Effects of Far-red Light on the Hormonal Control of Side Shoot Growth in the Tomato

Side shoot growth in young tomato plants was almost completelysuppressed by a 5 min period of far-red light immediately followinga 16 h photoperiod from fluorescent tubes, whereas plants givenan identical photoperiod but lacking the far-red treatment branchedprofusely. The influence of far-red light on the degree of sideshoot suppression and the correlated changes in the levels ofauxins, gibberellins, cytokinins and abscisic acid is presentedand discussed in relation to current hypotheses of correlativeinhibition. It is suggested that far-red light causes increasedauxin synthesis in the apex and young leaves, which in turninduces the formation of abscisic acid in or near the axillarybuds, and it is this hormone which inhibits bud outgrowth. Therole of cytokinins and gibberellins remains uncertain but theyprobably act in a sequential manner, the gibberellins promotingbud growth following cytokinin-mediated release from apicaldominance.     

Photocontrol of stem elongation in light-grown plants of Fuchsia hybrida

Stems of the caulescent long-day plant, Fuchsia hybrida cv Lord Byron, showed 2 types of response to light. In one, internode length was increased by far-red irradiation given at the end of an 8 h photoperiod: the response was no greater with prolonged exposure and was less when the start of far-red was delayed. The effect of far-red was reversible by a subsequent exposure to red light. Internode length was inversely proportional to the P_fr/P ratio established before entry to darkness and there was no evidence for loss of P_fr during a 16 h dark period. The inhibitory effect of P_fr acted at a relatively late stage of internode growth. With the development of successive internodes a second response appeared in which stems lengthened following prolonged daily exposures to red or far-red light, or mixtures of the two, or to brief breaks with red or white light. In these later internodes, a short exposure to far-red near the middle of the night was not reversible by red because red alone promoted elongation at this time. Internode length increased with increase in the daily duration of light and, when light was given throughout an otherwise dark period of 16 h, with increase in illuminance to a saturation value of 200 lx from tungsten lamps. Elongation increased as a linear function of decrease in photostationary state of phytochrome down to P_fr/P0.3; however, internodes were shorter in far-red light than in 25% red/red+far-red. It was concluded that stem length is a net response to two modes of phytochrome action. An inductive effect of P_fr inhibits a late stage in internode expansion, and a phytochrome reaction which operates only in light (and may involve pigment cycling) promotes an early stage of internode development. Stem elongation is thus a function both of the daily duration of light and its red/red+far-red content. The outgrowth of axillary buds was controlled by the first type of phytochrome action only.Abbreviations and symbols FR far red light - R red light - P phytochrome - P_fr phytochrome in the far-red light absorbing form - SD 8 h short days - LDP long-day plant - SDP short-day plant     

Effects of photoperiod on some vegetable species

Seven vegetable species grown in controlled environments were given similar daily amounts of visible radiation (500 J cm^-2) during three different photoperiod treatments. Plants were given (A) 12 h of visible irradiance (115 W m^-2) from fluorescent and tungsten lamps, (B) 16 h of the same light at 88 W m^-2 or (C) Treatment A extended to 16 h with 4 h of low-intensity incandescent light (3 W m?²) from tungsten lamps only. All seven species grew faster when daylength was extended with light of photosynthetic intensity (B), probably through increased net assimilation rates, and leaf area of these plants increased in proportion to change in plant dry weight. Daylength extension with a mixture of red and far-red light (C) induced photomorphogenic changes in specific leaf area in all species examined and increased leaf area and plant dry weight of lettuce, celery, beetroot and spinach beet but not of three members of the Cruciferae (radish, cabbage and oilseed rape).     

Photophysiology of Kalanchoë Seed Germination

Germination of Kalanchoé blossfeldiana seeds is absolutely light-requiring and needs repeated daily light periods. With increasing length of the photoperiod there was a gradual escape from the far-red inhibition. This escape depended also upon the duration of the far-red exposure: 10-second far-red caused a strong inhibition after a 10- to 30-minute photoperiod and did not inhibit after a 4-hour day, although the effect of the latter was completely suppressed by 5 minutes far-red. The action of a 12-hour photoperiod was not reversed by 10 minutes far-red but it was by 12 hours far-red. Light intensity and temperature during the photoperiod were two other important factors influencing the escape from far-red inhibition. The common features of this escape displayed in very different photomorphological responses are stressed. In order to explain our results in terms of phytochrome action, we distinguish two effects of white light: 1) on the initial photoconversion of the inactive to the active P_FR form 2) on the much slower transformation of P_FR to a reacted form P*_FR; the latter reaction can also proceed in darkness, but is enhanced by light and is dependent upon light intensity and temperature; this reacted phytochrome is not reversible by a brief far-red illumination.     

Effects of light quality on somatic embryogenesis in Araujia sericifera

The effects of photoperiod, light quality and end-of-day (EOD) phytochrome photoconversion on somatic embryogenesis (SE) of Araujia sericifera petals have been studied. Petals from immature flowers were cultured under 8- and 16-h photoperiods using Gro-lux fluorescent lamps. The photon fluence rate was 90–100 μmol m⁻² s⁻¹ and the red (R):far-red (FR) ratio was 98. R, FR, R followed by FR (R-FR) and FR followed by R (FR-R) light treatments were applied for 3 weeks at the end of the photoperiods. In a set of experiments, dl - α -difluoromethylarginine (DFMA) or methylglyoxal bis (guanylhydrazone) (MGBG), both inhibitors of polyamine biosynthesis, were added to the culture medium in order to study the involvement of polyamine metabolism. The level of SE was the same in long (LD) and short (SD) days. Thus, the light effect was accomplished after 8 h. All EOD treatments that decreased the P_fr level inhibited SE when applied after SD, but not after LD. The FR-R treatment after LD caused an additional stimulatory effect on SE, even in the presence of polyamine inhibitors. DFMA inhibited SE in both SD and LD, but MGBG did not modify SE in either SD or LD. The R, FR and R-FR treatments did not alter the level of SE when applied after LD in the presence of DFMA or MGBG. However, these treatments decreased SE after SD when the medium contained polyamine inhibitors. Our results suggest that Gro-lux lamps, which produce an extremely high R:FR ratio, promote SE in A. sericifera and a timing response to phytochrome photoconversion during photoperiodic induction. Thus, our data corroborate the involvement of phytochromes and polyamines in SE in A. sericifera, which responded as a light-dominant long-day plant.     

Effects of cytokinins and injury on the formation of shoot buds by leaves of Dendrophthoe falcata

Summary A number of cytokinins induce shoot buds on leaves obtained from embryonal tissue (diploid) or endosperm tissue (triploid) of Dendrophthoe falcata cultured in vitro. The buds develop either by division of an epidermal cell which finally is organized into a shoot meristem; or the epidermal cell first produces a callus which subsequently gives rise to shoot buds. Buds do not develop in the absence of cytokinins. Injury to the leaf plays a major role in the distribution and number of shoot buds formed. Normal leaves on the plant do not regenerate buds even in the presence of a cytokinin.     

Apical Dominanace in Xanthium strumarium: A DISCUSSION IN RELATION TO CURRENT HYPOTHESES OF CORRELATIVE INHIBITION

The influence of the spectral distribution of illumination onthe gibberellin, cytokinin, auxin, and abscisic acid levelsand the correlation with the degree of branching in Xanthiumstrumarium is presented and discussed. Gibberellins do not appearto play a major role in apical dominance but may be importantfor bud extension following the initial release from dominance.The cytokinin level was much higher in inhibited buds than inreleased buds. It is suggested that the cytokinins present wereprobably not able to participate in bud growth because of anauxin-induced accumulation of abscisic acid in the buds themselves.The concentration of abscisic acid as measured by bioassay andgas-liquid chromatography was between 50 and 250 times thatoccurring in all other plants parts examined. This level felldramatically following release from apical dominance by decapitation.The results are discussed in relation to current hypothesesof apical dominance.     

Axillary Bud Formation in Two Isogenic Lines of Tomato Showing Different Degrees of Apical Dominance

Grafting experiments have been carried out in which rootstocksof the cultivar Craigella were paired with scions of an isogenicline Craigella Lateral Suppressor (ls ls) and vice versa, andthe levels of hormones in the roots and shoots of the graftedplants examined. The roots of Craigella plants differed from those of LateralSuppressor in that they contained a higher proportion of a cytokininthat co-chromatographed with N⁶ - (²—isopentenyl) adenosine.Reciprocal grafts did not lead to any qualitative or quantitativechanges in the cytokinins in the roots of either line. GraftingLateral Suppressor scions on Craigella rootstocks led to anincrease in the IAA content of the apical region and the ABAcontent of the stem tissue immediately below it, but when Craigellascions were grafted on Lateral Suppressor rootstocks there wereno changes in the level of either hormone. Cytokinins applied to the leaf axils of Lateral Suppressor plantsresulted in lateral bud initiation in the axils above the pointof treatment but not if the plants were also given a short periodof far-red light at the end of the photoperiod. Cytokinins wereineffective in initiating lateral buds in grafted Lateral Suppressorscions. It is suggested that root-produced cytokinins influence lateralbud outgrowth indirectly by way of their effect on the levelsof IAA and ABA in the shoot. Lycopersicon esculentum Mill., tomato, apical dominance, growth regulation, indol-3yl acetic acid, abscisic acid, cytokinins     

The influence of high levels of brief or prolonged supplementary far-red illumination during growth on the photosynthetic characteristics, composition and morphology of Pisum sativum chloroplasts

Abstract. Peas were grown in controlled environments (12h white fluorescent light. ∼47 μmol photons m^-2 s 1/12 dark, 25 °C), using (1) 15-min far-red illumination at the end of each photoperiod (brief FR) to simulate the increase in the far-red/red ratio near the end of the day, and (2) high levels of supplementary far-red light (red:far-red ratio=0.04) during the entire photoperiod (long-term FR) to simulate extreme shade conditions under a plant canopy. Brief FR illumination led to marked morphological effects attributable to phytochrome regulation, namely, an increase in internodal length, but a decrease in leaflet area, chloroplast size and chlorophyll content per chloroplast compared with the control. Significantly, brief FR illumination had little or no effect on the amounts of the major chloroplast components (ribulose 1.5-biphosphate carboxylase, adenosine triphosphate synthase, cytochrome b/f complex and Photosystem II) relative to chlorophyll or Photosystem I, and the leaf photosynthetic capacities per unit chlorophyll were similar. In contrast, supplementing high levels of far-red light during the entire photoperiod not only led to the phytochrome effects above, but there was also a marked increase in leaf photosynthetic capacity per unit chlorophyll. due to increased amounts of the major chloroplast components relative to chlorophyll or Photosystem I. We hypothesize that supplementary far-red light, absorbed by Photosystem I, induced an increase in the major chloroplast components by a photosynthetic feedback mechanism. In fully greened leaves, we propose that the two photosystems themselves, rather than phytochrome, may be the predominent sensors of light quantity in triggering modulations of the stoichiometries of chloroplast components, which in turn lead to varying photosynthetic capacities.     

Photocontrol of petiole elongation in light-grown strawberry plants

Summary The mode of phytochrome control of elongation growth was studied in fully-green strawberry (Fragaria x Ananassa Duch.) plants. Petiole growth showed two distinct types of response to light. In one, the end-of-day response, petioles were lengthened by low-intensity far-red irradiation for 1 h immediately following the 8 h photoperiod. The response was little or no greater with prolonged exposure and less when the start of far-red was delayed. It was already evident in the first leaf to emerge after treatment began. With the development of successive leaves a second, photoperiodic, type of response appeared, in which petioles lengthened following only prolonged exposure to red, far-red, mixtures of the two, or tungsten lighting, all at low levels of intensity. As with the inhibition of flowering in previous experiments, irradiation with red light during the second half of the otherwise long dark period gave the greatest response.Abbreviations and Symbols FR far-red light - HIR high irradiance response - R red light - P_r phytochrome in the red light absorbing form - P_fr phytochrome in the far-red light absorbing form - SDP short-day plant - LDP long-day plant - PAR photosynthetically active radiation     

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)     

Phytochrome controlled C-sucrose uptake into etiolated pea buds: effects of gibberellic Acid and other substances

The previously reported red light enhancement of ¹⁴C-sucrose uptake into etiolated pea buds is inhibited by gibberellic acid applied no later than 2 hours after the light. Auxins, cytokinins and inhibitors of gibberellin biosynthesis are without effect, either alone or in the presence of gibberellic acid.     

Effect of Photoperiod on Growth of Sugar Beet

Sugar beet grown in controlled environments were given similardaily amounts of visible radiation during three different photoperiodtreatments. Plants were given (a) 115 W m^–2 visible irradiancefrom fluorescent and tungsten lamps for 12 h; (b) 88 W m^–2of the same light for 16 h or (c) 115 W m^–2 from fluorescentand tungsten lamps for 12 h extended to 16 h with low intensity(3 W m^–2) incandescent light from the tungsten lamps only.Plant growth was increased similarly in both long day treatments[(b) and (c)] and dry weights were 25 per cent greater thanin the 12 h photoperiod (a) after 6 weeks. Leaf area was increasedby 18 per cent and net assimilation rate by 10 per cent in the16 h photoperiod at 88 W ^m–2 (b). By contrast, extendingthe photoperiod with 4 h of incandescent light (c) triggereda photomorphogenic increase in leaf expansion which increasedleaf area per plant by 47 per cent and leaf-area ratio by 12per cent.     

Involvement of cytokinin,ABA and endogenous inhibitors in sprouting of basal buds in rose plants

Sprouting of basal buds in rose plants with a completely removed canopy was restrained by inhibitors in a water soluble fraction of an ethanolic extract from the bark tissue of the plant stump (the crown) extracted before the top removal. The inhibitory effect of this fraction disappeared if the bark was extracted one week after the removal of the plant canopy. A similar inhibition was obtained with extracts from stem segments. The organic fraction did not show any inhibitory activity. Both fractions, aqueous and organic, were inhitory in the lettuce hypocotyl elongation bioassay. High levels of ABA in the cis-configuration were found in extracts before the top removal. The amount of cis-ABA decreased in extracts made one week after top removal and a different peak of activity, probably corresponding to trans-ABA or phaseic acid was shown in the GLC-system used.A high content of cytokinins has been found in the bark tissue of the crown. There were qualitative differences in cytokinin activity between the tissue from plants after the removal of the canopy and from plants with main branches left intact. A balance between cytokinins and inhibitory complex was proposed to be involved in the regulation of basal buds outgrowth and plant regeneration.     

The Role of Light in Leaf and Flower Bud Break of the Peach (Prunus persica)

The effect of light on peach leaf and flower bud break was examined. It was found that leafless dormant shoots were light-perceptive organs. Darkness, after light preconditioning during dormancy, reduced leaf bud opening; however, light was obligatory when the shoots were preconditioned in the dark. Relatively short exposures to light were sufficient to stimulate leaf bud break. Terminal buds were less inhibited by darkness than were laterals. Flower bud break was inhibited in light after dark preconditioning. The red region of the spectrum was found to be active; the phytochorome system seems to be involved in the light reactions, as the red light effect was reversible with subsequent far-red illumination. Supplementary light, producing long-day conditions, could partly compensate for insufficient chilling. A possible sequence of reactions in the plant is suggested.     

Direct shoot bud induction and plant regeneration in <Emphasis Type="Italic">Capsicum frutescens</Emphasis> Mill.: influence of polyamines and polarity

Direct shoot bud induction and plant regeneration was achieved in Capsicum frutescens var. KTOC. Aseptically grown seedling explants devoid of roots, apical meristem and cotyledons were inoculated in an inverted position in medium comprising of Murashige and Skoog (Physiol Plant 15:472–497, 1962) basal medium supplemented with 2-(N-morpholine) ethanesulphonic acid buffer along with 2.28 μM indole-3-acetic acid, 10 μM silver nitrate and either of 13.31–89.77 μM benzyl adenine (BA), 9.29–23.23 μM kinetin, 0.91–9.12 μM zeatin, 2.46–9.84 μM 2-isopentenyl adenine. Profuse shoot bud induction was observed only in explants grown on a media supplemented with BA (26.63 μM) as a cytokinin source and 19.4 ± 4.2 shoot buds per explant was obtained in inverted mode under continuous light. Incorporation of polyamine inhibitors in the culture medium completely inhibited shoothoot bud induction. Incorporation of exogenous polyamines improved the induction of shoot buds under 24 h photoperiod. These buds were elongated in MS medium containing 2.8 μM gibberellic acid. Transfer of these shoots to hormone-free MS medium resulted in rooting and rooted plants were transferred to fields. This protocol can be efficiently used for mass propagation and presumably also for regeneration of genetically transformed C. frutescens.     

Comparison of endogenous cytokinins, ABA and metabolites during female cone bud differentiation in low and high cone-producing genotypes of lodgepole pine

In lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.), cone bud initiation within long-shoot buds varies according to genotype. We chose to study hormone profiles of two genotypes that differed significantly in cone yield. Phytohormone profiles were established by high performance liquid chromatography–electrospray ionization tandem mass spectrometry in multiple reaction monitoring mode with samples collected from genotypes 299 and 233, the typically high and low cone producers. Generally, concentrations of trans-zeatin-O-glucoside were higher in genotype 299, whereas dihydrozeatin concentrations were higher in genotype 233. Both isopentenyl adenine and isopentenyl adenosine were present at higher concentrations in genotype 233. The ratio of total quantifiable zeatin (Z)-type cytokinins to isopentenyl (iP)-type cytokinins was approximately threefold higher in genotype 299 during female cone bud differentiation. In genotype 299, ABA concentration was significantly lower than in genotype 233 on the first sampling date, while the phaseic acid concentration was lower consistently throughout the period investigated. Dihydrophaseic acid was present in low concentrations in most samples of genotype 233, but was not quantifiable in genotype 299. Our study reveals that long-shoot buds of the high cone-producing genotype had higher ratios of Z-type cytokinins to iP-type cytokinins than were found in the low cone-producing genotype. High cone-producing buds also contained less ABA, phaseic acid and dihydrophaseic acid during female cone bud differentiation.     

Changes in endogenous level of auxins and cytokinins in axillary buds ofpisutn sativum L. in relation to apical dominance

Decapitation of the stem in one-week-old pea seedlings below the first node causes a rapid outgrowth of the two cotyledonary buds. One of them soon becomes dominant, while the other one is inhibited, but can be released from inhibition by cutting off the dominant bud. The level of endogenous auxins and cytokinins was determined in dominant and inhibited buds, as well as in released buds at different time intervals after deinhibition. It was found that the inhibited buds contained very little acidic, ether soluble auxins, a high level of tryptophan and also a high level of cytokinins, in comparison with dominant buds. When the inhibited buda were released from inhibition, their auxin content rose, while that of tryptophan and cytokinins decreased, reaching the level found in dominant buds within six days. Specific changes in content of two undetermined auxin-like substances were found in released buds during de-inhibition. These results are discussed in relation to the current views on the regulation of apical dominance.     

Photocontrol of seed germination in the hemiparasite Buchnera hispida (Scrophulariaceae)

Abstract Buchnera hispida, a facultative root parasite of grasses and graminaceous crops, has a light requirement for germination. Studies were carried out on the effects of varying photoperiods with or without preceding dark incubation, on seed germination. Buchnera seeds showed long-day behaviour, since they germinated at all photoperiods including continuous light, and longer photoperiods were more effective in triggering seed germination than shorter photoperiods. Also, effects of red and far-red light indicated that the phytochrome system is operative in the light-induced germination of Buchnera. Although dark incubation in water before illumination was not absolutely necessary for germination, it caused the seeds to respond more rapidly to light. The longer the time of the dark incubation the more responsive the seeds were to photoperiod except when 15 min light was given. The effectiveness of a preceding dark incubation in making Buchnera seeds sensitive to rapid light action was completely inhibited at 4°C. This is in agreement with the hypothesis that a reaction partner of the far-red absorbing form of phytochrome is produced during dark incubation of Buchnera seeds. Such an intermediate has also been reported in some positively photoblastic seeds of non-parasitic flowering plants.     

Phytochromes A1 and B1 have distinct functions in the photoperiodic control of flowering in the obligate long-day plant Nicotiana sylvestris

The obligate long-day plant Nicotiana sylvestris with a nominal critical day length of 12 h was used to dissect the roles of two major phytochromes (phyA1 and phyB1) in the photoperiodic control of flowering using transgenic plants under-expressing PHYA1 (SUA2), over-expressing PHYB1 (SOB36), or cosuppressing the PHYB1 gene (SCB35). When tungsten filament lamps were used to extend an 8 h main photoperiod, SCB35 and SOB36 flowered earlier and later, respectively, than wild-type plants, while flowering was greatly delayed in SUA2. These results are consistent with those obtained with other long-day plants in that phyB has a negative role in the control of flowering, while phyA is required for sensing day-length extensions. However, evidence was obtained for a positive role for PHYB1 in the control of flowering. Firstly, transgenic plants under-expressing both PHYA1 and PHYB1 exhibited extreme insensitivity to day-length extensions. Secondly, flowering in SCB35 was completely repressed under 8 h extensions with far-red-deficient light from fluorescent lamps. This indicates that the dual requirement for both far-red and red for maximum floral induction is mediated by an interaction between phyA1 and phyB1. In addition, a diurnal periodicity to the sensitivity of both negative and positive light signals was observed. This is consistent with existing models in which photoperiodic time measurement is not based on the actual measurement of the duration of either the light or dark period, but rather the coincidence of endogenous rhythms of sensitivity - both positive and negative - and the presence of light cues.