. as Affected by the Photoperiod and Age During Flower Induction of the Parent Plants

Seed germination in Amaranthtis retroflexus, a facultative shortday plant, was affected by the parental photoperiodic conditions.Seeds from parents grown continuously in short days (SD, 8 h)had a higher dark germination and a greater response (at 30°C) to a short irradiation or low temperature pretreatmentthan seeds from plants grown continuously in long days (LD,16 h). Daily night breaks of 1 h in the middle of the long-nightinhibited the SD induction of flowering as well as the SD promotionof germinability. Germinability of seeds produced by plantsinduced to flower in LD by 1, 2, or 3 SD was lower than thatof seeds produced by plants grown continuously in SD, and decreasedwith the age of the parent plants at the time of flower induction.     

Effects of photoperiod and temperature on sugars and fructans in leaf blades, leaf sheaths and stems, and roots in relation to growth of Poa pratensis

Vegetative plants of Poa pratensis L. cv. Holt were cultivated in short days (SD; 8 h summer daylight) and in long days (LD; 8 h summer daylight + low intensity extension of 5 μmol m^-2 s^-1) at 12, 18 and 24°C in one experiment and at 9, 12, 15 and 18°C in another. Relative growth rate (RGR) as the mean of both experiments and all temperatures was 32% higher in LD than in SD between start of daylength treatment and first harvest, and 18% higher in LD than in SD between first and second harvest. Early in the daylength treatment period, more assimilates were allocated to storage in SD than in LD, so that at first harvest leaf sheaths and stems had 175% higher concentration of fructans in SD. Later this allocation pattern changed, and for the larger plants at the second harvest the differences in fructan concentrations were much smaller between the two daylengths. Both sugar and fructan concentrations were highest at low temperatures. The distribution of sugars and fructans varied from mostly sugars in the leaves to mostly fructans in leaf sheaths and stems and roots. The fructans were mainly high degree polymerization fructans. At least two series of fructans were present, and the dominant one was probably based upon kestose. It is concluded that allocation of assimilates to growth in leaf area instead of to storage may be important for the observed LD stimulation of dry matter production.     

Temperature and daylength control of flower initiation in winter oilseed rape (Brassica napus L.)

The influence of low temperature and daylength on pre-floral growth and flower initiation in winter oilseed rape cv. Mikado was examined under controlled environment conditions at the University of Newcastle upon Tyne during 1985 and 1986.
The vernalisation requirement of Mikado was most effectively fulfilled by temperatures of 6 °C and 9 °C. Plants maintained at both higher and lower temperatures had an extended pre-floral growth phase. The transition from vegetative to reproductive growth in plants maintained at 12 °C was delayed by slow accumulation of the cold requirement, whereas flower initiation appeared to be delayed by limited leaf production, dry matter accumulation and/or assimilate availability in plants grown at 3 °C. The mechanism of floral induction remained unresolved but it was clear that flower initiation was not controlled by low temperature per se . Short days partially substituted for the cold requirement at 12 °C but photoperiodic induction of flower initiation was less important than the influence of low temperature.     

Diapause in a Glasgow strain of the flour moth, Ephestia kuehniella

ABSTRACT. The incidence of diapause in Ephestia kuehniella Zeller from an unhealed granary in Scotland was influenced by both photoperiod and temperature. At 25°C, nearly 50% of larvae entered diapause when reared in continuous darkness (DD) and up to 30% did so in short photoperiods. Little diapause was detected around LD 14:10 but a second, smaller peak of about 20% occurred at LD 16:8 and LD 18:6, falling away again to nearly zero in continuous light. More larvae entered diapause when reared continuously at 15 or 20°C than at 25 and 30°C. However, when larvae reared from hatch at 25°C in LD 16:8 were transferred after 1 week to 15°C in LD 9:15, almost twice as many entered diapause as did those reared at 15°C throughout. The sensitive phase for diapause induction occurred near the start of the final instar. The mean duration of diapause was between 2 and 3 months in most photoperiods at 20 and 25°C, and was shorter at 15°C. However, in DD at 25°C, it lasted about 7 months. Termination of diapause was hastened in larvae reared at 25°C in DD by transferring them to LD 14:10, and also by chilling them at 7.5°C for 6 weeks before returning to 25°C in DD. In an unhealed store in southern England, viable adults emerged from May to July and originated from larvae which terminated diapause relatively late. It would appear from the results of transferring larvae back to the laboratory at various times during the winter that some phases of diapause development were completed quite early after exposure to low temperatures, although no further development took place in the store until temperatures rose again in April.     

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.     

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     

The effect of temperature and light pulses on the induction of diapause in the Toyama strain of the Indian meal moth, Plodia interpunctella

Abstract.The photoperiodic response in Plodia interpunctella collected at Toyama (36.7°N) was of long-day type and highly sensitive to temperature. The critical photoperiod giving 50% diapause was between 14 and 16 h at 20°C, between 12 and 14 h at 25°C and between 6 and 8 h at 30°C. Effects of night interruption by a 2-h light pulse on the diapause response were examined at 25°C on different background photoperiods ranging from LD 12:12 h to LD 2:22 h. Percentage diapause was very low when the middle portion of dark period was interrupted, so that U- or V-shaped response curves were obtained with background scotophases longer than 12 h. In these curves, the descending slopes were less steep than the ascending slopes. The critical dark period measured from dusk to an interrupting light pulse was about 1.5 h longer than the critical dark period ( c . 10 h) in the normal photoperiodic response. The critical dark period from the interrupting light pulse to dawn, on the other hand, was not parallel to dawn but shorter than the normal critical period in LD 12:12 h and LD 10:14 h and longer than that in LD 7:17 h to LD 4:20 h, indicating that the priming effects of the light pulse might be under the influence of the photophase.     

Effects of photoperiod and temperature on the development of Bonagota cranaodes

Abstract.  The Brazilian apple leafroller, Bonagota cranaodes (Meyrick) (Lepidoptera: Tortricidae) is reared in the laboratory under a long-day (LD 14 : 10 h) and a short-day (LD 7 : 17 h) photoperiod at 22 °C, and under two different temperatures (10–13 °C and 21–22 °C). The development time from larval to adult eclosion do not differ between the two photoperiods, but did between the two temperature regimes. However, the larvae do not enter diapause, even under short day conditions and low temperatures. The number of adults obtained does not differ with temperature and light conditions. Field captures with pheromone traps show that Brazilian apple leafroller occurs in apple orchards throughout the year and the population densities are lower in winter. Accordingly, control measures should be taken during the off-season.     

Influence of photoperiod on low temperature acclimation for cold-hardiness in Drosophila auraria

ABSTRACT. Imagines of Drosophila auraria Peng, a reproductive diapause species, developed cold-hardiness at low temperatures to a greater extent when exposed to a diapause-inducing photoperiod (LD10:14 h) than when exposed to a diapause-preventing photoperiod (LD 16:8h). Imagines kept at 18°C, which was the temperature at which they were reared to eclosion, did not survive a test exposure to -5°C for 8 days regardless of age or photoperiod. When transferred to 10 or 5°C, either from eclosion or from 8 days after eclosion, the survival rate, on testing, rose with time since transfer and rose faster and higher with a photoperiod of LD 10:14h than with LD16:8h. Flies transferred to 15°C only showed improved ability to survive the test if they were kept in LD 10:14h. When cultured at 18°C to the age of 8 days after eclosion, diapause was terminated in about 30% of females even at LD 10:14h. In these post-diapause females the ability to develop cold-hardiness at lower temperatures was somewhat less than in the diapausing females, but apparently greater than in the non-diapause females. These results suggest that the physiological mechanism which promotes cold-hardiness under a diapause-inducing photoperiod is not directly linked to the process causing reproductive diapause.
In Sapporo, flies from a natural population became tolerant to cold in October when they entered diapause and daily mean temperature fell below 15°C and the light/dark cycle fell below LD 12:12h.