Photoregulation of seed germination of wild-type and of an aurea-mutant of tomato

Seed germination of an aurea mutant of tomato ( Lycopersicon esculentum Mill.) is promoted by continuous irradiation with red, far-red or long-wavelength far-red (758 nm) light as well as by cyclic irradiations (5 min red or 5 min far-red/25 min darkness). Far-red light applied immediately after each red does not change the germination behaviour. Seed germination of the isogenic wild-type, cv. UC-105, is promoted by continuous and cyclic red light while it is inhibited by continuous and cyclic far-red light and by continious 758 nm irradiation. Far-red irradiation reverses almost completely the promoting effect of red light. The promoting effect (in the aurea mutant) and the inhibitory effect (in the wild-type) of continuous far-red light do not show photon fluence rate dependency above 20 nmol m⁻² s⁻¹. It is concluded that phytochrome controls tomato seed germination throgh low energy responses in both the wild type and the au mutant. The promoting effect of continuous and cyclic far-red light in the au mutant can be attributed to a greater sensitivity to P_fr.     

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.     

An action spectrum in the blue for inhibition of germination of lettuce seed

Summary Using a 2-h irradiation period at constant quantum irradiance, a complete action spectrum for inhibition of germination of lettuce seed has been obtained. Action maxima were near 470 and 720 nm, the latter being the most active wavelength. It was also shown, under conditions where light inhibition cannot occur, that phytochrome potentiation of germination is maximal at all wavelengths below 700 nm, including the highly active blue region. Evidence was presented for promotion of germination by a 2-h irradiation in the red which cannot be explained on the basis of conversion of phytochrome to the active form.Abbreviations Bl blue - FR far-red, P_FR far-red-absorbing form of phytochrome - R red Supported in part by funds provided for biological and medical research by the State of Washington Initiative Measure No. 171 and the Graduate School Research Funds.     

Synergism between gibberellic acid and low Pfr levels inducing germination of Kalanchoë seeds

Sown on water, seeds of Kalanchoëbiossfetdiana Poelln. cv. Feuerblute are absolutely light-requiring and show full red/far-red reversibility. In seeds, sown on 2 ×10^-3 M gibberellic acid, red/far-red reversibility disappears and both short red and far-red irradiations induce germination. Gibberellic acid alone does not induce germination, but it increases the physiological activity of P_fr to the extent, that the low P_fr level obtained by far-red irradiation becomes very effective. The synergism between gibberellic acid and far-red light appears after a two-day incubation; period. The nature of this lag phase was examined by measuring both germination and uptake of labelled gibberellic acid in intact seeds and seeds with a punctured seed coat. The lag phase was shown to be independent of the uptake kinetics of gibberellic acid and allows development to a specific stage, necessary for germination after phytochrome-phototransformation. The kinetics of the uptake of gibberellic acid by intact seeds and embryos of intact seeds are different. In intact seeds most of the gibberellic acid is retained in the seed coat; only a small fraction actually penetrates to the embryo where it can exert its physiological activity.     

Photoregulation of germination in seed of transgenic lines of tobacco and Arabidopsis which express an introduced cDNA encoding phytochrome A or phytochrome B

Photoinduction and photoinhibition of germination in seed from a homozygous tobacco (Nicotiana tabacum L.) line containing an introduced oat phyA cDNA (encoding phytochrome A) is compared with that of isogenic wild-type (WT) tobacco. Under continuous irradiation by a light source with a low redfar-red (RFR) ratio the transgenic tobacco seed appeared to be less susceptible to photoinhibition of germination compared with WT seed. However, induction of germination following a short pulse by R (666 nm) was not enhanced in the genotype transformed by oat phyA cDNA compared with the WT; neither did germination of the transgenic tobacco seed show an increased sensitivity to saturating pulses of light of longer wavelengths (666–730 nm). In seeds of transgenic Arabidopsis thaliana (L.) Heynh. which contained an introduced phytochrome-B-encoding cDNA, levels of dark germination were enhanced, consistent with mediation of response by phytochrome B-P_fr. The germination behaviour of Arabidopsis genotypes wich contained an introduced cDNA encoding phytochrome A, however, did not significantly differ from that of the WT.Abbreviations ABO seed transformed with Arabidopsis phyB - cDNA; CaMV cauliflower mosaic virus - FR far-red light - P_fr far-red-absorbing form of phytochrome - P_tot total phytochrome - P_fr/P_tot phytochrome photoequilibrium - R red light - RBO seed transformed with rice phyB cDNA - RFR quantum ratio of red and far-red light - WL white light - WL + FR whitelight supplemented with far-red light - WT wild type The authors wish to thank R.D. Vierstra (Department of Horticulture, University of Wisconsin-Madison, USA) for providing the transgenic tobacco line, and M.T. Boylan, D. Wagner and P.H. Quail (U.C. Berkeley/USDA Plant Gene Expression Center, Albany, Calif. USA) for providing the transgenic Arabidopsis lines. The work presented in this paper was funded by grants from the Agricultural and Food Research Council (H.S., A.C.M., G.C.W.).     

Growth and Dormancy in Lunularia cruciata (L.) Dum: VI. GROWTH REGULATION BY DAYLENGTH, BY RED, FAR-RED, AND BLUE LIGHT, AND BY APPLIED GROWTH REGULATORS AND CHELATING AGENTS

Experiments with photoperiods ranging from 2 to 24 h confirmthat 8 h light per day is optimal for Lunularia: there is nogrowth in the dark or in continuous light, which causes therapid onset of dormancy. Short-day cycles intercalated amonga series of continuous light cycles promote growth; in cycleslonger than 24 h very long dark periods are detrimental. Withvery short photoperiods (5 min) red light promotes growth moreeffectively than white light at higher intensity; far-red actsas dark. The growth effects of red and far-red light breaks(3 min) depended on the time of application; red light inhibitedin the middle but promoted at the beginning of the 16-h darkperiod of a short day; far-red light had the opposite effect;in each case red and far-red effects were reversible by theother wavelength. Blue light gave the same response as red includingthe reversibility of far-red effects and vice versa. Surprisingly,significant effects of 5 min red, blue, and far-red irradiationwere also found in the middle of the main high-intensity white-lightperiod, red and blue promoting growth, far-red reducing it;again there was ready reversibility of the effects. Growth promoters of higher plants are generally inhibitory toLunularia or have little effect; among growth retardants TIBA,Phosphon D, and CCC gave a slight promotion of growth. EDTApromoted growth (cell numbers) very significantly while 8-hydroxyquinolinewas initially inhibitory, but had a marked latent promotingeffect when subsequently washed from the thalli.     

Antagonistic action of low-fluence and high-irradiance modes of response of phytochrome on germination and beta-mannanase activity in Datura ferox seeds

Seed germination is often induced by a pulse of red light perceived by phytochrome and cancelled by a subsequent pulse of far-red light. When the pulse of red light is followed by several hours of darkness, a pulse of far-red light is no longer effective and prolonged far-red is necessary to block germination. The aim was to investigate whether the red light pulse and prolonged far-red light act on the same or different processes during germination of Datura ferox seeds. Forty-five hours after the inductive red light pulse, germination could not be blocked by one pulse or six hourly pulses of far-red light, but was significantly reduced by 6 h of continuous far-red light. The pulse of red light increased embryo growth potential and the activities of beta-mannanase and beta-mannosidase extracted from the micropylar region of the endosperm. Continuous far-red light had no effect on embryo growth potential or beta-mannosidase activity, but severely reduced the activity of beta-mannanase. The effect of far-red light had the features of a high-irradiance response of phytochrome. Both germination and beta-mannanase activity were restored by a pulse of red light given after the end of the continuous far-red treatment. It is concluded that the low-fluence response and the high-irradiance response modes of phytochrome have antagonistic effects on seed germination and that the control of beta-mannanase activity is one process where this antagonism is established.     

Germination characteristics of some plant species from calcareous fens in southern Germany and their implications for the seed bank

Seeds of 25 plant species from calcareous fen hay meadows were exposed to different experimental conditions and their germination was characterised. Constant temperature inhibited germination especially in Cyperaceae . Both gibberellic acid and potassium nitrate failed to terminate dormancy. Increased germination rates were found in dicot species after treatment with gibberellic acid. Temperature fluctuations increased germination of Cyperaceae as well as dicotyledons. Treatment with gibberellic acid removed the chilling requirement in some of the species. Dormancy of small seeds with thin seed coats was broken by the application of gibberellic acid or fluctuating temperature; large thick-coated seeds were unaffected by gibberellic acid. No obligatory darkness requirement was found in any species; three species germinated irrespective of light treatments. All other species achieved higher percentage germination in daylight or in red (670 nm) light. Permanent darkness and far-red light (730 nm) reduced germination drastically. The results indicate that germination characteristics of the species investigated can be related to their seed bank types.     

Photocontrol of spore germination in the fern Thelypteris kunthii

The light requirement for germination in spores of the fern Thelypteris kunthii (Desv.) Morton was fully satisfied by a long period of continuous red light or partially by intermittent, short periods of red light. Red light-potentiated spore germination was inhibited by brief far-red light irradiation, indicating phytochrome involvement. Repeated exposure of spores to prolonged red and short far-red irradiations, or exposure of red-potentiated spores to far-red light after an extended period in darkness, led to their escape from inhibition of germination by far-red light. Prolonged irradiation of spores with blue light before or after red light treatment partially antagonized the effect of red light.     

Phytochrome Control of Turion Formation in Spirodela polyrhiza L. Schleiden

Turion yield in Spirodela polyrhiza, strain SJ, is increasedby increasing the daily light period. This effect is more pronouncedin autotrophic than in mixotrophic conditions. Night-break irradiation(15 mins) increased turion yield by 150 % under the conditionsof an 8-h daily light period. Besides the effect of night-breakirradiation, end-of-day far-red irradiation decreased turionyield with increasing photoperiod, whereas end-of-day red irradiationwas without any effect. This demonstrates the promoting effectof the P_fr form of phytochrome on formation of light-grown turions. Formation of dark-grown turions was increased by about 240%by a single red light pulse and was reversed by an immediatelyapplied far-red light pulse. Consequently, under heterotrophicconditions phytochrome modulates the turion formation process. Spirodela polyrhiza L. Schleiden, duckweed, Lemnaceae, photomorphogenesis, phytochrome, turion     

Mediation of phytochrome in the inductive action of low temperature on dark germination of lettuce seed at supra-optimal temperature

The induction of dark germination in light-requiring lettuce (Lactuca sativa) seed at supraoptimal temperatures by cold treatment (in darkness) was partly reversed by a brief far-red irradiation made at time of transfer, and even more so when the irradiation was made at the beginning of the cold pretreatment. When the inhibitory far-red irradiation was followed by additional cold treatment, the promotion was greatly restored. The promotive effects of brief irradiations with red light were further enhanced by a following cold period, before transfer to the supraoptimal temperature. These results are interpreted as indicating that the active (far-red absorbing) form of phytochrome is pre-existing in the dry seed, and interacts with a co-factor which is built-up during imbibition. The rate of build-up of this co-factor, as well as of the dark inactivation of active phytochrome increase with temperature. The products of the interaction pass through a photo-labile thermo-stable phase, before becoming photo-stable as well.     

Photoresponses of Light-Grown phyA Mutants of Arabidopsis (Phytochrome A Is Required for the Perception of Daylength Extensions)

Several aspects of the photophysiology of wild-type Arabidopsis thaliana seedlings were compared with those of a phytochrome A null mutant, phyA-1, and a mutant, fhy1, that is putatively involved in the transduction of light signals from phytochrome A. Although phyA seedlings display a near wild-type phenotype when grown in white light (W), they nevertheless display several photomorphogenic abnormalities. Thus, whereas the germination of wild-type and fhy1 seeds is almost fully promoted by a pulse of red light (R) or by continuous far-red light (FR), phyA seed germination is responsive only to R. Following growth under day/night cycles, but not under continuous W, the hypocotyls of light-grown phyA and fhy1 seedlings are more elongated than those of wild-type seedlings. For seedlings grown under low red/far-red (R/FR) ratio light conditions, phyA and fhy1 seedlings display a more marked promotion of hypocotyl elongation than wild-type seedlings. Similarly, seedlings that are doubly null for phytochrome A and phytochrome B(phyA phyB) also have more elongated hypocotyls under low R/FR ratio conditions than phyB seedlings. This indicates that phytochrome A action in light-grown seedlings is antagonistic to the action of phytochrome B. Although wild-type, fhy1, and phyA seedlings flower at essentially the same time under both short-day and long-day conditions, an obvious consequence of phytochrome A deficiency is a pronounced late flowering under conditions where a short day of 8 h of fluorescent W is extended by 8 h of low-fluence-rate incandescent light. The evidence thus indicates that phytochrome A plays a role in seed germination, in the control of elongation growth of light-grown seedlings, and in the perception of daylength.     

Phytochrome regulation of seed germination

Seed germination of many plant species is influenced by light. Of the various photoreceptor systems, phytochrome plays an especially important role in seed germination. The existence of at least five phytochrome genes has led to the proposal that different members of the family have different roles in the photoregulation of seed germination. Physiological analysis of seed germination ofArabidopsis thaliana (L.) Heynh. with phytochrome-deficient mutants showed for the first time that phytochrome A and phytochrome B modulate the timing of seed germination in distinct actions. Phytochrome A photo-irreversibly triggers the photoinduction of seed germination after irradiation with extremely low fluence light in a wide range of wavelengths, from UV-A, to visible, to far-red. In contrast, phytochrome B mediates the well-characterized photoreversible reaction, responding to red and far-red light of fluences four orders of magnitude higher than those to which PhyA responds. Wild plants, such asA. thaliana, survive under ground as dormant seeds for long periods, and the timing of seed germination is crucial for optimizing growth and reproduction. It therefore seems reasonable for plants to possess at least two different physiological systems for sensing the light environment over a wide spectral range with exquisite sensitivity of different phytochromes. This redundancy seems to enhance plant survival in a fluctuating environment.     

Rhizome transition to storage organ is under phytochrome control in lotus <Emphasis Type="Italic">(Nelumbo nucifera)</Emphasis>

We examined photoperiodic response of lotus (Nelumbo nucifera) rhizome morphogenesis (its transition to a storage organ) by using seed-derived plants. Rhizome enlargement (increase in girth) was brought about under 8, 10 and 12 h photoperiods, whereas the rhizomes elongated under 13 and 14 h photoperiods. Rhizomes elongated under 14 h light regimes supplied as 8 h of natural light plus 6 h supplemental hours of white, yellow or red light, but similar treatments with supplemental blue, green or far red light, caused enlargement in girth of the rhizomes. A 2 h interruption of the night with white, yellow or red light, in plants entrained to 8 h photoperiod brought rhizome elongation, whereas 2 h-blue, green or far red light night breaks still resulted in rhizome increase in girth. The inhibitory effect of a red (R) light night break on rhizome increase in girth was reversed by a far-red (FR) light given immediately afterwards. Irradiation with R/FR/R inhibited the rhizome increase in girth. FR light irradiation following R/FR/R irradiation cancelled the effect of the last R light irradiation. It was demonstrated that the critical photoperiod for rhizome transition to storage organ is between 12 and 13 h photoperiod. It was also evident that the optimal light quality range for interruption of dark period (night break) is between yellow and red light and that a R/FR reversible reaction is observed. From these results, we propose that phytochrome plays an important role in photoperiodic response of rhizome increase in girth in lotus. This is the first report on phytochrome-dependent morphogenesis of storage organs in rhizomous plants.     

Phytochrome-mediated germination control of maize caryopses

Maize caryopses sown in water germinate equally well either in darkness or under any light regime. However, when they are imbibed in mannitol solutions, continuous far-red light proves to be strongly inhibitory on the final germination as compared to darkness. Similar but less pronounced inhibition is also exhibited by continuous red or blue light. Intermittent far-red light can partially substitute for continuous far-red light in inhibiting maize caryopsis germination, and its effect is reversed to the intermittent red light level when red light is given immediately after each far-red illumination. These results are interpreted as a proof of existence and involvement of phytochrome in the germination control of maize caryopses, though its manifestation is realized only under osmotic stress.Abbreviations D darkness - FR far-red - R red - B blue - c-FR, c-R, c-B continuous FR, R, B, resp. - i-FR, i-R intermittent FR, R, resp.     

Wavelength dependence of suppression of potato sprout growth by light

Abstract Light is used commercially to prevent the sprouts on seed potatoes from growing so long that they are knocked off at planting. We investigated the relationship between wavelength and growth inhibition, using a very long irradiation time of 23 d with a 12-h photoperiod. The inhibition showed a narrow peak of far-red activity centered on 707 nm, with a shoulder in the red whose size depended on the degree of inhibition chosen as standard, because the log photon-fluence rate/response lines were not parallel. There was also inhibitory activity in the blue (< 500 nm). In many respects the wavelength relationship resembled the action spectra for growth inhibition in dark-grown seedlings, but in the latter the red peak is, or quickly becomes, more pronounced than the far-red peak: this did not happen with potato sprouts. Blue light caused a positive phototropic response at similar or lower fluence rates. Greening became visible only at the highest fluence rates, between the two spectral regions inhibitory to growth. Broadband sources had much less inhibitory activity in the 650–750-nm region.     

ACTION OF KINETIN ON PHOTOSENSITIVE GERMINATION OF LETTUCE SEED AS COMPARED WITH THAT OF GIBBERELLIC ACID

1. Experiments with the seeds of Grand Rapids lettuce showedthat the germination induced by gibberellic acid or by red lightis strongly accelerated by kinetin, although the latter itselfcan promote the germination in the dark only slightly. The reversiblelight reactions of the phytochrome system interact with kinetintreatment just as effectively as with water-imbibed controls.The site of primary action of red light is not altered by kinetintreatment. Kinetin does not modify the water uptake of the seedsfor at least 8 hours. 2. Kinetin was found to inhibit the growth of the hypocotyland root of the seed, but to promote very markedly the expansionof the cotyledons. This effect was observed not only with cotyledonsin intact seeds but also with isolated cotyledons. The expansionof kinetin-treated cotyledons is further promoted by red light,but not by far-red, as is also the case with germination itself. 3. A number of purine derivatives which have been reported topromote germination also cause expansion of isolated cotyledons. 4. Gibberellic acid promotes both hypocotyl elongation and cotyledonexpansion in the dark, but this effect does not interact withthe phytochrome system. The site of action of gibberellic acidprobably lies in the axis. 5. It is concluded, therefore, that the site of kinetin actionis in the cotyledons, whose expansion helps to break the seed-coatwhen light or gibberellin has contributed the primary stimulus. ¹ Present address: Johnson Foundation for Medical Physics, Universityof Pennsylvania, Philadelphia, Pa. (Received January 16, 1963; )     

Blue light induced inhibition of seed germination: The necessity of the fruit coats for the blue light response

The seeds (achenes) of Laportea bulbifera require a chilling to break their dormancy and are negatively photoblastic. Their germination is inhibited by both continuous blue light and continuous or prolonged far-red radiation. The germination of de-coated seeds, prepared by removing the fruit coats, however, was strongly inhibited by continuous far-red, but not by continuous blue light. Photoreversible germination by a brief irradiation with red light occurred when the chilled seeds were exposed to prolonged far-red light. These results suggest that far-red light may regulate the germination of L. bulbifera seeds through the phytochrome system which exists in the regions other than fruit coats and that the blue light reaction may be governed by other photoreceptor system(s).     

Photocontrol of flowering and stem extension of the intermediate-day plant Echinacea purpurea

Intermediate-day plants (IDP) flower most rapidly and completely under intermediate photoperiods (e.g., 12–14 h of light), but few species have been identified and their flowering responses are not well understood. We identified Echinacea purpurea Moench as an IDP and, based on our results, propose a novel mechanism for flowering of IDP. Two genotypes of E. purpurea ('Bravado' and 'Magnus') flowered most completely (≥79%) and rapidly and at the youngest physiological age under intermediate photoperiods of 13–15 h. Few (≤14%) plants flowered under 10- or 24-h photoperiods, indicating E . purpurea is a strongly quantitative IDP. Plants were also induced to flower when 15-h dark periods were interrupted with as few as 7.5 min of low-intensity lighting (night interruption, NI). Flowering was progressively earlier as the NI increased to 1 h, but was delayed when the NI was extended to 4 h. Stem length increased by ≥230% as the photoperiod or NI duration increased, until plants received a saturating duration (at 14 or 1 h, respectively). Flowering was inhibited when 16-h photoperiods were deficient in red (R, 600–700 nm) light, and was promoted when photoperiods were deficient in far-red (FR, 700–800 nm) light. Because of our results, we propose the flowering behavior of IDP such as E . purpurea is composed of two mechanisms: a light-dependent response operating through light-labile (type I) phytochrome in which flowering is inhibited by an LD, and a light-stable (type II) phytochrome (i.e., phyB, D and E) response in which flowering is promoted by a short-night.     

Photomanipulation of phytochrome in lettuce seeds

Seeds of lettuce (Lactuca sativa L. cv. Grand Rapids) were imbibed and given either short irradiation with red or far red light prior to drying or dried under continuous red or far red light. Seeds treated with either short or continuous red germinate in darkness, whereas seeds treated with either short or continuous far red require a short exposure to red light, after a period of imbibition, to stimulate germination. Irradiation of dry red seeds with far red light immediately before sowing results in a marked inhibition of germination. This result was predicted since far red-absorbing form phytochrome can be photoconverted to the intermediate P650 (absorbance maximum 650 nm) in freeze-dried tissue. A similar far red treatment to continuous red seeds is less effective and it is concluded that in these seeds a proportion of total phytochrome is blocked as intermediates between red-absorbing and far red-absorbing form phytochrome, which only form the far red-absorbing form of phytochrome on imbibition. The inhibition of dry short red seeds by far red light can be reversed by an irradiation with short red light given immediately before sowing, confirming that P650 can be photoconverted back to the far red-absorbing form of phytochrome. The results are discussed in relation to seed maturation (dehydration) on the parent plant.