Photoperiodic studies on growth and development of Impatiens balsamina L.

Summary In the short-day plant Impatiens balsamina it was found that, while floral buds are initiated with 3 short-day (SD) cycles, at least 8 such cycles are required for flowering. The numbers of floral buds and open flowers bear a linear relationship with the number of SD cycles. The induced floral buds revert to vegetative growth unless the plants receive the minimum number of SD cycles needed for flowering, this reversion occurring in a basipetal direction. The rate of extension growth of the stem increases with increasing numbers of SD cycles. The high rate is maintained longer in plants receiving 32 or more SD cycles, but the subsequent fall is also steeper in these plants than in plants receiving less inductive cycles. Senescence also occurs in these plants and appears to be related to the magnitude of reproductive development and the high rate of extension growth.     

Significance of timing and number of short-day cycles for initiation and subsequent development of floral buds in Lemna perpusilla 6746

Time courses of the flowering process in Lemna perpusilla 6746,a short-day plant, were studied using selected fronds in relationto the order of emergence. Various numbers of short-day cycleswere interposed during continuous light. The floral buds evokedby short-day cycles developed to a floral stage determined bythe number of short-day cycles 3 days after the transfer toconsecutive long-day cycles, but aborted on the next day, regardlessof the floral stages. At least 2 long-day cycles were requiredfor the abortion of the floral buds at any stage of development.These results suggest the importance of the number of short-daycycles not only for initiation but also for development of floralbuds. (Received February 4, 1977; )     

Photoperiod and the Determination of Potential Seed Number in Chenopodium rubrum L.

Individuals from two latitudinal populations of Chenopodiumrubrum, a short-day annual, were induced in two inductive photoperiods,15 h and 12 h, to examine the dynamics of reproductive developmentthat determine the potential number of seeds produced. The northernpopulation (50° N) is induced in both photoperiods, whilethe southern population (34° N) is induced only in the 12h photoperiod. Individuals were given either 2, 6, 10 or continuousinductive cycles and dissected at intervals after the startof inductive treatments to determine the rates of initiationand differentiation of primordia on the main axis and selectedaxillary buds. Initial reproductive data indicated that the duration of reproductivedevelopment among individuals of the northern population, whengrown in the longer photoperiod, was 25 per cent greater, butthe number of seeds was increased by a factor of 46. Likewisethe duration of reproductive development among individuals ofthe southern population, when grown in the same photoperiodas the northern population, was 50 per cent longer and the numberof seeds was increased by a factor of 66. Dissection of reproductively-developing individuals revealedthat induction leads to a stimulation in the rate of initiationof primordia followed by a complete inhibition coincident withthe differentiation of terminal floral structures. The timingof this stimulation-inhibition process on each axillary buddepends upon its age relative to the timing of induction. Thuscertain primordia on individuals prematurely removed from inductivetreatments escape floral differentiation and remain vegetative.The total number of floral primordia (potential number of seeds)is determined early in reproductive development by (1) the numberof axillary buds at the start of induction, (2) the stimulatedrate of initiation of primordia after induction, and (3) therate of differentiation of induced primordia. Among individuals of the northern population, the longer inductivephotoperiod leads to a greater stimulation in the rate of primordialinitiation and a decrease in the rate of floral differentiation,which together lead to the production of more primordia. Likewiseamong individuals of the southern population in the shorterphotoperiod, the rate of organ initiation is similar to thatof the northern population in the same photoperiod, but therate of floral differentiation is lower, leading to the initiationof many more primordia. The effects of photoperiod on seed number are discussed in termsof physiological and ecological criteria of optimality. Sincenatural induction occurs in the longest, physiologically-'sub-optimarphotoperiods possible, and this leads to the greatest productionof seeds, it is suggested that the critical photoperiod is amore meaningful focus of interpretation than the physiologically‘optimal’ photoperiod. Of the factors influencing potential seed number, the most significantdifference between the two latitudinal populations is the responseof the rate of floral differentiation to the photoperiod ofinduction. Thus potential seed number in natural populationsis intimately related to the prevailing photoperiods throughthe rates of developmental events. Selection for changes inor maintenance of a particular reproductive ecology must bemediated through developmental responses. Limitations on the potential growth rate of plants are discussedin terms of the ratio of meristematic cells to the total cellpopulation in the plant. Thus the number of growing axillarybuds greatly contributes to the potential growth rate, and thedegree of correlative inhibition is interpreted as a cost ofselection for vertical growth among terrestrial plants competingfor light. By means of a simple model the cost of correlativeinhibition is also discussed in terms of potential seed number.Reproduction is seen as a release from the developmental constraintsrequired by plant form and a stimulation of growth that leadsto a very high production of potential seeds.     

Flowering of Stylosanthes guianensis in Relation to Juvenility and the Long-Short Day Requirement

Seedlings of Stylosanthes guianensis var. guianensis were grownin long (14 h) days in five temperature regimes for varyingperiods before transfer to short (11 h) days at 30 ?C/21 ?C.The juvenile phase before seedlings responded to inductive conditionswas c. 45–50 d, 50–60 d and 60–70 d for cv.Schofield, cv. Cook and C.P.I. 34906 respectively, which ispositively related to their critical photoperiod for flowering.Temperatures favourable for growth (e.g. 30 ?C/26 ?C) reducedthe juvenile phase in C.P.I. 34906 and in Cook, which did notflower in 11 h days unless previously exposed to more than 18long days. In a second experiment cv. Cook was confirmed as a long-shortday plant. Seedlings were grown for 50 d in a glasshouse withnatural daylength extended to 13, 14, 16 or 24 h before transferto 12 h photoperiods. Cook floral development was positivelyrelated to daylength provenance before transfer and plants incontinuous 12 h did not flower. Shortening daylength after 48 cycles of 12 h to 11.75 h didnot result in continued floral development in Cook plants butcv. Graham plants were initiated or transitional by 75 d. Key words: Stylosanthes guianensis, Photoperiod, Temperature, Flowering     

Photoperiod Sensitivity during Flower Development of Opium Poppy (Papaver somniferum L.)

Photoperiod is a major factor in flower development of the opiumpoppy (Papaver somniferum L. ‘album DC’) which isa long-day plant. Predicting time to flower in field-grown opiumpoppy requires knowledge of which stages of growth are sensitiveto photoperiod and how the rate of flower development is influencedby photoperiod. The objective of this work was to determinewhen poppy plants first become sensitive to photoperiod andhow long photoperiod continues to influence the time to firstflower under consistent temperature conditions. Plants weregrown in artificially-lit growth chambers with either a 16-hphotoperiod (highly flower inductive) or a 9-h photoperiod (non-inductive).Plants were transferred at 1 to 3-d intervals from a 16- toa 9-h photoperiod andvice versa . All chambers were maintainedat a 12-h thermoperiod of 25/20 °C. Poppy plants becamesensitive to photoperiod 4 d after emergence and required aminimum of four inductive cycles (short dark periods) beforethe plant flowered. Additional inductive cycles, up to a maximumof nine, hastened flowering. After 13 inductive cycles, floweringtime was no longer influenced by photoperiod. These resultsindicate that the interval between emergence and first flowercan be divided into four phases: (1) a photoperiod-insensitivejuvenile phase (JP); (2) a photoperiod-sensitive inductive phase(PSP); (3) a photoperiod-sensitive post-inductive phase (PSPP);and (4) a photoperiod-insensitive post-inductive phase (PIPP).The minimum durations of these phases forPapaver somniferum‘album DC’ under the conditions of our experimentwere determined as 4 d, 4 d, 9 d, and 14 d, respectively. Anthesis; days to flowering; flower bud; opium poppy; Papaver somniferum L.; photoperiod; photoperiod sensitivity; predicting time to flowering; transfer     

Competence to respond to floral inductive signals requires the homeobox genes PENNYWISE and POUND-FOOLISH

The transition from vegetative to reproductive development establishes new growth patterns required for flowering. This switch is controlled by environmental and/or intrinsic developmental cues that converge at the shoot apical meristem (SAM). During this developmental transition, floral inductive signals cause the vegetative meristem to undergo morphological changes that are essential for flowering. Arabidopsis plants containing null mutations in two paralogous BEL1-like (BELL) homeobox genes, PENNYWISE (PNY) and POUND-FOOLISH (PNF), disrupt the transition from vegetative to reproductive development. These double mutants are completely unable to flower even though the SAM displays morphological and molecular changes that are consistent with having received floral inductive signals. These studies establish a link between the competence to receive floral inductive signals and restructuring of the SAM during floral evocation.     

Development of the shoot apex ofChenopodium rubrum L. after photoperiodic induction in the cotyledon stage

Development of the shoot apex up to floral differentiation was investigated in the short-day plantChenopodium rubrum. The changes occurring in the apex from energence until full opening of the cotyledons (Figs 1–4), development during photoperiodic induction (Figs. 5–8), as well as the resulting floral differentiation (Figs. 9–10) are described. It was aimed at excluding the influence of plastochron changes on the interpretation of ontogeny of the shoot apex. For that reason two planes of longitudinal sections and two plastochron stages were compared. In young plants zonation does not become fully evident prior to floral differentiation. The anatomical structure of the shoot apex does not change substantially during the first two inductive cycles which proved to be obligatory under the given experimental conditions. The changes occurring during two further inductive cycles correspond to the total activation of the meristems as manifested by the growth and branching of the apex preceeding floral differentiation proper.     

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.     

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     

Effect of varying lengths of inductive and supplementary non-inductive photoperiods on vegetative growth and flowering of Impatiens balsamina

The photoperiodic requirement for flowering in Impatiens balsaminachanges with the length of the photoperiod. Floral buds wereinitiated with two 8 hr but with four 15 hr photoperiods andflowers opened with four 8 hr but twenty-eight 15 hr photoperiods.A part of the photoperiodic requirement for floral inductionin this plant can be substituted by LDs containing 4 or morehours of darkness (10). It indicates the identical nature ofthe floral stimulus produced during the dark period, whetherit forms a part of the inductive or non-inductive cycles. Theeffect of these supplementary non-inductive photoperiodic cyclesin causing floral bud initiation also depends on the lengthof the first inductive obligatory cycle. More floral buds andflowers were produced on plants exposed to 15 hr than 8 hr photoperiods,probably due to the higher number of leaves that were producedunder the former condition of weaker induction. The shorterthe dark period in the photoperiodic cycle, the weaker the induction,the slower the rate of extension growth but the more differentiationof leaves. ¹ Present address: Department of Biology, Guru Nanak Dev University,Amritsar-143005, India. (Received November 9, 1977; )     

Short-day and Low-temperature Control of Floral Induction in Festuca

The conditions necessary for floral induction to occur in tallfescue (Festuca arundinacea), meadow fescue (Festuca pratensis),and red fescue (Festuca rubra), have been investigated. Onlya Tunisian ecotype of tall fescue produced inflorescences undershort-day conditions when air temperatures were above 8 °C.Under short days with low temperatures nearly all plants ofS. 170 tall fescue and S. 215 meadow fescue produced inflorescencesafter 15 weeks' exposure, but S. 59 red fescue showed only asmall response. Evidence was obtained for the existence in bothtall fescue and meadow fescue of a juvenile stage during whichplants showed a reduced response to inductive conditions. Avariation of 35 days in the required length of exposure to inductiveconditions was demonstrated between families within the S. 170variety of tall fescue, indicating the possibility of selectingfor larger or smaller inductive requirements. A second generationof seed was produced within a 12-month period from inflorescenceswhich had developed in a heated glasshouse during the wintermonths.     

Rhythmicity of Flowering in Pharbitis nil

When young seedlings of Pharbitis nil are grown under continuous light, except for a single inductive dark period, they flower to a varying degree, depending on when this dark period is given. Plants become sensitive to this induction approximately three days after the seedlings emerge from the soil. The expression of flowering varies in a rhythmic fashion for three or more cycles, when an inductive dark period is given at progressively later times. The time between maximum expression of flowering is 24 hours or somewhat longer. It appears necessary that the inductive dark period be of sufficient duration, to only partially induce the plants to flower for this rhythm to be expressed. Under the conditions employed in this study, this duration is 12 hours. If this rhythm is endogenous, it exists at least from when the plants emerged from the soil since no environmental cues are given after that time, and it raises questions of the interpretations of data from previous studies with this organism.     

IAA-oxidase in Impatiens balsamina Affected by GA3 and Tannic Acid

The activity of IAA-oxidase increased in the leaves of Impatiensbalsamina plants receiving inductive photoperiodic cycles andin plants receiving treatments with gibberellic acid (GA₃) and/ortannic acid (TA), even under non-inductive photoperiods; theactivity also increased in the stem receiving inductive photoperiodiccycles (8 h). Treatment with GA₃ and TA mimics the effect ofSD cycles in the development of some isoenzymes of IAA-oxidase.Thus a new isoenzyme at R_f 0.48 developed in the leaves andone at R_f 0.82 developed in both the stem and the leaves ofall plants receiving inductive treatments – photoperiodicor chemical – but not in water-treated controls undernon-inductive photoperiods. Another isoenzyme at R_f 0.68 developedonly in the stems. Flowering, gibberellic acid, IAA oxidase, Impatiens, phenols, photoperiod     

Role of the Photoperiod Preceding a Flower-Inductive Dark Period in Dark-Grown Seedlings of Pharbitis nil Choisy

Flowering responses to a single photoperiod, of various durationsand irradiances, followed by an inductive dark period were investigatedwith dark-grown seedlings of Pharbitis nil Choisy. The numberof flower buds induced in each plant (NFB) increased with theincrease of both duration and irradiance of the photoperiod.Reciprocity did not hold for this photoresponse within the rangeof 0-16 h and 2.5-10 W-m^-2, NFB depending on the duration ratherthan the irradiance. With lengthening of the dark period followinga photoperiod of 8 h or less, two different phases alternatelyappeared so that NFB sharply increased at 20-24 h and 40-43h after the onset of the photoperiod, then gradually decreased.When the photoperiod was longer than 8 h, NFB sharply increasedat 12–16 h after the end of the photoperiod and remainedaround the saturated value with longer dark periods. Far-redlight given immediately after the photoperiod inhibited flowering,the inhibitory effect being stronger the shorter the photoperiod.This far-red effect is mediated by phytochrome and P_FR seemsto be required during the inductive dark period following ashort photoperiod for floral induction. (Received December 23, 1983; Accepted April 12, 1984)     

Bimodal Floral Response of Lemna gibba G3 to Night Interruption: Photoperiodic Time Measurement

Lemna gibba G3 (M-1% sucrose medium, 26°C) showed a bimodalfloral response to a 2-hr light pulse scanning 21-, 18-, 15-and 12-hr nyctoperiods. With the simplified min-LD method, thelight pulse given early or late in these nyctoperiods was foundto signal false dusk or false dawn after two transient cycles.The magnitude of floral response to the light pulse dependedon the length of the asymmetric skeleton photoperiod comprisingeither the preceding main photoperiod and the false dusk orthe false dawn and the subsequent main photoperiod. No flowerwas induced by asymmetric skeleton photoperiods shorter thanthe critical daylength, 12 hr. In duckweed previously entrainedto an interrupted 15-hr nyctoperiod, false dawn or false duskwas physiologically equivalent to the light-requiring L1- orL2-phase of the critical photoperiod. Another light-requiringphase occurred 12 hr after or before the false dawn or falsedusk. These and relevant findings suggest that the timing ofthe L1- and L2-phases is under the control of the endogenouscircadian oscillator and that the skeleton as well as completephotoperiods are inductive only when both the L1- and L2-phases,whether they are shifted or not due to night interruption, areilluminated. (Received October 3, 1980; Accepted December 3, 1980)     

Requirements for Floral Induction in Contrasting White Clover (Trifolium repens) Populations

Norm, I. B. 1987. Requirements for floral induction in contrastingwhite clover (Trifolium repens) populations.—J. exp. Bot.38: 900–907. Floral initiation and development of four contrasting whiteclover (Trifolium repens) populations was examined after differentinduction treatments (16 h, 5 ?C and 8 h, 5 ?C. The number of reproductive stolons and of reproductive budsper stolon was increased after cold induction. Varietal differencesin response to daylength were large; some varieties respondingbetter to a long day cold period, others to a short day coldperiod while one variety required no induction at all. Whetherthe daylength effect was due to photoperiod, irradiance or totheir interaction was not known. The induction periods had a subsequent effect upon pedunclelength, floret and ovule number. Short days and chilling reducedpeduncle length but increased ovule number, whereas long daysand chilling tended to increase floret number. Nectar concentrationwas highest after short day induction. Key words: White clover, floral initiation, floral induction     

Darkness promotes flowering in the absolute long-day requiring plant, Lolium temulentum L. Ceres

Vegetative plants of the long-day grass Lolium temulentum L.Ceres were exposed to threshold long days or light breaks. Protracteddarkness given just afterwards clearly promoted flowering andwas weakly inductive on its own. The promotive effect of darknesswas restricted to floral induction since further apical developmentwas weak. Key words: Lolium temulentum, flowering, photoperiodism, darkness     

delayed flowering1 Encodes a basic leucine zipper protein that mediates floral inductive signals at the shoot apex in maize

Separation of the life cycle of flowering plants into two distinct growth phases, vegetative and reproductive, is marked by the floral transition. The initial floral inductive signals are perceived in the leaves and transmitted to the shoot apex, where the vegetative shoot apical meristem is restructured into a reproductive meristem. In this study, we report cloning and characterization of the maize (Zea mays) flowering time gene delayed flowering1 (dlf1). Loss of dlf1 function results in late flowering, indicating dlf1 is required for timely promotion of the floral transition. dlf1 encodes a protein with a basic leucine zipper domain belonging to an evolutionarily conserved family. Three-dimensional protein modeling of a missense mutation within the basic domain suggests DLF1 protein functions through DNA binding. The spatial and temporal expression pattern of dlf1 indicates a threshold level of dlf1 is required in the shoot apex for proper timing of the floral transition. Double mutant analysis of dlf1 and indeterminate1 (id1), another late flowering mutation, places dlf1 downstream of id1 function and suggests dlf1 mediates floral inductive signals transmitted from leaves to the shoot apex. This study establishes an emergent framework for the genetic control of floral induction in maize and highlights the conserved topology of the floral transition network in flowering plants.     

Photoperiod and Temperature Regulation of Floral Initiation and Anthesis in Soya Bean

Floral development includes initiation of floral primordia andsubsequent anthesis as discrete events, even though in manyinvestigations only anthesis is considered. For ‘Ransom’soya bean [Glycine max (L.) Merrill] grown at day/night temperaturesof 18/14, 22/18, 26/22, 30/26, and 34/30 °C and exposedto photoperiods of 10, 12, 14, 15, and 16 h, time of anthesisranged from less than 21 days after exposure at the shorterphotoperiods and warmer temperatures to more than 60 days atlonger photoperiods and cooler temperatures. For all temperatureregimes, however, floral primordia were initiated under shorterphotopenods within 3 to 5 days after exposure and after notmore than 7 to 10 days exposure to longer photoperiods. Onceinitiation had begun, time required for differentiation of individualfloral primordia and the duration of leaf initiation at shootapices increased with increasing length of photoperiod. Whileproduction of nodes ceased abruptly under photoperiods of 10and 12 h, new nodes continued to be formed concurrently withinitiation of axillary floral primordia under photoperiods of14, 15 and 16 h. The vegetative condition at the main stem shootapex was prolonged under the three longer photoperiods and issuggestive of the existence of an intermediate apex under theseconditions. The results indicate that initiation and anthesisare controlled independently rather than collectively by photoperiod,and that floral initiation consists of two independent steps—onefor the first-initiated flower in an axil of a main stem leafand a second for transformation of the terminal shoot apex fromthe vegetative to reproductive condition. Apical meristem, intermediate apex, floral initiation, anthesis, photoinduction, Glycine max(L.) Merrill, soya bean, photoperiod, temperature