Qualitative analysis by isoelectric focusing of the protein content of Pharbitis nil apices and cotyledons during floral induction

The protein content of apices and cotyledons in Morally inducedor vegetative plants of Pharbitis nilwas examined using isoelectricfocusing. No differences were found in the protein patternsproduced by apical tissue with and without floral induction.Cotyledons, however, repeatedly showed the distinct loss ofa single protein band on floral induction. ¹Current address: Department of Radiation Biology and Biophysics,The University of Rochester School of Medicine and Dentristry,Rochester, N.Y. 14642, U.S.A. (Received March 29, 1976; )     

Rates of Growth and Cell Division in the Shoot Apex of Silene During the Transition to Flowering

The growth rates of the shoot apex during and after floral inductionwere measured in Silene, a long-day plant. Plants were inducedto flower with 4 or more long days (LD) but not with 3 longdays or with short days (SD). The rate of increase of cell numberin the apical dome, above the youngest leaf pair, was exponentialand in plants given 3 LD remained the same as in plants in SD.In plants induced to flower with 7 LD, until the end of theinductive period the rate of increase of cell number in theapical dome remained the same as in plants in SD. Only whenthe apex began to enlarge as the first stage in the formationof the flower did the growth rate of the apical dome increase.The rates of increase of cell numbers in the apex correspondedto mean cell generation times of 20 to 33 h for plants in SD,for plants given 3 LD, and during the 7 days of induction forplants given 7 LD, and 6 to 8 h for induced plants when flowerformation was beginning. The distribution of cell division in the apex was examined bytreating plants with colchicine and noting in sections the positionsof the resulting metaphases. In vegetative apices and also inapices undergoing transition to flowering the whole of the apicaldome appeared to consist of cells dividing at a similar rate. The rate of leaf initiation during induction was the same asin vegetative, non-induced plants.     

Initial Anatomical Changes Associated with Tuber Formation on Single-node Potato (Solanum tuberosum L.) Cuttings: A Re-evaluation

Cell division and cell expansion during early stages of tuberdevelopment were studied using developing axillary buds on single-leafcuttings from potato (Solanum tuberosum L.). Cuttings takenfrom plants induced to form tubers, by short day (SD) treatment,were compared with cuttings from non-induced (long day, LD)plants. In the apical zone of the buds, cell division occurredfrom the first day after cutting, in both LD and SD cuttings.The planes of these divisions were transverse, associated withelongation of the buds. At day 5, a new orientation of celldivision was observed in the subapical zone of SD cuttings only.These divisions were longitudinal, associated with radial growth.Cell expansion occurred in both SD and LD cuttings, and wasnot uniquely related to the onset of tuber formation. Copyright1999 Annals of Botany Company Solanum tuberosum L., potato, tuber formation, cell division, cell expansion.     

The Pattern of Reproductive Development in Chenopodium rubrum L.

Individual plants of Chenopodium rubrum were given differentnumbers of inductive cycles in a 12 h photoperiod and the patternof reproductive development was analysed after 40 d of growth.At least 2 inductive cycles are required to form any determinatereproductive organs and at least 12 cycles are required fornormal reproductive development. Individuals given a singleinductive cycle display a loss of apical dominance at thosenodes formed immediately after the treatment without the subsequentformation of any floral structures. Plants given between 2 and12 mductive cycles display both determinate reproductive organsand indeter minate vegetative shoots. The pattern of reproductivedevelopment on such plants depends upon the number of cyclesrelative to the developmental age of newly initiated primordia.It is suggested that the early events of floral induction mayinvolve a radical decrease in the ratio of auxin to cytokinin.     

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     

Effects of Abscisic Acid on the Growth Pattern of the Shoot Apical Meristem and on Flowering in Chenopodium rubrum L.

Abscisic acid (ABA) at 1 x 10^–4 M or 3 x 10^–4 Mwas applied to the apical buds of Chenopodium rubrum plantsexposed to different photoperiodic treatments and showing differentpatterns of floral differentiation. Stimulation of growth inwidth of the apical meristem of the shoot and/or inhibitionof growth in length was obtained under all photoperiodic treatments.This change of growth pattern was followed by different effectson flowering. In non-induced plants grown under continuous light ABA stimulatedpericlinal divisions in the peripheral zone and the initiationof leaves as well as the growth in width of bud primordia. Inplants induced by two short days reduced growth of the meristemcoincided with ABA application. Longitudinal growth of the meristemwas inhibited in this case and only a temporary stimulationof inflorescence formation took place. In plants induced ata very early stage, ABA exerted a strong inhibitory effect onflowering. A permanent and reproducible stimulatory effect onflowering was obtained in plants induced by three sub-criticalphotoperiodic cycles if ABA was applied to apices released fromapical dominance. In this case formation of lateral organs andinternodes was promoted by ABA and was followed by stimulatedinflorescence formation. Gibberellic acid (GA2) at 1x 10^–4M or 3 x 10^–4 M brought about a similar effect on floweringas ABA, although the primary growth effect was different, i.e.GA₂ stimulated longitudinal growth. The effects of ABA and GA₂ on floral differentiation have beencompared with earlier results obtained from auxin and kinetinapplications. These growth hormones are believed to regulateflowering by changing cellular growth within the shoot apex.Depending on the actual state of the meristem identical growthresponses may result in different patterns of organogenesisand even in opposite effects on flowering. Shoot apex, flowering, photoperiodic induction, abscisic acid, gibberellic acid, Chenopodium rubrum L.     

Cell Cycle Changes in the Shoot Apex of Silene coeli-rosa During the Second and Third Days of Floral Induction

The cell cycle in Silene coeli-rosa shoot apices was measuredto test whether or not early components of the floral stimulus,produced during the 2nd and 3rd long days (LD) of an inductiveLD treatment, resulted in an increase in the duration of G₂phase in constant 20–24 h cell cycles. Plants were grownat 20°C in short days (SD) of 8 h light and 16 h darknessfor 28 d (day 0). Starting on day 0, plants were given SD or3 LD each comprising an identical 8 h day and 16 h photo-extension,or 3 dark-interrupted (d.i.) non-inductive LD, interrupted at1700 h of each day with 1 h of darkness. The cell cycle (percentagelabelled mitoses method) and changes in cell number were determinedin the shoot apical meristem. During days 1–2 of the SDtreatment, the cell cycle and mean cell generation time (MCGT)was 18 and 32 h, respectively, giving a growth fraction of 56%.During days 2–3, the cell cycle and MCGT shortened to15 and 23 h, respectively (growth fraction = 65%). During days1–2 of the LD and d.i. LD treatments, cell cycles andMCGTs were 9–10 and 27–29 h, respectively, resultingin smaller growth fractions (about 33%). Thus, shortened cellcycles and altered growth fractions occurred regardless of whetheror not the treatment was inductive. The LD treatment resultedin a marked shortening of G₁ and, to a lesser extent, S-phase,whilst G₂ remained constant. These changes were consistent withincreases in the proportion of cells in G₂ during the photoextensionof each LD which were suppressed during the comparable periodsof the d.i. LD treatment. The latter treatment resulted in eachphase occupying virtually identical proportions of the cellcycle as in the SD treatment. Thus, the unique cell cycle responsesto the initial part of the inductive LD treatment were increasesin the proportion of cells in G₂ coupled with G₁ and G₂ beingof similar duration. Cell cycle, mean cell generation time, shoot apex, Silene coeli-rosa     

Genetic Regulation of Development in Sorghum bicolor: IV. GA(3) Hastens Floral Differentiation but Not Floral Development under Nonfavorable Photoperiods

Sorghum bicolor (L.) Moench lines with genetic differences in photoperiod requirement were planted in the field near Plainview, Texas (about 34° northern latitude) around June 1 and treated with gibberellic acid (GA₃) solutions applied in the apical leaf whorl. GA₃ hastened the date of floral differentiation (initiation). The greatest responses to GA₃ were by 90M and 100M, the latest of the genotypes, for which floral initiation dates were hastened an average of 19.5 and 21.7 days, respectively, for the 4 years beginning in 1980. There were very small differences in dates of anthesis between control and GA₃-treated plants. Microscopic examination of apical meristems collected between the date of floral initiation of GA₃-treated plants and the later date of initiation of control plants revealed: (a) several morphological characteristics of floral differentiation in the apical meristem of treated plants, (b) consistent occurrence of vegetative morphology in control plants, (c) a few meristems from GA₃-treated plants that appeared to be regressing in floral development and thus possibly exhibiting dedifferentiation. Dedifferentiation of prepanicle primordia into leaves would explain the observed equal or greater number of leaves in GA₃-treated plants rather than the expected smaller number. It is apparent that the presence of a morphological differentiated floral meristem in sorghum does not drive subsequent floral development in the absence of inductive photoperiods. This further suggests that initial floral differentiation and subsequent floral development may be controlled separately in sorghum.     

Changes in the Pattern of Cell Division in the Shoot and Root Meristems of Lolium perenne during the Transition from Vegetative to Floral Growth

For Lolium perenne cv. Cropper, a system which resulted in 100%flowering comprised 90 short days (SD) at 4 ?C (vernalization)and 30 SD at 18 ?C followed by 8 long days (LD). The mitoticindex and G1 and G2 percentages were measured in the shoot androot apices of plants following 2, 5 or 8 LD and in SD controlssampled at the beginning and end of induction. Identical measurementswere made in plants given 48 SD at 18 ?C followed by 2, 5 or8 LD; plants remained vegetative in response to this treatmentlacking vernalization. Significant increases in both mitoticindex and meristem size occurred in the shoot apex in LD followingthe vernalizing, but not the non-vernalizing, treatment. A clusterof mitoses in the apical dome of the shoot apex was unique tothe vernalized plants given 5 or 8 LD. However, an increasein root meristem size occurred regardless of vernalization,but a significant increase in the mitotic index was limitedto vernalized plants given 5 or 8 LD. Whilst the vernalization-LDtreatment resulted in an increase in the G2 percentage in theshoot apex following 2, 5 or 8 LD, no such alteration was observedin the root meristem. Thus, the changes to the cell cycle whichcorrelated with flowering were increased mitotic indices andG2 percentages in the shoot apex at each sampling time and increasedmitotic indices in the root apex following 5 and 8 LD. Key words: Cell division, flowering, Lolium perenne L.     

Effect of Photoperiod and Electrical Potentials Applied to Petioles on the Glucose-6-phosphate Dehydrogenase Activity used as Marker of Floral Induction in Shoot Apices of Spinach I

The increase of glucose-6-phosphate dehydrogenase (G-6-PD) activityhas been proposed as an early marker of floral evocation inthe shoot apical meristem of spinach. This induction is obtainedby the transfer of vegetative plants from short days to continuouslight. The exposure of a single leaf to continuous light inducedthe same effect as that produced with the whole plant. In vegetativeconditions (short days), an electrical potential of ten volts(electrical current: 12·5 µA) applied to the petioleof a single leaf induced a weak increase of G-6-PD activityin the shoot apex, while under inductive conditions, this increasewas similar to that of control plants (transferred to continuouslight). Application of an electrical potential to the petioleand the root inhibits the increase linked to the inductive transfer.These results show that an externally applied electrical potentialmay interact with the natural electrical gradients along theshoot and consequently inhibit the normal floral developmentby an as yet unspecified mechanism. Spinacia oleracea, floral induction, interorganic relations, glucose-6-phosphate dehydrogenase, shoot apex, electrical potentials     

Development of Floral Apex After Floral Induction in Stylo (Stylosanthes guianensis (Aubl.) Sw. var. guianensis)

The apex morphology of stylo (Stylosanthes guianensis var. guianensis)is described in four developmental phases (vegetative, transitional,initiated and floral) further subdivided into a total of tenstages. The apical dome broadens and flattens as induction proceedsuntil the initiation phase when apical diameter within 0.05mm of the dome apex is 55 per cent greater than in the vegetativeapex. Changes in vegetative morphology during induction aredescribed. Stylosanthes guianensis var. guianensis, stylo, flowering, reproductive anatomy, developmental stages     

Occurrence of the Transition of Apical Architecture and Expression Patterns of Related Genes during Conversion of Apical Meristem Identity in G2 Pea

G2 pea exhibits an apical senescence delaying phenotype under short-day (SD) conditions; however, the structural basis for its apical development is still largely unknown. In the present study, the apical meristem of SD-grown G2 pea plants underwent a transition from vegetative to indeterminate inflorescence meristem, but the apical meristem of long-day (LD)-grown G2 pea plants would be further converted to determinate floral meristem. Both SD signal and GA3 treatment enhanced expression of the putative calcium transporter PPF1, and pea homologs of TFL1 (LF and DET), whereas LD signal suppressed their expression at 60 d post-flowering compared with those at 40 d post-flowering. Both PPF1 and LF expressed at the vegetative and reproductive phases in SD-grown apical buds, but floral initiation obviously increased the expression level of PPF1 compared with the unchanged expression level of LF from 40 to 60 d post-flowering. In addition, although the floral initiation significantly enhanced the expression levels of PPF1 and DET, DET was mainly expressed after floral initiation in SD-grown apical buds. Therefore, the main structural difference between LD- and SD-grown apical meristem in G2 pea lies in whether their apical indeterminate inflorescence medstem could be converted to the determinate structure.     

Gibberellin Is Involved in the Regulation of Cell Death-mediated Apical Senescence in G2 Pea

Senescence is the process of programmed degradation. The G2 line of pea exhibits apical senescence-delaying phenotype under short-day （SD） conditions, but the mechanism regulating the apical senescence is still largely unknown. Gibberellin （GA） was proved to be able to delay this apical senescence phenotype in G2 pea grown under long-day （LD） conditions. Here we show that the initiation of cell death signals in the terminal floral meristem was involved in the regulation of apical senescence in pea plants. SD signals prevented the formation of the cell death region in the apical mersitem. Moreover, GA3 treatment could effectively inhibit the occurrence of cell death-mediated apical senescence in LD-grown apical buds. Therefore, our data suggest that the prevention of apical senescence in SD-grown G2 pea through GA3 treatment may be largely responsible for the regulation of occurrence of the DNA fragmentation in apical meristem.     

Environmental Control of Flowering in some Northern Carex Species

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

Effects of Indol-3yl acetic Acid on Floral Induction and Apical Differentiation in Chenopodium rubrum L.

Indol-3yl acetic acid (10^–4M) was applied to the plumulesof Chenopodium rubrum. Effects on the anatomical structure andthe growth pattern in the apical meristem, as well as DNA synthesisand nucleolus size were investigated. When auxin is applied before or during photoperiodic inductionit inhibits DNA synthesis and meristematic activity. The axillarymeristem (i.e. a group of cells in the axils of the leaf primordia)is most affected. A similar inhibition of the axillary meristemwas also observed in non-induced control plants grown in continuouslight. Auxin applied simultaneously with photoperiodic inductioncounteracts the reduction of apical dominance in the apex andthus inhibits the onset of floral differentiation. Auxin appliedfollowing induction inhibits the previously-formed buds andmakes possible a more complete development of the apical flower. The dual effect of IAA on flowering, inhibitory and stimulatory,manifests itself as a growth response at different stages ofthe changing shoot apex.     

Effect of Gibberellic Acid and Monophenols on Flowering of lmpatiens balsamina in Relation to the Number of Inductive and Non-inductive Photoperiodic Cycles

A single treatment of plants with GA₃ (gibberellic acid) is not adequate to cause induction under LD (long day: 24-h photo-period) condition, but its effect is added to the sub-threshold induction caused by one SD (short day: 8-h photoperiod) cycle. Floral bud initiation is hastened, and the number of floral buds and flowers per flowering plant increases in plants receiving a single treatment with the combination GA₃+ SA (salicylic acid) accompanying a single SD cycle. However, the increase on 10 replicate basis is more marked in plants receiving three treatments with the combination GA₃+β-N (β-naphthol) and five treatments with the combination GA₃+ SA accompanying six and 10 SD cycles, respectively. The number of floral buds and flowers decreases with an increase hi the number of SD cycles, but it is higher in plants treated with GA₃, SA or GA₃+β-N than in the water-treated controls. — Under long days, treatment of plants with the combinations GA₃+ SA or GA₃+β-N accelerates the initiation as well as increases the number of floral buds. While a minimum of five treatments with GA₃ or of 25 with SA or β-N alone is needed for floral bud initiation under a 24-h photoperiod, three treatments are adequate to induce floral buds with the combination GA₃+ SA or GA₃+β-N under continuous illumination. Ten or more treatments with these combinations under a 24-h photoperiod produce more flowers than the same treatments under an 8-h photoperiod.     

Developmental physiology of floral initiation in Nicotiana tabacum L.

The central process in the making of a multicellular organismis the fating of cells and tissues for their terminal phenotypes.The formation of a flower from a shoot apical meristem completesa sequence of fating processes initiated in embryogenesis. Thefating of a vegetative meristem of Nicotiana tabacum L. to initiatea flower involves at least two signals and two developmentalstates. A signal from the roots maintains vegetative growth,or prevents flowering, in the young seedling. As the plant grows,the vegetative meristem gains greater competence to respondto the floral stimulus from the leaves until it is evoked, byfloral stimulus, into a florally determined state. The florallydetermined state is then expressed. These developmental processesnot only establish the time of floral initiation, but also regulateplant size as measured by the number of nodes produced. Key words: Plant size, floral stimulus, competence, floral determination, induction     

Induction of flowering ofImpatiens balsamina treated with gibberellic acid and resorcinol

Under strictly non-inductive photoperiods (24-h photoperiods) floral buds were initiated on plants receiving 25 treatments with Reso (resorcinol) or 8 treatments with GA₃ (gibberellic acid) or GA₃ + Reso, while water treated control plants did not flower at all. Although a single treatment of plants with GA₃ or GA₃ + Reso is not adequate to cause induction under LD conditions, its effect is added to the sub-threshold induction caused by one SD (short day: 8-h photoperiod) cycle. The initiation of floral buds was hastened with an increasing number of SD cycles accompanying respective number of treatments, the effect of GA₃ alone or together with Reso being more pronounced than that of Reso alone. GA₃ increased the number of floral buds more than Reso, the number being the highest in plants receiving the respective number of treatments with the combination GA₃ + Reso under both inductive as well as non-inductive photoperiods. Deceased.     

The effect of the floral repressor FLC on the timing and progression of vegetative phase change in Arabidopsis

Plants undergo two major post-embryonic developmental transitions--the juvenile-to-adult vegetative transition (vegetative phase change) and the adult-to-reproductive transition (flowering). In woody plants, these transitions can be separated by years, but in herbaceous species they are often very close together, making it difficult to differentiate the effects of vegetative phase change and floral induction on vegetative development. To distinguish between these factors, we have compared the vegetative morphology of plants highly expressing the floral repressor FLC (FRI;FLC) with plants mutant for this gene (FRI;flc-3) under both photoinductive (long day, LD and night interruption, NI) and non-photoinductive (short day, SD) conditions. We show that the onset of abaxial trichome production is insensitive to floral induction, but the distribution and overall number of abaxial trichomes, as well as several other leaf traits associated with vegetative change, are strongly influenced by flowering. Most of the major differences in leaf morphology between FRI;FLC and FRI;flc-3 plants grown in LD can be attributed to the early flowering phenotype of FRI;flc-3, because these differences are not apparent in plants grown in SD. These include differences in leaf size, hydathode number and the distribution of abaxial trichomes along the length of the leaf. Leaf shape and the total number of abaxial trichomes are affected by FLC independently of its effect on flowering. Our results demonstrate that the onset and the progression of vegetative phase change are regulated by different combinations of endogenous and environmental factors, and reveal a role for FLC in vegetative development.