Genetic Regulation of Development in Sorghum bicolor: II. Effect of the ma(3) Allele Mimicked by GA(3)

The photoperiodic behavior and other developmental and morphological differences of 11 maturity genotypes (as identified by JR Quinby 1967, Adv Agron 19: 267-305) of the milo group of Sorghum bicolor (L.) Moench were studied under 8, 10, 12, and 14 hour photoperiods. Sorghum is a quantitative short day plant. The genotypes studied differ in genes which modify photoperiodic behavior and thus maturity; the alleles are designated as Ma₁, ma₁, Ma₂, ma₂, Ma₃, ma₃, and ma₃^R (single symbols indicate homozygosity at the indicated gene loci). Based on floral initiation (differentiation) under 10, 12, and 14 hour photoperiods the 11 genotypes were assigned to three clases: (I) flower initiation delayed by 12 hour photoperiods (all genotypes with Ma₁Ma₂ but not ma₃^R), (II) flower initiation delayed by 14 hour photoperiods (all genotypes with Ma₁ma₂, ma₁Ma₂, or ma₁ma₂ but not ma₃^R), (III) flower initiation not drastically delayed by 14 hour photoperiods (all genotypes with ma₃^R). All of the class III genotypes were taller, had longer leaf sheaths, narrower and longer leaf blades, and less leaf area, than the other genotypes. In addition, the class III genotypes initiated rapid culm and thus internode elongation sooner after floral initiation than any of the class I or II genotypes. Dry weight did not differ between the class III genotypes and the others. The rate of leaf emergence in the class III genotypes and all others was indistinguishable until after floral initiation in the former. The allelic combination unique to class I, Ma₁Ma₂, makes plants very photoperiod sensitive without causing observable changes in morphology or other development events. The allelic combination unique to class III, ma₃^R, makes plants relatively photoperiod insensitive and results in several differences in morphology and development.     

Photoperiod Control of Gibberellin Levels and Flowering in Sorghum

Regulation of rhythmic peaks in levels of endogenous gibberellins (GAs) by photoperiod was studied in the short-day monocot sorghum (Sorghum bicolor [L.] Moench). Comparisons were made between three maturity (Ma) genotypes: 58M (Ma₁Ma₁, Ma₂Ma₂, phyB-1phyB-1, and Ma₄Ma₄ [a phytochrome B null mutant]); 90M (Ma₁Ma₁, Ma₂Ma₂, phyB-2phyB-2, and Ma₄Ma₄); and 100M (Ma₁Ma₁, Ma₂Ma₂, PHYBPHYB, and Ma₄Ma₄). Plants were grown for 14 d under 10-, 14-, 16-, 18-, and 20-h photoperiods, and GA levels were assayed by gas chromatography-mass spectrometry every 3 h for 24 h. Under inductive 10-h photoperiods, the peak of GA₂₀ and GA₁ levels in 90M and 100M was shifted from midday, observed earlier with 12-h photoperiods, to an early morning peak, and flowering was hastened. In addition, the early morning peaks in levels of GA₂₀ and GA₁ in 58M under conditions allowing early flowering (10-, 12-, and 14-h photoperiods) were shifted to midday by noninductive (18- and 20-h) photoperiods, and flowering was delayed. These results are consistent with the possibility that the diurnal rhythm of GA levels plays a role in floral initiation and may be one way by which the absence of phytochrome B causes early flowering in 58M under most photoperiods.     

Effect of Gibberellic Acid and Phosfon on the Critical Dark Period Reqnirement for Flowering of Impatiens balsamina cv. Rose

The critical dark period requirement for flowering of Impatiens balsamina L. cv. Rose, an obligate short day plant, is about 8.5 hours. While GA₃ completely substituted for the dark period requirement, Phosfon prolonged it to 9.5 hours. GA₃ hastened and Phosfon delayed the initiation of floral buds under all photoperiods. Floral buds opened into flowers only during 8 and 14 hour photoperiods in control and Phosfon-treated plants but during all photoperiods in GA₃-treated ones. The delay in floral bud initiation and flowering was correlated with shifting up of the node bearing the first floral bud and flower respectively. While GA₃ increased the numher of floral buds and flowers in all photoperiods except 8-hour, Phosfon increased their number in the 14-hour photoperiod only. The number of flowering plants decreased with increasing photoperiod regardless of GA₃ and Phosfon application. The effect of Phosfon was completely or partially overcome, depending upon the photoperiod, by simultaneous application of GA₃.     

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.     

Effects of Photoperiod, Temperature and Asynchrony between Thermoperiod and Photoperiod on Development to Panicle Initiation in Sorghum

The duration of the vegetative phase (i.e. days from sowingto panicle initiation) in sorghum [Sorghum bicolor (L.) Moench]is affected by photoperiod and temperature. Plants of severalcontrasting genotypes of sorghum were grown in controlled-environmentgrowth cabinets with either synchronous or asynchronous photoperiodsand thermoperiods. Apical development was recorded. Diurnalasynchrony between photoperiod and thermoperiod reduced durationsto panicle initiation when the temperature warmed after lightswent on and cooled after lights went off, but increased thesedurations when the temperature warmed before lights went onand cooled before lights went off. These effects were shownin the maturity lines 60M and SM100 and also in the USA cv.RS610 and the Sudanese landrace IS22365, but their magnitudevaried with genotype, photothermal regime, and the degree ofasynchrony. The greatest effect was detected in IS22365 grownat 30/21 °C (12 h/12 h) with a 12 h d^-1photoperiod whenthe temperature warmed 2.5 h before lights went on and cooled2.5 h before lights went off, when the duration from sowingto panicle initiation was 69 d compared with 37 d in the control(synchronous photoperiod and thermoperiod in each diurnal cycle). Reciprocal transfers of plants of IS22365 between short andlong days revealed that asynchrony principally affected theduration of the photoperiod-insensitive pre-inductive phaseof development; i.e. asynchrony affected the time (age) at whichthe plants were first able to respond to photoperiod. In thatinvestigation in controlled-environment growth chambers, thesubsequent photoperiod-sensitive inductive phase continued untilpanicle initiation. Subsequent reciprocal transfer experimentsin controlled-environment glasshouses in four different alternatingtemperature regimes employed synchronous photoperiods and thermoperiodsin short (11 h) days with temperature warming 1.5 h after thebeginning of the day in long (12.5 h) days. In those investigations,photoperiod sensitivity ended some time before (2.5–8.1d, mean 5.7 d) panicle initiation in IS22365, Naga White andSeredo. Moreover, whereas the duration of the photoperiod-insensitivepre-inductive phase was affected by temperature, the durationsof the photoperiod-sensitive inductive and the photoperiod-insensitivepost-inductive phases were not. Sorghum bicolor (L.) Moench; sorghum; asynchrony; photoperiod; thermoperiod; vegetative phase; panicle initiation     

Molecular Evolution of the Sorghum Maturity Gene Ma3

Time to maturity is a critical trait in sorghum (Sorghum bicolor) breeding, as it determines whether a variety can be grown in a particular cropping system or ecosystem. Understanding the nucleotide variation and the mechanisms of molecular evolution of the maturity genes would be helpful for breeding programs. In this study, we analyzed the nucleotide diversity of Ma₃, an important maturity gene in sorghum, using 252 cultivated and wild sorghum materials from all over the world. The nucleotide variation and diversity were analyzed based both on race- and usage-based groups. We also sequenced 12 genes around the Ma₃ gene in 185 of these materials to search for a selective sweep and found that purifying selection was the strongest force on Ma₃, as low nucleotide diversity and low-frequency amino acid variants were observed. However, a very special mutation, described as ma₃^R, seemed to be under positive selection, as indicated by dramatically reduced nucleotide variation not only at the loci but also in the surrounding regions among individuals carrying the mutations. In addition, in an association study using the Ma₃ nucleotide variations, we detected 3 significant SNPs for the heading date at a high-latitude environment (Beijing) and 17 at a low-latitude environment (Hainan). The results of this study increases our understanding of the evolutionary mechanisms of the maturity genes in sorghum and will be useful in sorghum breeding.     

Floral initiation in sorghum hastened by gibberellic acid and far-red light

Combinations of far-red light (FR) (4 min) and gibberellic acid (GA₃), given at the beginning of a daily 12-h dark period in a growth room, were used to study floral induction in four maturity genotypes of the milo group of sorghum (Sorghum bicolor (L.) Moench). The 12-h dark period without GA₃ application or FR induced flowering in only the early genotype; FR hastened initiation in the early genotype, while GA₃ hastened floral initiation in the two intermidiate-flowering genotypes. GA₃ and FR together had a strong synergistic effect, hastening floral initiation by 30 to more than 80 d in the early and intermediate genotypes. Red light (R) did not hasten flowering; FR preceded by R gave the same effect as FR alone. GA₃ promoted stem elongation equally whether floral initiation occurred or not; thus, its effect on stem elongation was independent of floral initiation. The capacity of GA₃ to induce flowering in sorghum, a short-day plant, seems to be enhanced by phytochrome being in the P_R form at the beginning of the night when GA₃ was applied.Abbreviations FR far-red light - GA(s) gibberellin(s) - GA₃ gibberellic acid - R red light     

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.     

Inheritance of Factors Affecting Floral Primordia Initiation in Cestrum; Hybrids of C. elegans and C. nocturnum

Photoperiod patterns of hybrids of Cestrum elegans (Brongn.) Schlect., a day neutral plant, and C. nocturnum L., a long-short day and long day plant, were investigated. Plants of the F₁ generation, F₂ generation, and backcrosses to each parent were tested on short day, long day, continuous light, long-short day and short-long day for floral primordia initiation. The data recorded suggest 2 independent genes or gene groups controlling floral primordia initiation in C. nocturnum, a single dominant gene that is activated by long-short day treatment and a recessive gene or genes responding to long day treatment. Further, these data suggest that the day neutral condition in C. elegans is the result of the series of independent genes or gene groups that respond to various photoperiods, the combination of these genes resulting in floral primordia initiation on all photoperiods.     

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     

Phase Shift in the Circadian Rhythm of Floral Promotion by Far Red Energy in Hordeum vulgare L

Eight-day-old barley seedlings (Hordeum vulgare L. cv. Wintex) were pretreated with a single 24-hour daylight fluorescent photoperiod that was supplemented with sufficient far-red energy (FR) to produce a relative red (R)/FR ratio of 0.5. These plants undergo floral initiation about a week after they are returned to 12-hour daylight fluorescent photoperiods (R/FR ratio, 5.5), but floral development does not begin for an additional 2 weeks. Addition of FR light to a subsequent 12-hour photoperiod decreases the lag period between initiation and development by 10 days without affecting the rate of development. Extending the photoperiod to 24 hours has the same effect on the lag period, but this treatment also increases the rate of development. FR present during the second half of this 24-hour photoperiod only further increases the rate of development. Thus, the presence of FR during the first half of the photoperiod appears to affect the time of onset of floral development, while its presence during the second half of the photoperiod affects the rate of this development.     

Carbon Dioxide and Flowering in Pharbitis nil Choisy

The effects of photoperiod on floral and vegetative development of Pharbitis nil were modified by atmospheric CO₂ concentrations maintained during plant growth. Short day (SD) photoperiods caused rapid flowering in Pharbitis plants growing in 0.03 or 0.1% CO₂, while plants in long day (LD) conditions remained vegetative. At 1 or 5% CO₂, however, flower buds were developed under both the SD and LD photoperiods. Flowering was earliest in the plants exposed to SD at low CO₂ concentrations which formed floral buds at stem node 3 or 4. At high CO₂ concentrations, floral buds did not form until stem node 6 or 7. Both high CO₂ concentrations and LD photoperiods tended to enhance stem elongation and leaf formation.     

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.     

Studies on the Physiology of Flowering of Timothy (Phleutm pratense L.): I. Influence of Daylength and Temperature on Initiation and Differentiation of the Inflorescence

Spikelet initiation is advanced and the proportion of plantswhich attain the reproductive condition is increased in S. 48timothy by lengthening the photo-period from 14 to 24 hours.In shorter periods of light, reproduction is almost completelyinhibited, and in 8-hour short days plants remain vegetativeeven after 35 weeks. Spikelet initiation at the shoot apex occursafter exposure to 3–5 long days followed by short days.Initiation also occurs when extended daylength is replaced by‘light-breaks’ during long nights, or when a singleleaf is photo-induced while the remainder of the plant receivesshort days. High temperatures promote spikelet initiation incontinuous light; in photoperiods nearer the threshold for floweringthis response is reversed and a rise in temperature from 55°to 75° F. increasingly inhibits reproduction. Once initiationhas occurred, spike differentiation is hastened by increasesin temperature or photoperiod. Internode elongation begins atthe time of spikelet initiation, and is promoted by temperatureand photoperiod. Elongated vegetative shoots may be producedwhen spikelet initiation fails in threshold photoperiods orhigh temperatures.     

Genetic Regulation of Development in Sorghum bicolar: I. Role of the Maturity Genes

The photoperiodic behavior and other developmental and morphological differences of 11 maturity genotypes (as identified by JR Quinby 1967, Adv Agron 19: 267-305) of the milo group of Sorghum bicolor (L.) Moench were studied under 8, 10, 12, and 14 hour photoperiods. Sorghum is a quantitative short day plant. The genotypes studied differ in genes which modify photoperiodic behavior and thus maturity; the alleles are designated as Ma(1), ma(1), Ma(2), ma(2), Ma(3), ma(3), and ma(3) (R) (single symbols indicate homozygosity at the indicated gene loci). Based on floral initiation (differentiation) under 10, 12, and 14 hour photoperiods the 11 genotypes were assigned to three clases: (I) flower initiation delayed by 12 hour photoperiods (all genotypes with Ma(1)Ma(2) but not ma(3) (R)), (II) flower initiation delayed by 14 hour photoperiods (all genotypes with Ma(1)ma(2), ma(1)Ma(2), or ma(1)ma(2) but not ma(3) (R)), (III) flower initiation not drastically delayed by 14 hour photoperiods (all genotypes with ma(3) (R)). All of the class III genotypes were taller, had longer leaf sheaths, narrower and longer leaf blades, and less leaf area, than the other genotypes. In addition, the class III genotypes initiated rapid culm and thus internode elongation sooner after floral initiation than any of the class I or II genotypes. Dry weight did not differ between the class III genotypes and the others. The rate of leaf emergence in the class III genotypes and all others was indistinguishable until after floral initiation in the former. The allelic combination unique to class I, Ma(1)Ma(2), makes plants very photoperiod sensitive without causing observable changes in morphology or other development events. The allelic combination unique to class III, ma(3) (R), makes plants relatively photoperiod insensitive and results in several differences in morphology and development.     

Growth responses of Typha orientalis presl to controlled temperatures and photoperiods

Experiments are described in which seedlings of Typha orientalis Presls were grown for up to 6 months under precise conditions of temperature and photoperiod; photosynthesis was by natural daylight and did not vary between treatments. Variable treatments were imposed either from the seedling stage or on large plants raised under constant conditions.In general, total dry matter production increased as photoperiod increased from 8 to 16 h and also as day or night temperature increased, maximum production occurring when there was a warm day (30 or 27°C) and a small temperature drop (to 22°C) at night. The distribution of dry matter was also markedly affected by the imposed variables, leaf growth being favoured by high temperatures (to 30°C) and long photoperiods, and production of roots and rhizomes by low temperatures (to 10°C) and short photoperiods. None of the treatments resulted in floral initiation. The results are considered in relation to growth in the natural habitat.     

Enzyme activity and electrophoretic pattern of isoenzymes of peroxidase,esterase and alkaline and acid phosphatase in relation to flowering inAmaranthus viridis L. - a quantitative SD plant

Amaranthus viridis is a quantitative SD plant in which inflorescence primordia are initiated under both 24- and 8-h photoperiods after 12 and 10 days, when 8 and 7 leaves are differentiated, respectively. Photoperiod plays a non-determinate role, whereas the maturity of plants linked with the attainment of minimum leaf number is significant and of primary importance in floral induction. This is further confirmed by the more or less identical nature of changes in the total enzyme activity and isoenzyme patterns of peroxidase, esterase and alkaline and acid phosphatase under the two photoperiods. These changes occur once the minimum vegetative growth has been achieved prior to the reproductive transformation, irrespective of the photoperiod, pointing to the activation of a general common pathway of events leading to floral induction.     

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; )     

The Effect of Gibberellic Acid and Guanosine Monophosphates on Extension Growth, Leaf Production and Flowering of Impatiens balsamina

Gibberellic acid (GA₃) increases the height of Impatiens balsamina under both 8- and 24-h photoperiods. The height also increases with all guanosine monophosphates (GMPs) under 8-h photoperiods but only with 5′-GMP under 24-h photoperiods. GA₃ as well as GMPs increase the number of leaves under 8-h but not under 24-h photoperiods. GA₃ as well as GMPs induce floral buds under strictly non-inductive photoperiods and increase the number of floral buds under 8-h photoperiods. The floral bud initiation occurs earlier when cGMP is used in combination with 100 mg/l GA₃.     

Effects of gibberellic acid and some diphenols on the flowering of Impatiens balsamina L., a qualitative short day plant

Res, DOPA and CA resemble GA₃ in inducing floral buds in Impatiensbalsamina under strictly non-inductive photoperiods, while Catdoes not do so. 1 mg/liter Res and 100 mg/liter CA in combinationwith 100 mg/liter GA₃ even hastened the initiation of floralbuds. All the tested phenols, in combination with 100 mg/literGA₃, caused a synergistic increase in the number of floral buds. (Received November 24, 1977; )