Temperature and photoperiodic control of developmental responses in climatic races of Mimulus

Two strongly differentiated climatic races of the Mimulus cardinalis-lewisiicomplex were grown at a variety of temperatures (3–27°C)and photoperiods (8 and 16 hr) under controlled environmentalconditions. M. cardinalis (the lowland race, 400 m) and M. lewisii(the sub-alpine race, 3200 m) were found to differ in theirphysiological responses to the varied environments in severalsignificant ways: 1) At 27°C (16 hrphotoperiod), M. lewisiisustained 100% mortality in contrast to the substantial growthand flowering of M. cardinalis under these conditions; 2) In8 hr photoperiods at all temperatures, there was little growthand no flowering in M. lewisii whereas there was considerablegrowth at all temperatures, and flowering at 23 and 27°Cin M. cardinalis; 3) At low temperatures (7–15°C),16 hr photoperiods, flowering occurred a week or two earlierin M. lewisii than in M. cardinalis. The lowland race has asignificantly wider temperature and photoperiodic tolerancethan has the sub-alpine race. Applications of gibberellic acidto rosette Mimulus plants under non-inductive conditions (15°C,8 hr photoperiod) promoted vigorous stem elongation withoutflowering. The application of steroids, other hormones and metaboliteshad no observable effects. ¹Present address: Faculty of Botany, Ohio State University,Columbus, Ohio, U.S.A. (Received March 16, 1970; )     

Light Quality and Vernalization Interact in Controlling Late Flowering in Arabidopsis Ecotypes and Mutants

The late flowering ecotypes of Arabidopsis thaliana L. (Heyn.)Eifel, Pitztal and Innsbruck responded to 10 d vernalization(cold treatment) by flowering earlier with less with less thanhalf the number of leaves of non-induced plants. The vernalizationresponse was cumulative: increased numbers of days of vernalizationinduced earlier flowering up to an apparent saturation in responseafter 30 to 40 d. The ratio of red:far-red (R:FR) light alsoaffected non-vernalized time-to-flower. When grown under fluorescentplus incandescent lamps (R:FR = 1·0), time-to-flowerwas approximately half that required by plants grown under fluorescentlamps (R:FR = 5·8) at the same photon flux density andphotoperiod. Leaf production rate was unaffected by either vernalizationor light quality changes and time-to-flower and leaf numberwere highly correlated (r² = 0·973). The late flowering mutants of Landsberg erecta were grown underlighting which displayed a gradient of R:FR. Some mutants likeco, flowered at the same time in all R:FR treatment, while otherlike fca took nearly twice as long to flower, with double thenumber of leaves at R:FR ratio of 5·8 compared with theR:FR = 1 treatment. The ranking of the response from least tomost responsive was co, fe, gi, WT, fd, fwa, ft, fha, fpa, fy,fve and fca. Vernalization of these Landsberg mutants always resulted inearlier flowering, although only fca, fve, fy and fpa were significantlymore sensitive to thermoinduction than the wild type parent.There was a high correlation (r² = 0·89 between the responseto thermoinduction and to R:FR ratio. Vernalization of fca for24 d largely eliminated the R:FR time-to-flower response. Vernalizationand photoinduction similarly affect late flowering and can substitutefor each another.Copyright 1993, 1999 Academic Press Light quality, vernalization, flowering, Arabidopsis thaliana, phytochrome, thermoinduction, photoperiod, photoinduction, growth conditions, photon flux density, daylength, spectral quality, far-red light     

Photoreversibility of flower initiation in Lemna perpusilla as affected by the light quality of the main light period

Reversible floral responses of Lemna perpusilla to red and far-redlights appeared only at the beginning of the inductive darkperiod when the 8 hr photoperiod consisted of white or red light.When blue or far-red light was given during the 8 hr photoperiod,the far-red given at the beginning of the dark period scarcelyinhibited flowering; red/far-red reversibility newly appearedat the middle of the dark period. This indicates that the photoregulationsystem in the flowering of L. perpusilla can be converted fromthe Pharbitis type to the Xanthium type by changing the lightquality of the main photoperiod from white or red to blue orto far-red, which is known to be effective for the so-calledhigh-energy photoreaction of photomorphogenesis. (Received July 2, 1975; )     

Growth and flowering of Lemna paucicostata I. General aspects and role of chelating agents in flowering

Lemna paucicostata HEGELM. is normally a short-day plant andflowers only in the presence of a chelating agent (EDTA or EDDHA)in the medium. The plant can be induced to flower even by asingle long night treatment; the flowering percentage, however,increases with further inductive cycles. The length of the criticaldark period depends upon the chelating agent employed in themedium. It is between 10 and 12 hr in the medium containingEDTA and about 8 hr in the EDDHA-supplemented medium. Red lightinterruption in the middle of the dark period—even fora minute—is inhibitory for flowering. Attempts to identify the metal ion(s) chelated reveal that thechelating agents affect flowering by facilitating iron uptake.This is also supported by the fact that the requirement of achelating agent for flowering can be overcome with an excessof iron in the medium. Interestingly, provision of EDDHA andexcess of ferric citrate, together, can bring about floweringeven under long days. ¹Originally HEGELMAIER (1) designated L. paucicostata as a separatespecies; however, THOMPSON (2) and DAUBS (3) have treated itsynonymous to L. perpusilla. More recently, based on physiologicaland chemotaxonomic studies, the distinctiveness of L. paucicostatafrom L. perpusilla has been favoured (4, 5). (Received September 8, 1969; )     

A STUDY ON VARIETAL DIFFERENCE IN FLOWERING HABITS OF SOYBEAN PLANTS AS FOLLOWED BY GRAFTING EXPERIMENTS

The material basis of varietal differences in flowering habitwas investigated from the standpoint that flowering is determinedby the balance in amounts of flowering promotor and inhibitor.The grafting method was used throughout the experiments. Late variety of Glycine max L. seems to produce flowering inhibitor(or inhibitors) under the conditions under which midseason varietyproduces flower. Early variety seems to produce flowering hormonewhich overcomes the flower-inhibiting action in the late variety. The amounts of flowering hormone produced under short day conditionbymidseason and late varieties were compared. The results showthat, under the short day condition, the midseason variety producesequal or smaller amounts of flowering hormone as compared withthe late variety. On the basis of these results, the mechanismby which the flowering habits are determined in various varietiesof soybean plants was discussed. (Received June 18, 1962; )     

Flowering Responses of Contrasting Ecotypes of Poa annua and their Putative Ancestors Poa infirma andPoa supina

Flowering responses of two Australian and six Norwegian populationsof Poa annua and their putative ancestors P. infirma and P.supina were studied in controlled environments. The two Australianpopulations originating from suburban parks in Canberra hadopposite daylength flowering responses across the range of temperaturestested (9–21 °C), one being a quantitative short-day(SD) plant with no response to vernalization, the other a quantitativelong-day (LD) plant with a quantitative vernalization requirement(winter annual type). Variation in earliness of flowering withinthe former population was shown to be genetically determined,and testing of selfed progenies indicated that the populationis an aggregate of several largely homozygous lines with divergentflowering responses. Two lowland populations from southern Norwaywere both quantitative LD plants with no vernalization response,while two alpine snowbed populations from southern Norway andtwo high-latitude, subarctic populations from northern Norwaywere quantitative SD plants with an obligatory plant vernalizationor SD requirement for flowering. Two populations of P. supinaexhibited the same flowering responses as the alpine and high-latitudepopulations of P. annua with an obligatory plant vernalizationor SD requirement for flowering. A combination of SD and lowtemperature (9–12 °C) for 8–10 weeks was optimalfor induction and inflorescence initiation. On the other hand,P. infirma was found to be an early-flowering quantitative SDplant which flowered freely across the range of temperatures(9–21 °C) as a typical summer annual. The experimentsdemonstrate that virtually any kind of photoperiodic and vernalizationresponses can be found among populations of P. annua. Theseversatile flowering responses reflect the contrasting floweringresponses of P. supina and P. infirma, and add strong supportto the hypothesis that P. annua has originated from these species.Copyright 2001 Annals of Botany Company Adaptation, evolution, flowering, Poa annua, P. infirma, P. supina, photoperiod, vernalization     

Flowering of S. 24 Perennial Ryegrass (Lolium perenne L.) in Short Days at Low Temperature

The flowering of the quantitative long-day plant perennial ryegrass(Lolium perenne L.) in short days (8 h), when grown at low temperature(9/4 °C) and under natural summer daylight, is presentedas evidence for the replacement of specific environmental requirementsfor flowering by alternative stimuli in a grass.     

Flowering in Pisum: the Effect of Age on the Gene Sn and the Site of Action of Gene Hr

Late cultivars of peas behave as quantitative long day plants.The reason that they flower between nodes 20 and 35 under an8 h photoperiod is shown to be because the leaves and maturestem produce a more promotory ratio of the flowering hormonesas they age. Later formed leaves may also start with a slightlymore promotory ratio than the leaves produced at a lower node.The gene Sn controls the production of a flower inhibitor andit is suggested that the activity of this gene in a leaf isgradually reduced as the leaf ages. From grafting experiments,the site of action of the gene Hr is shown to be in the leavesor mature stem and not at the shoot apex. This supports a previoussuggestion that the gene Hr is a specific inhibitor of the ageingresponse of gene Sn. Gene Hr is shown to cause a substantial delay in the floweringnode of decotyledonized plants of genotype If e sn hr undershort day conditions, suggesting that Hr has little effect inthe cotyledons. It is argued that the gene sn is a leaky mutantand that gene Hr does not control a photoperiod response inits own right but has its effect through the Sn locus. From a comparison of intact plants and self-grafts of the lategenotype If e Sn hr it is shown that under the conditions usedphysiological age may be of more importance than chronologicalage in determining flowering in peas. Reasons for the smalleffect of defoliation treatments on flowering are discussedas well as possible reasons for the promotory effect of decotyledonizationon the flowering node of late lines. Pisum sativum L, flowering, ageing, genetic control     

Modelling the Masting Behaviour of Betula platyphylla var. japonica using the Resource Budget Model

A masting model of Betula platyphylla var. japonica, a majorwind-pollinated tree in Hokkaido, northern Japan, was constructedfrom the perspective of the resource budget model. An 11-yeardata series of pollen dispersal by birch showed marked annualfluctuations in flowering. Although flowering fluctuated widelyamong years, a reciprocal relationship was observed betweentwo consecutive years; very little flowering occurred aftera mast year. This suggests that there is a negative relationshipbetween current fruiting and flower bud induction. There wasa significant positive relationship in flowering between regions,which suggests that weather conditions regulate flowering inbirch. A model that included weather variables and resourceallocation systems among years explained 94·5% of theobserved annual fluctuations in flowering. In the model, warmspring conditions from bud burst to flower bud development andlittle flowering in the previous year (t-1) resulted in increasedflowering in the current year (t), whereas abundant floweringin year t-2 resulted in a decrease in flowering in the currentyear (t). The latter indicates that flowering in year t-2 affectsresource storage after overwintering; the balance is carriedforward to year t-1 and affects the initiation of flower primordiabefore bud burst. In the model, fluctuating weather conditionsin the previous spring strongly affected the masting behaviourof birch. Copyright 2001 Annals of Botany Company Betula platyphylla var. japonica, flowering, masting, modelling, pollen dispersal, resource allocation, resource budget model, weather conditions     

Effects of Exogenous Amino Acids on Iron Uptake in Relation to their Effects on Photoperiodic Flowering in Lemna paucicostata 6746

The flowering of Lemna paucicostata 6746 grown on 14-h photoperiodwas enhanced by the addition of high concentrations of ironto the medium, which also increased the endogenous iron concentration.The addition of asparagine, aspartate, glutamate, -alanine,glycine or serine to the medium also increased the endogenousiron level, resulting in the promotion of flowering. In contrast,the addition of cysteine, cystine, glutamine, arginine, threonineor phenylalanine lowered the endogenous iron level, resultingin the inhibition of flowering. Glycine and asparagine added to the medium during an inductive96-h dark period did not promote iron uptake and had no effecton flowering, but when added during the subsequent 120-h lightperiod, they promoted both iron uptake and flowering response.The increase in the endogenous iron level seems to favor floraldevelopment rather than induction of photoperiodic floweringof Lemna paucicostata 6746. (Received September 8, 1986; Accepted March 31, 1987)     

Isolation and Identification of L-Pipecolic Acid and Nicotinamide as Flower-Inducing Substances in Lemna

Flower-inducing factors in extracts of flowering Lemna gibbaG3 were investigated using Lemna paucicostata 151 as the bioassayplant. Fractions with flower-inducing activity were obtainedafter several purification steps. Two of the active substanceswere identified as L-pipecolic acid and nicotinamide by MS andNMR analyses. Both L-pipecolic acid and nicotinamide exhibited flower-inducingactivity in L. paucicostata 151 grown on one-tenth-strengthM medium containing benzyladenine, the former being ten timesas active as the latter. L-Pipecolic acid was active even at0.01 ppm (7.8 ? 10^–8 M). The effect of L-pipecolic acidon flowering strongly depended upon the presence of exogenouscytokinin. The coexistence of cytokinin seemed to be essentialfor L-pipecolic acid to exhibit flower-inducing activity. Incontrast, the effect of nicotinamide on flowering was basicallythe same as that of benzoic acid or nicotinic acid. (Received February 9, 1987; Accepted May 21, 1987)     

Cell Cycle Responses to Red or Far-red Light, or Darkness, in the Shoot Apex of Silene coeli-rosa L. During Floral Induction

Twenty-eight-day-old plants of Silene coeli-rosa L. were maintainedin short days (SD) for 9 d (0–8) or exposed to 7 longdays (LD), or 7 SD with a 5 min exposure at 1700 h of each dayto far-red (FR), red (R) or 5 min FR/5 min R, or 7 dark-interrupted(di = 1700–1720 h) LD. Treatments were followed by twofurther SD. The mitotic index and G1 and G2 proportions weremeasured in the shoot apices of plants sampled at 2000 h ofeach day of each replicated treatment. Exposure to 7 LD (= 100per cent flowering) resulted in significant increases, relativeto the SD controls, in both the G2 proportion and the mitoticindex on d 0 to 3, 7 and 8. Five minute FR (= 0 per cent flowering)resulted in cell cycle responses similar to those in LD onlyfrom d 0 to 2. R and FR/R (both = 0 per cent flowering) didnot result in any increases in the G2 proportion in the apexapart from d 3 of FR/R. However 5 min FR/5 min R, and to a lesserextent 5 min R, did result in significant increases in the mitoticindex on d 0, 1, 7, and 8. diLD (= 8–10 per cent flowering)also prevented any significant increases in the G2 proportionon d 0 to 3, and 5 to 8 but the mitotic index was again higheron these days compared with control data. Thus the transitionto floral growth for 90 per cent of the plants is associatedwith changes in the cell cycle in the shoot apex measured asincreases in the G2 proportion at 2000 h of LD 0 to 3 and 7to 8. Silene coeli-rosa L., cell cycle, flowering, phytochrome, shoot apex     

Relationships Between Climate and Flowering of Eight Herbs in a Swedish Deciduous Forest

Effects of annual variation in rainfall, temperature and humidityon flowering abundance of eight temperate woodland plants (Anemonenemorosa, Cardamine bulbifera, Lamiastrum galeobdolon,Oxalisacetosella , Ranunculus ficaria, Stellaria holostea, Viola reichenbachianaand Viola riviniana) were studied during 12 consecutive years(1989–2000) in a hornbeam (Carpinus betulus) forest insoutheast Sweden. Above-average rainfall/humidity in late summerto early autumn of the preceding year increased flowering abundancein L. galeobdolon, O. acetosella, V. reichenbachiana, V. rivinianaand, especially, in R. ficaria, but not in S. holostea and A.nemorosa. Moreover, flowering of R. ficaria and O. acetosellawas positively related to rainfall/humidity during several partsof, or the entire, preceding year. On the contrary, floweringof S. holostea and A. nemorosa was closely related to low valuesof rainfall/humidity in autumn and/or winter of the precedingyear and also to low humidity in the current year in A. nemorosa.Two long periods (3–4 years) of increasing rainfall deficitcoincided with decreasing flowering abundance in most of thespecies, but not with decreasing vegetative development. Temperaturevariability was less consistently related to flowering. A coolperiod during the preceding summer or autumn seemed importantfor flowering in L. galeobdolon, O. acetosella and the Violaspecies, although these relations were, at least partly, causedby interactions with rainfall/humidity. No significant (P <0.05) correlations were found between flowering and the conditionsprevailing in April to May—the main flowering season—ofthe current year. Copyright 2001 Annals of Botany Company Climate, flowering, rainfall, temperature, Anemone nemorosa, Cardamine bulbifera, Lamiastrum galeobdolon, Oxalis acetosella, Ranunculus ficaria, Stellaria holostea, Viola reichenbachiana, Viola riviniana     

Effect of 8-Hydroxyquinoline on Flowering and Endogenous Levels of Iron and Copper in Lemna paucicostata, Strain LP6

Lemna paucicostata LP₆, a strain of duckweed isolated locally,does not flower under any photoperiodic schedule when grownin Bonner and Devirian or other media routinely employed invarious laboratories for studies on flowering in Lemnaceae.Flowering in this strain could be induced, however, by 8-hydroxyquinoline(8-HQ)—a well-known copper chelating agent—irrespectiveof the length of the photoperiod. To our knowledge, this isthe first report where a direct induction of flowering in aduckweed by 8-HQ has been observed. Atomic absorption analysisof the plant material revealed that the endogenous level ofcopper is significantly higher in the plants treated with 8-HQ.This is contrary to the general assumption that chelating agentsinfluence flowering of duckweeds by causing a reduction in theuptake of copper ions and making them less available to theplants. (Received May 23, 1983; Accepted July 22, 1983)     

Flower Formation from Citrus unshiu Buds Cultured In Vitro

Flowers were formed in vitro when buds of Satsuma mandarin (Citrusunshiu Mare) from the summer flush of growth harvested duringthe winter rest period before the onset of flower initiation,were grown on a solid Murashige and Skoog medium supplementedwith sucrose and a cytokinin. Flower development was dependentupon illumination, and was enhanced when a piece of stem wasattached to the bud. The percentage of flowering explants wasalways lower than the percentage of naturally flowering budsin spring, but treatments such as ringing which increase floweringin vitro, increased the number of explants flowering int vitroas well. Citrus unshiu Marc, ‘Satsuma’ mandarin, in vitro flowering, ringing     

Obligate Requirement of Salicylic Acid for Short-Day Induction of Flowering in a New Duckweed, Wolffiella hyalina 7378

Wolffiella hyalina 7378, a member of the Lemnaceae, does notflower in the basal media of Bonner-Devirian, Hoagland, Hutner(1/2 strength), M medium of Hillman or Pirson-Seidel under eitherlong or short day photoperiodic regimes. Well-known chelatingagents, such as EDTA, EDDHA and 8-hydroxyquinoline, which havebeen shown to significantly influence flowering in other membersof the Lemnaceae, also do not show any effect in this strain.It is, however, noteworthy that W. hyalina 7378 flowers profuselyunder a short-day photoperiodic schedule of 8 h light and 16h darkness provided that 10^–5 M salicylic acid is supplementedto modified Bonner and Devirian medium. The critical daylengthof W. hyalina 7378 is 13 h. Further, at least two inductivephotocycles are required for initiation of flowering. As tothe mechanism of SA action, it appears that SA effect on floweringis hormonal in nature rather than due to chelation of metalions. (Received August 25, 1986; Accepted January 9, 1987)     

Developmental and photobiological factors affecting photoperiodic induction in Arabidopsis thaliana Heynh. Landsberg erecta

We have tested whether the promotion of flowering by long days(LD) in Arabidopsis thaliana is a consequence of photoperiodicinduction. To achieve this, the flowering responses of Arabidopsisthaliana (L.) Heynh. Landsberg erecta (Ler) and the long-hypocotylmutants hy2, hy3 and hy4 were determined with respect to age,daylength and light quality. Ler was capable of distinguishingbetween short days (SD) and long days (LD) from about 4 d aftersowing at 20 C, the time at which cotyledons were expandingand greening. At this stage, the critical daylength was between8 h and 10 h. At 7 d, seedlings required five LD for inductionand, as the seedlings aged, they became more sensitive so thatby day 20, one LD was fully inductive. The response to SD innewly germinated seedlings was to delay flowering without alteringleaf number, but after about 10 d, delay of flowering by SDwas accompanied by extra leaves. In light quality experiments,blue light (B) was inductive for 5-d-old plants and in all subsequenttreatments, far-red (FR) caused induction in treatments at 12d and 18 d and low pressure sodium, equivalent to red, was notinductive at 5 d and 12 d, but partially inductive at day 18.Hence, both a specific blue-light photoreceptor and phytochromeA in High Irradiance Response mode promote floral induction.In daylength transfer experiments all three hy mutants respondedto LD by earlier flowering. Both hy2 and hy3 produced substantiallyfewer leaves than Ler in SD and hy3 flowered slightly earlierthan Ler. The hy4 mutants flowered later than Ler in SD andhad a higher leaf number. A scheme is proposed in which photoperiodicinduction depends on the ability of the plant to sense photoperiod,the stage of development and the photobiological input. We alsopropose that phytochrome A and the blue photoreceptor promoteflowering whereas phytochrome B promotes vegetative development. Key words: Arabidopsis thaliana, blue-absorbing photoreceptor, flowering, photoperiodic induction, phytochrome     

Developmental Responses to Temperature and Photoperiod in Ecotypes of Medicago polymorpha L. Collected Along an Environmental Gradient in Central Chile

The phenological development of nine Chilean accessions of Medicagopolymorpha, collected along a north–south aridity gradient,and of two commercial cultivars of the same species, were comparedin 12 sequential outdoor sowings at Cauquenes (35°58'S,72°17'W, elev. 177 m), in the sub-humid Mediterranean climatezone of Chile. A glasshouse experiment was also conducted toevaluate the effect of photoperiod on phenophase timing. Therewas a clear gradient in precocity among the Chilean accessionsin both experiments: accessions MPO-9-88 and MPO-7-88, fromthe arid zone, were the earliest-flowering accessions, whereasMPO-36-88 from the humid Mediterranean zone was the latest.Both experiments revealed significant variation among the Chileanaccessions in the response of flowering time to variation inphotoperiod regime. Differences in days to flowering betweenthe least- (8 h) and the most- (16 h) inductive photoperiodswere lower in precocious accessions from arid and semi-aridzones, than in late-flowering accessions from more humid zones.Rate of progress to flowering, defined as the inverse of timefrom emergence to first flower appearance (1/ f), was relatedto mean diurnal temperature, or to both mean diurnal temperatureand mean photoperiod. In two early-flowering accessions fromthe arid zone, and in the Australian cultivar ‘CircleValley’, 1/ f was affected significantly (P < 0.05)by both temperature and photoperiod. In the remaining accessions,no significant responses to temperatures were detected; 1/ fwas influenced significantly by photoperiod only. Copyright2000 Annals of Botany Company Annual medic, aridity gradient, Medicago polymorpha, flowering time, rate of development     

Effects of Interactions between Low-temperature Treatments, Gibberellin (GA3) and Photoperiod on Flowering and Stem Height of Spring Rape (Brassica napus var. annua)

Exogenous gibberellin A₃(GA₃) reduced the number of leaf nodesat flowering and time to flowering and increased the stem heightat flowering in three genotypes of spring rape (Brassica napusvar.annua L.). The responses to GA₃were similar to those forlong days (LD) and low-temperature treatments, suggesting thatthe effect of photoperiod and the vernalization response areprobably mediated through gibberellins. The response to exogenousGA₃was greatest in non-cold-treated plants in short days (SD)suggesting that endogenous GAs are limiting in these conditions.CCC, an inhibitor of gibberellin biosynthesis, caused a smallincrease in the number of leaf nodes at flowering and time toflowering and a small decrease in the stem height at flowering,but unexpectedly, its effect was hardly influenced by the applicationof exogenous GA₃. Genotypes that showed the clearest responsesto the treatments with regard to the number of leaf nodes atflowering and time to flowering did not show the clearest responseswith regard to the stem height at flowering; the pattern ofresponses of the number of leaf nodes at flowering and timeto flowering was distinct from that of stem height at flowering.This indicates that flower formation and stem elongation areseparable developmental processes which may be controlled bydifferent endogenous gibberellins, different levels of a specificendogenous gibberellin, or different responses to gibberellin.Copyright 1999 Annals of Botany Company Brassica napus var. annua, gibberellin, photoperiod, spring rape, vernalization.