Long-day control of flowering in everbearing strawberries

Summary

Photoperiod and temperature control of flowering in a number of perpetual-flowering or everbearing strawberry cultivars of widely varying pedigree has been studied in controlled environments. Flower bud initiation in the cultivars ‘Flamenco’, ‘Ridder’, ‘Rita’ and ‘Rondo’ was significantly advanced by long-day (LD) conditions at temperatures of 15°C and 21ºC; while, at 27ºC, flowering took place under LD conditions only. Some plants of the seed-propagated F₁-hybrid ‘Elan’, raised at 21°C, also flowered under short-day (SD) conditions at 27°C, but reverted to the vegetative state after a few weeks when maintained under these conditions. When vegetative plants growing in SD at 27°C were transferred to LD conditions at the same temperature, they consistently initiated flower buds and started flowering after about 4 weeks. At such a high temperature, flowering could thus be turned on and off by switching between SD and LD conditions. This applied to all the cultivars studied. Also the cultivar ‘Everest’, which was tested only at 21°C, produced similar results. Night interruption for 2 h was effective in bringing about the LD response. At 9°C, flowering was substantially delayed, especially in ‘Flamenco’ and, at this temperature, flowering was unaffected by photoperiod. Runner formation was generally promoted by high temperature and SD conditions, but the photoperiodic effect varied between experiments. We conclude that everbearing strawberry cultivars, in general, whether of the older European-type or the modern Californian-type originating from crosses with selections of Fragaria virginiana ssp. glauca, are qualitative (obligatory) LD plants at high temperature (27°C), and quantitative LD plants at intermediate temperatures. Only at temperatures below 10°C are these cultivars day-neutral.     

Quantitative long-day flowering response in the perpetual-flowering F1 strawberry cultivar Elan

Summary

Perpetual-flowering strawberry cultivars are commonly classified as photoperiodically day-neutral, even though early investigations demonstrated long-day (LD) regulation. An important reason for this inconsistency is that these freely flowering plants are difficult to establish in a true vegetative state, and experiments have therefore often been started using runner plants with pre-formed inflorescences. In order to circumvent this problem, we have used the perpetual-flowering F₁-hybrid ‘Elan’ that is propagated by seed, and is thus not pre-conditioned by its earlier life history. The results demonstrated a marked quantitative LD response across a range of temperatures from 9° – 27°C. Seedlings were responsive to the LD stimulus at an early stage, and early flowering required LD exposure almost from germination. The critical daylength for the early flowering response was about 15 h at 18°C. Because of this threshold LD response, it is concluded that regulation of flowering is truly photoperiodic in nature, and not merely an effect of additional light. Flower development was also slightly advanced by LD conditions. Stolon formation was strongly enhanced by short-day conditions in combination with high temperature. Thus, not only flowering, but also runnering, was oppositely affected by photoperiod in ‘Elan’ compared with mono-flowering cultivars. The results are discussed in relation to the photoperiodic classification of strawberries.     

Bulbing responses of two cultivars of red tropical onions to photoperiod,light integral and temperature under controlled growth conditions

Summary

The effects of photoperiod, light intensity and temperature on bulb formation and bulb structure of two tropical onion cultivars were investigated. From an initial experiment it was observed that the number of true scales and sheath scales differed significantly between the cultivars ‘Red Creole’ and ‘Agrifound Dark Red’. When these two cultivars were given 11, 12 and 13 h photoperiod treatments, it was found that both cultivars needed at least 12 h photoperiod for bulb formation. Modify the R/FR ratio from 1.22 to 1.16 in the final hour of the 11 h light period did not induce bulbing. The 13 h photoperiod increased the number of true scales and decreased the number of sheath scales compared with the 12 h photoperiod in both cultivars but total scale + leaf sheath numbers remained nearly constant. When onion plants were grown under 0%, 25%, 50% and 75% shading treatments, (12 h photoperiod), only plants receiving 0% and 25% shading bulbed. Low light intensity decreased the number of true scales and increased the number of sheath scales. Four temperature regimes were compared in a growth room experiment. Plants under the 29°348C treatment bulbed within two weeks and matured within six weeks. However, plants receiving the 25°308C treatment delayed bulb initiation more than those plants receiving 17°228C and 21°268C treatments. At the lowest temperature, bigger bulbs with thick necks were produced. This may be due to changes in bulb structure since at low temperature, the number of sheath scales was increased, however the number of true scales remained relatively constant in both cultivars. Dormant leaf initials decreased with decreasing temperature while the number of secondary meristems significantly increased. The results suggest that ‘Red Creole’ was more responsive to shorter photoperiods, bulbing earlier than ‘Agrifound Dark Red’. There was no significant difference in time to bulbing in response to temperature between the two cultivars if measured by bulbing ratio however there were differences in bulb structure which suggested that ‘Red Creole’ bulbed earlier. These effects may be due to the breeding histories of the two cultivars. It is suggested that studying bulb structure may provide a useful method of interpreting onion bulbing responses.     

Effect of temperature on flowering in Primula vulgaris

When grown in a glasshouse, flowering in Primula vulgaris ‘Aalsmeer Giant’ (yellow) and ‘Ducat’ (blue) was delayed with increasing temperature from approximately 12°C to 18°C. In addition, size of the first open flower and the number of flowering axillary shoots decreased, whereas the number of leaves and leaf area increased with the temperature increase. All temperature responses were greater in ‘Aalsmeer’ than in ‘Ducat’.When grown in growth rooms at 9°C, flowering in P. vulgaris ‘Aalsmeer Giant’ (yellow) was inhibited compared with 15°C. However, when 9 weeks of 15°C was applied to plants grown for 9 weeks at 9°C, the inhibition was overcome; longer periods of 15°C being no more effective. This indicates than an early stage of flower formation, probably the initiation, in Primula vulgaris is inhibited by 9°C, and not the further development of the buds towards open flowers.     

Effect of photoperiod and light integral on flowering and growth of Eustoma grandiflorum (Raf.) Shinn.

The aim of the study was to examine the effects of different photoperiod and light integral on floral initiation, development and subsequent growth of Eustoma grandiflorum (Raf.) Shinn. Six-weeks-old seedlings of ‘Echo Blue’ and ‘Fuji Deep Blue’ were placed under short day (SD, 10 h) and were transferred to long days (LD, 20 h) at 2-week intervals from 6 to 14 weeks after seeding. Plants initiated flower buds regardless of light regimes. Flower bud initiation was delayed by SD compared to LD; plants transferred after 6 weeks from seeding initiated flower buds at least 21 and 10 days earlier at LD at high (HL) and low (LL) daily light integral, respectively, compared to those at SD. Light regimes had little or no effect on time to flower bud development after initiation. Thus, it seems likely that LD and HL affected the initiation rather than development. Both the photoperiod and light integral strongly influenced the subsequent growth after initiation. SD delayed the time to visible bud (VB), increased the number of nodes to first open flower, number of branches, stem diameter and shoot dry weight compared to LD. HL promoted flowering and increased several shoot characteristics and flowering compared to LL.The results indicate that Eustoma is a quantitative long-day plant. LD, and more specifically HL, enhanced flower bud initiation, development and subsequent growth. An initial SD period is preferred to increase the number of branches, number of flowering buds and flowers, stem diameter and shoot dry weight.     

Flowering of Primula malacoides in response to photoperiod and temperature

Primula malacoides Franch. ‘Prima Lilac’ was grown at 16 or 20 °C in combination with short days (SD, 8 h) or long days (LD, 16 h). In addition to uninterrupted growing conditions, plants within each temperature were moved at weekly intervals to the other photoperiod and left until termination. Temperature, but not photoperiod, significantly affected the rate of development from start of treatments (51 days from seeding) to 2 mm visible flower bud (VB). At 16 °C, VB averaged 30 days and at 20 °C, 48 days. Time to flower (first horizontal petals) at 16 °C increased from 56 to 64 days as SD increased from 1 week to continuous conditions while LD decreased time to flower from 64 to 56 days. Time to flower at 20 °C varied from 73 to 87 days with additional SD exposure resulting in slower and LD in faster flowering. These observations of the flowering response in ‘Prima’ are contrary to the photoperiodic classification of P. malacoides as a SD plant.     

Short days and temperature effects on growth and flowering in strawberry (Fragaria × ananassa Duch.)

Summary

‘Korona’, ‘Elsanta’, ‘Bounty’ and ‘Senga Sengana’ strawberry (Fragaria × ananassa Duch.) plants, were placed at constant temperatures of 9, 15 or 21°C and daylengths of 8 h (short day) or 24 h (long day). The plants were given different numbers of short-day (SD) cycles, and flowering and growth were studied. ‘Korona’ and ‘Elsanta’ were responsive to both short-day treatment and temperature, with optimum flowering at 15°C and 24 SD. ‘Bounty’ was more responsive to temperature, inducing flowers independently of the number of SD cycles at 9°C and 15°C. In ‘Senga Sengana’ flowering was induced independently of temperature and the number of SD cycles, indicating that it had a stronger dependence on other environmental effects. The effect of the number of short-day cycles and the temperature on vegetative growth variâtes such as the number of stolons and daughter plants, the length of flower trusses and petiole length were also studied.     

Flower formation of Chinese cabbage. I. Response to vernalization and photoperiods

The influence of temperature, vernalization duration and photoperiods on the leaf number and beginning of bolting of Chinese cabbage was tested in growth chambers. An increasing effect of vernalization was indicated by decreasing leaf number and premature bolting. Sensitiveness for vernalization started with germination and remained constant with increasing plant age. During their development on mother plants, seeds were not vernalized.Inductive temperature ranged between 0 and 20°C. The necessary vernalization duration was lowest at 5–8°C. For slight vernalization effects one week was sufficient; for complete vernalization about 3 weeks were needed. To achieve the same effect, the necessary vernalization duration increased considerably at temperatures below 5°C, but increased only slightly at temperatures above 8°C.Long days promoted bolting when the temperature rose after incomplete vernalization. In constant low or high temperature, no photoperiodic effect was perceptible.     

Inhibition of flower stalk elongation and abnormal flower development by short-day treatment in a Japanese variety of Chinese chive (Allium tuberosum Rottler ex Sprengel)

Chinese chive is produced year-round in Japan, therefore it is important to demonstrate the relationships of short-day (SD) conditions with flower stalk elongation and flower formation for optimizing the cropping system, e.g. the increase in harvests per year. To clarify the influence of short photoperiod on flower stalk elongation and flower formation in Chinese chive, 8 h SD treatments were applied at different developmental stages of flower stalk elongation and flower formation. When the SD treatment started from vegetative or floral-initiated stages, the earlier the SD treatment started, the fewer flower stalks appeared. Also, the earlier the SD treatment started, the less the flowers bloomed and the more the flower stalk elongation was inhibited at the end of SD treatments. Many involucres did not open and withered with death of florets in SD when the SD treatment started between the umbel or flower bud differentiation and the perianth to stamen-formation stages. Also, all or part of the florets aborted and there were no complete inflorescences in the later SD treatments. We found that, in Chinese chive, the development of flower stalk elongation and flower formation were inhibited with the earlier SD treatment, after vegetative or floral-initiated stages. Furthermore, it is considered that Chinese chive needs long-day (LD) for the flower stalk elongation and inflorescence formation after the initiation of the flower bud. The plant has a qualitative LD requirement with the same photoperiodic requirement for both flower bud initiation and flower development.     

The responses of early-flowering chrysanthemums to daylength

In a study of photoperiod response 30 early (summer)-flowering chrysanthemum cultivars from Britain and Japan reacted as quantitative short-day plants. Both flower bud initiation and development occurred more rapidly in short days (SD) as compared with long days (LD). The early-flowering cultivars differed from late-flowering chrysanthemums in that lateral flower buds, and often the terminal flower buds, develop to anthesis in LD. Internodes were shorter in SD and flowers had fewer florets. There was an increased number and proportion of disc florets in SD in all cultivars except one. Two cultivars, ‘Mezame’ and ‘Pennine Yellow’, closely approached day-neutrality.All cultivars produced fewer leaves in LD on upper lateral shoots than on main stems. Either the presence of the terminal flower bud induces earlier flowering of the upper lateral shoots, or the change which causes the terminal apical meristem to initiate a flower in LD influences the axillary meristem also.SD could be used to speed up the flowering of glasshouse-grown crops of early-flowering cultivars. The advantages and disadvantages which may be associated with treatment at different stages of growth are discussed.     

The temperature and photoperiod regulation of flowering and runnering in the strawberries,Fragaria chiloensis,F. virginiana,and F. x ananassa

The temperature and photoperiod interactions of a number of elite genotypes of Fragaria virginiana, F. x ananassa, and F. chiloensis were studied in a series of growth chamber experiments. Several parameters were evaluated including: (1) the critical day-length (CDL) for flowering of short day (SD) genotypes under 8, 9, 10, and 11 h days at 18 °C, (2) the floral and runnering response of single and multiple cropping genotypes under 8 and 16 h days at 18 °C, and (3) the effect of temperature on flower bud formation in day-neutral (DN) genotypes held at 18, 22, 26, and 30 °C under 12 h day-lengths. The same number of flowers were initiated under 15 and 30 day induction periods, regardless of photoperiod. Frederick 9, LH 50-4 and RH 30 (F. virginiana), ‘Aromas’ and ‘Tribute’ (F. x ananassa) and CFRA 0368 of F. chiloensis flowered under both long days (LDs) and SDs; while Eagle-14 (F. virginiana), ‘Fort Laramie’ and ‘Quinalt’ (F. x ananassa) flowered only under long days. While those genotypes that flowered under both LD and SD can be considered day-neutral, they varied in the degree of floral response to the two photoperiods. CFRA 0368 and Frederick 9 produced the same number of flowers under both LDs and SDs, while ‘Aromas’ and ‘Tribute’ had more flowers under LDs and RH 30 had more under SDs. Of the DN genotypes, LH 50-4 and RH 30 were the only ones that produced runners under SDs. Flowering in ‘Fort Laramie’ was least affected of any genotype by high temperature, although its dry weight was negatively impacted. Based on these data, several genotypes show promise as breeding parents: CFRA 0368 and Frederick 9 to equalize flower production under LD and SD conditions, LH 50-4 and RH 30 to produce more freely runnering DNs, and ‘Fort Laramie’ for floral heat tolerance.     

两种春化方式对不同萝卜品种抽薹开花的影响

以4种不同生态类型的萝卜品种为试材,研究了种子春化及种子春化结合幼苗春化两种处理方式对萝卜抽墓开花的影响,以探讨在北京地区如何实现不同萝卜品种冬季温室加代繁殖的目的.实验结果表明:对于不耐抽薹即春性强的的萝卜品种,单一的种子春化处理即可实现加代目的;但对于耐抽薹即春性弱的萝卜品种,必须将种子春化与幼苗春化结合起来,才能实现加代目的.     

Flower formation of Chinese cabbage. II. Anti-vernalization and short-day treatment

The influence of high temperature (anti-vernalization) and short-day treatment during the raising of transplants on the bolting of Chinese cabbage having early planting-dates has been tested. A temperature rise from 18 to 26°C resulted in a delay in bolting, but only caused a slight increase in leaf number. Hence, Chinese cabbage shows anti-vernalization effects. The raising of transplants below 20°C and with short-day treatment for energy saving cannot be recommended, since the effect of the short-day treatment is too small.     

Inhibition of flowering of Mexican- and Guatemalan-type avocados under tropical conditions

Avocado trees of a range of cultivars growing in Darwin, northern Australia (average yearly maximum 33°C, minimum 23°C), were observed for flower and shoot development. Terminal buds of the cultivars ‘Fuerte’, ‘Rincon’ and ‘Edranol’ sampled in July were not floral. Buds which did not burst were sampled in September and they contained developing flowers with perianth primordia. Vegetative extension growth resulted from laterals proximal to the inhibited terminal buds.Avocado trees of the cultivars ‘Fuerte’ and ‘Hass’ which had initiated floral buds were transferred to controlled environment chambers with 33°C day, 23°C night ($\text{33}\text{23}$) or 25°C day, 15°C night ($\text{25}\text{15}$) with a 12-h photoperiod and photon flux density of 400 μmol m^?2 s^?1 (400–700 nm). At $\text{33}\text{23}$ the trees had fewer flowers and a shorter flowering period than at $\text{25}\text{15}$. Inhibited floral buds and lateral vegetative extension resulted at $\text{33}\text{23}$, as observed in northern Australia. The unburst buds had developing flowers with perianth and stamen primordia.The controlled environment experiments showed that the abnormal flushing behaviour of Mexican- and Guatemalan-type avocados growing in northern Australia was due to high temperature. Floral development was inhibited at the stage of stamen differentiation.     

Flowering in pyrethrum (Tanacetum cinerariaefolium L.). I. Environmental requirements

SUMMARY

Floral evocation in pyrethrum {Tanacetum cinerariaefolium) is stimulated by a period of vernalization. Night temperatures of 6°C and 12°C for two weeks and three weeks, respectively, promote rapid inflorescence initiation and development. A night temperature of 18°C does not satisfy the vernalization requirement. Longer periods of vernalization stimulate more rapid inflorescence initiation and development and result in a larger number of inflorescences being initiated. The vernalization stimulus may be modified by the daily light integral. Both inflorescence initiation and inflorescence development are promoted by long days. The number of leaves formed before flower bud initiation is not affected by daylength. Night-break lighting does not promote flowering. It is suggested that the stimulatory effect of long days may be in supplying photosynthetic assimilates to the developing meristems. Floral development is retarded by low photon flux density conditions regardless of day temperature. High day temperatures (25°C) combined with low photon flux (350 umol m"2 s"' or less) prevented pyrethrum from flowering in otherwise inductive conditions.     

Effect of photoperiod on flower bud initiation and development in myoga (Zingiber mioga Roscoe)

Selection of suitable production locations in Australia and New Zealand for production of myoga (Zingiber mioga Roscoe) has been limited by lack of information on climatic influences on flowering. This study focused on photoperiod as potential production sites within Australia differ considerably in daylength due to the geographical range. The two cultivars available in Australia (Inferior and Superior) were examined in this trial due to previously observed differences in vegetative and reproductive development.Plants grown under long-day conditions (16 h) and short-day conditions (8 h) with a night break produced flower buds, while those under short-day conditions (8 h) did not. The failure of plants under short-day conditions to produce flower buds was due to abortion of developing floral primordia rather than a failure to initiate inflorescences. It was concluded that for flower development in myoga a qualitative long-day requirement must be satisfied, but that flower initiation was day-neutral.Short-day conditions resulted in abortion of flower primordia, premature senescence of foliage and reduced foliage dry weight in both cultivars. Early senescence and low flower bud yield of the Inferior cultivar, but not the Superior cultivar have been observed in crop evaluation trials in Southern Australia and New Zealand. Differences in critical photoperiod between the two cultivars may explain this observation and therefore photoperiodic requirements may be an important consideration in site and planting date selection for different cultivars.     

Effects of vernalization and gibberellic acid on flower bud formation in different genotypes of pea under different photoperiods

The influences of vernalization, photoperiods of 8, 12 or 16 hours, and gibberellic acid (GA) on the flower bud formation of the pea were studied in a Late, a Medium and an Early flowering genotype, each genotypically either Tall or Dwarf, but for the rest genetically identical. The detailed experimental results are summarized on p. 188 (Section 3.4).Just as Medium and Early, Late is considered as a quantitative long-day plant, but its necessary duration of exposure is exceedingly long under short-day conditions. It is suggested to replace ‘critical day length’ by ‘critical duration of exposure’ in the pea and similarly reacting plants. Flower bud formation becomes increasingly more difficult from Early to Medium to Late, from Tall to Dwarf, from photoperiods of 16 hours to 12 hours to 8 hours, and certain combinations become marginal.Vernalization and GA had dramatic effects under the same marginal conditions, but GA was sometimes effective, where vernalization was not. In case of effect of both factors, strong interactions were found: vernalization reduced the effect of GA, and the reverse. This suggests similarity of action.The implications of these results for the mechanism of flower bud formation were discussed. It is presumed that the triple allelic genes for Late, Medium and Early determine a strong, an intermediate and a weak inhibition of flower bud formation, respectively. Inducing factors remove these inhibitions. Long day induces so strongly that vernalization and GA have hardly any supplementary effects, but under decreasing photoperiods, or more general, under marginal conditions, vernalization and gibberellic acid become more and more important as inducing factors.     

Juvenile Phase and Flower Initiation in Brilliant Stocks (Matthiola Inc Ana R.Br.)

The effect of temperature and day-length on growth and flowering of the Scandinavian Brilliant-type stock (Mattkiola incana R. Br.) has been studied. Flower differentiation and development were examined.

Flower initiation was favoured by low temperature and long days, but a temperature as high as 20° C. (68 °F.) did not prevent flowering even under short-day (9 hours) conditions. A close interaction between temperature and day-length was observed.

High temperature was favourable for further development of the flowers, but abnormal flowers were developed if the plants were removed to high temperature before the first flower primordium was visible under the microscope.

Flower initiation was associated with accelerated stem elongation. Therefore, optimal temperature conditions for stem elongation changed with the developmental stages of the plants.

The plants responded to a low-temperature treatment as early as 12 days after germination, at which stage they had only two small leaves in addition'to the cotyledons. The Brilliant stocks have evidently a much shorter juvenile phase than that previously described for the Column type. This might be the main reason why the Brilliant stocks flower very early.

In single-flowered plants the differentiation of floral organs took place in the following sequence: sepals, stamens, carpels, petals. In this respect the stock showed a similar behaviour to that previously described for certain other cruciferous genera.

The results of the experiments are discussed in the light of previous studies concerning low-temperature effects, vernalization and related phenomena, on flowering. It is concluded that no true vernalization process is involved. The facts that the initiation took place immediately after exposure to low temperature, and that removal to high temperature caused abnormal flower development unless differentiation had reached a certain stage, form the main basis for this conclusion.     

Promotion of floral initiation in ‘Fuerte’ avocado by low temperature and short daylength

Potted avocado (Persea americana Mill., cv. ‘Fuerte’) plants were maintained in growth cabinets for up to 32 weeks and new growth observed for flower formation. Flowers were formed if temperatures were 20°C or below, but with 25° or 30°, even if only for 1 hour per day, flower formation was inhibited. Time to flowering was accelerated, but number of flowers reduced, if daylength was shortened from 15 h to 9 h. With low temperature and short days, full bloom was about 4 months after starting experiments. Spring flowering of cv. ‘Fuerte’ in the field could follow flower induction about 4 months previously with the onset of winter temperatures and daylengths.