Tomato Seed Germination. Osmotic Pretreatment and Far Red Inhibition

At 25 °C germination of tomato (Lycopersicon lycopersicum)seeds is inhibited by continuous and intermittent far red illumination.It is also inhibited by a single 30 min far red irradiationgiven 8 h from the start of imbibition. The incubation of seedsin a mannitol solution inhibitory for germination has no effecton the final germination percentage after seeds are subsequentlytransferred to water. A 30 min far red irradiation at the timeof transfer results in partial inhibition of germination. Thisinhibition can be released by the continuation of osmotic incubationfor several days before the transfer to water. At the end ofa 7 d dark period of osmotic incubation, inhibition of subsequentgermination in water can be realized only by continuous farred illumination. Seeds osmotically pretreated for 7 d and afterwardsdried-back show a mean time to 50% germination significantlylower than that of untreated seeds. Moreover, besides singleand intermittent, even continuous far red light has no inhibitoryeffect on the germination of these seeds. It is concluded that,in addition to the already known germination advantages, osmoticpresowing treatment also induces the ability of seeds to germinateunder unfavourable light conditi.     

Studies of the Mechanism of Enhancement of Phytochrome-dependent Lettuce Seed Germination by Prechilling

Temperature and kinetic studies were performed to examine the mechanism by which prechilling stimulates phytochrome-dependent seed germination in lettuce, Lactuca sativa, L. cv. Grand Rapids. Imbibed seeds were given a short far red irradiation and one day of dark incubation at 20 C to establish very low levels of the far red-absorbing form of phytochrome—(Pfr). Germination was greatly stimulated by subsequent prechilling treatments when they were followed by a second short far red irradiation. Prechilling therefore increased germination sensitivity to the low, normally inhibitory Pfr levels established by far red irradiation. This sensitivity increased with lowered prechilling temperature to a maximum near 4 C. It was linearly dependent upon duration of prechilling at 4 C up to a near maximal response at 10 hours, and it decayed in a converse manner when seeds were returned to 20 C after 10 hours at 4 C. Prechilling also increased germination responses to subsequent periods of high levels of Pfr which were initiated by red and terminated by far red irradiations. High Pfr periods adequate to promote the germination of unchilled seeds produced sharp inflections at 18 C in the dependence of germination on prechilling temperature. Rates of phytochrome potentiation of germination were not affected by prechilling. The response to prechilling fit a mechanism involving homeoviscous adaptation of membrane lipids to temperature.     

Phytochrome and Seed Germination: VI. Phytochrome and Temperature Interaction in the Control of Cucumber Seed Germination

Phytochrome control of cucumber seed germination is temperature-dependent. A prolonged exposure to radiation from broad spectrum far red sources (Pfr/P = 0.05 to 0.07) prevents germination at temperatures below 20 C. Above 20 C there is no inhibition and it appears as if there is an escape from phytochrome control. However, radiation from a monochromatic, narrow band 730 nanometer source (Pfr/P < 0.02) inhibits germination at temperatures above 20 C. This result supports the idea that, even at high temperatures, Pfr is responsible for the activation of germination. After 4 days of exposure to far red, a short red irradiation is quite effective in promoting germination if temperatures during the dark incubation periods are maintained below 20 C; red becomes effective at temperatures above 20 C. Promotion of germination will take place at a temperature of 25 C or higher without red irradiation. Again, we have an apparent escape from phytochrome control at high temperatures. However, if higher temperatures are used for only short periods, 2 to 6 hours, in combination with short red irradiation, one can demonstrate that activation of germination at high temperatures is still dependent on phytochrome. Phytochrome is probably destroyed during prolonged exposure to far red. Thus, the subsequent short red irradiation establishes levels of Pfr which may not be sufficient to promote germination at low temperatures but are probably adequate at high temperatures.     

Inhibition of Prechill-induced Dark Germination in Sorghum halepense (L.) Pers. Seeds by Phytochrome Transformations

A 10 C dark prechilling of johnsongrass [Sorghum halepense (L.) Pers.] seeds, when terminated by a 2-hr, 40 C temperature shift, potentiates about 40% germination at 20 C in darkness. Irradiation of the seeds before, during, and at the end of prechilling with far red light reduces the subsequent germination, although red irradiation after the far red can overcome some of the inhibition. However, either brief red or far red irradiation given immediately after the temperature shift inhibits subsequent germination by one-third to one-half. The results suggest that the far red-absorbing form of phytochrome is a factor in the prechill-induced dark germination and that phytochrome participates in the inhibition of germination by irradiations immediately after the temperature shift.     

Phytochrome and Seed Germination. I. Temperature Dependence and Relative P(FR) Levels in the Germination of Dark-germinating Tomato Seeds

Germination of the dark-germinating seeds of 3 varieties of tomato is controlled by the phytochrome system. Germination is inhibited by far red radiation and repromoted by red applied after far red. At low temperatures, 17 to 20°, a single, low energy far red irradiation is sufficient to inhibit germination in all 3 varieties. At higher temperatures far red is less effective in the inhibition of the germination of the tomato seeds. The phytochrome fraction present as P_FR in the dark-germinating seeds of the Ace variety is about 40% of the total phytochrome present.     

Photophysiology of Kalanchoë Seed Germination

The effects of irradiations with different proportions of red/farred light and of gibberellic acid on the phytochrome-mediated seed germination of Kalanchoë blossfeldiana cv. Feuerblüte, were studied. The seed coat transmits much more red than far-red light, and therefore the energy ratio between 660 nm and 730 nm is given only for the transmitted light. Decreasing this ratio from 65 to 1.0 caused only a very slight inhibition. If this ratio is further lowered to 0.64, a 10 min terminal irradiation after a 3-h white light photoperiod is inhibitory, but a 12-h photoperiod or continuous irradiation is not. If the ratio is decreased to 0.44 or 0.31, a 12-h photoperiod is now also inhibitory, although continuous irradiation and 10 min terminal irradiation are still more inhibitory. These results are discussed in terms of phytochrome phototransformations. Although gibberellic acid is unable to cause any germination in complete darkness, it can result in a very high germination percentage, if combined with treatments which by themselves do not induce any germination such as continuous far-red, terminal far-red after short photoperiods, or very short photoperiods at 25°C. These results point to a strong synergism between gibberellic acid and the so-called stabilized form of phytochrome, P*_FR.     

Phytochrome control of skotodormancy release in Grand Rapids lettuce achenes

Light-requiring Grand Rapids lettuce ( Lactuca sativa L.) achenes develop skotodormancy when imbibed in darkness for 7 days at 25°C. Redried skotodormant achenes maintain this type of dormancy upon subsequent rehydration. At 25°C full germination of skotodormant achenes can be induced by continuous and intermittent red light illumination as well as by several brief red irradiations given daily. One brief (10 min) red light irradiation can partly break skotodormancy at 20°C, while at lower temperatures the same treatment results in full induction of germination. Phytochrome control of the release from skotodormancy is proven by a) the dependence of the germination response on the relative sequence of red and far-red light in cyclic irradiations, and b) the reversion of red action by subsequent far-red irradiation. The time course of germination of skotodormant achenes treated with intermittent red light depends upon the length of dark interval between the light pulses. Germination is considerably delayed compared to that of non-skotodormant ones, induced by a single brief red light treatment. This fact in combination with the requirement, over a long period of time, of P_fr action for full manifestation of germination, indicates that skotodormancy is a deeper form of dormancy. It is concluded that the germination of lettuce achenes may always be subjected to phytochrome control.     

Changes in Phytochrome Expressed by Germination of Amaranthus retroflexus L. Seeds

Effects of red (600 to 680 nanometers) and far red (700 to 760 nanometers) irradiances on Amaranthus retroflexus L. seeds indicate that synthesis of phytochrome in the red-absorbing form takes place in water-imbibed nongerminating seeds at 35 C. After 96 hours in darkness, conversion of about 0.10% phytochrome to the far red-absorbing form induces 50% germination. Continuous far red radiation at 35 C with an irradiance of 0.4 × 10⁻¹⁰ Einsteins per square centimeter per second caused photoinactivation of phytochrome about equal to the rate of synthesis. Germination of seeds at 35 C, following far red irradiation adequate to establish the photostationary state, is enhanced by holding at 26 C for 16 minutes. Germination is unaffected relative to controls at constant temperature, if the period at 26 C precedes irradiation. The results indicate a quick response to action of phytochrome in a germination process.     

Photoinduced Seed Germination of Oenothera biennis L: II. Analysis of the Photoinduction Period

The photoinduction period of Oenothera biennis L. seed germination was analyzed by varying the photoinduction temperature and by substituting red light pulses for continuous red light. At 24°C, seeds require 36 hours of continuous red light for maximal percent germination. The optimal photoinduction temperature is 32°C, with higher and lower temperatures being strongly inhibitory. A 30 minute exposure to far-red light, given immediately after a red light period of 1 to 36 hours, reduces germination by about 25%. Seeds escape from far-red inhibition with a half-time of 5 to 10 hours, depending on the length of the red exposure that precedes the far-red light. Periodic 15 minute pulses of red light can substitute for continuous red light in stimulating germination. Ted red light pulses, with 6 hours of darkness between successive pulses, cause maximal germination. The response to periodic red light is fully reversible by far-red light. Probit analysis of the periodic light response shows that as the length of the dark periods between successive pulses increases, less incident light is needed to induce germination but the population variance in light sensitivity remains constant. Probit analysis of the temperature response shows that as the photoinduction temperature increases from 16 to 32°C, less incident light is needed to induce germination and the population variance in light sensitivity also increases.     

Photoinduction of Spore Germination in Marchantia polymorpha L. is Mediated by Photosynthesis

The effects of light on spore germination (protrusion of protonemata)in the liverwort Marchantia polymorpha L. were examined. Sporegermination was found to be light dependent and light irradiationfor 10 h or longer was necessary. Test using specific wavelengthsshowed that the entire spectrum from near UV to red light waseffective, red light being the most effective. Spore germinationcould be induced by intermittent irradiation with 15-min redlight pulses given every 1 or 2 h for 24 h. The effect of intermittentred light was not reversed by subsequent or simultaneous far-redlight irradiation. However, spore germination was inhibitedby the photosynthesis inhibitor DCMU (100 µM). Completeinhibition of spore germination was found when DCMU was givenduring the light period. When DCMU was applied during the darkperiods, only a slight reduction of germination rate was observed.Further, it was found that Chl formed in the spores during imbibitionin darkness. Light sensitivity increased at nearly the samerate as the appearance of Chl. Moreover, spore germination wasinduced in total darkness by the addition of glucose to themedium. These results clearly indicate that photosynthesis mediatesthe photoinduction of spore germination in Marchantia polymorpha. (Received May 13, 1999; Accepted July 14, 1999)     

Photocontrol of the Germination of Onoclea Spores: IV. Metabolic Changes during Germination

The changes in levels of metabolites during photoinduced germination of Onoclea sensibilis L. spores are described. Proteins and lipids, which constitute 25 and 20%, respectively, of the unimbibed spores on a dry weight basis, are hydrolyzed at the time of differentiation and elongation of the germling cells and may be utilized for these processes. Sucrose degradation, starch synthesis, and active respiration occur during dark imbibition, but these processes are accelerated by red or far red irradiation. Endogenous sucrose is the probable source of the carbon skeleton for starch synthesis.     

Photocontrol of Anthocyanin Synthesis: IV. Dose Dependence and Reciprocity Relationships in Anthocyanin Synthesis

Under continuous far red light, anthocyanin synthesis in young, dark-grown cabbage seedlings (Brassica oleracea cv. Red Acre) is irradiance-dependent and fails to follow the reciprocity (irradiance × time = constant) relationships. Under intermittent far red treatments extended over a prolonged period of time, anthocyanin synthesis becomes dose dependent, and reciprocity relationships are valid. Intermittent far red treatments with short dark intervals between successive irradiations are as effective as continuous treatments, if the total radiation doses applied with the two types of treatments are equal and are applied over equally long periods of time. The high effectiveness of inter-mittent treatments, the dose dependence, and the validity of the reciprocity relationships suggest that cycling between red-absorbing form of phytochrome and far red-absorbing form of phytochrome and the formation of electronically excited far red-absorbing form of phytochrome, or the involvement of a second photoreactive system, besides phytochrome, may play only a minor role in high irradiance reaction anthocyanin synthesis brought about by prolonged exposures to far red irradiation.     

Phytochrome Control of Germination of Rumex crispus L. Seeds Induced by Temperature Shifts

High germination of curly dock (Rumex crispus L.) seeds is evident after suitable imbibition and temperature shift treatment, but germination at constant temperatures fails without an input of far red-absorbing form of phytochrome. Preliminary imbibitions at high temperatures (30 C) sharply reduce germination induced by temperature shifts. High germination may be restored by low energies of red radiation, or by brief far red adequate for the photosteady state. Prolonged far red during imbibition also nullifies temperature shift-induced germination. After prolonged far red, high germination may be restored by red radiation of an energy dependent upon the duration of the far red treatment. The evidence supports the conclusion that dark germination induced by temperature shifts arises from the interaction of pre-existent far red-absorbing form of phytochrome in the mature seeds with the temperature shift.     

Light Actions in the Germination of Cocklebur Seeds: I. DIFFERENCES IN THE LIGHT RESPONSES OF THE UPPER AND LOWER SEEDS

Germination responses to light were studied in the upper andlower seeds of cocklebur (Xanthium pennsylvanicum Wallr.). Thelower seed was dark-germinating and negatively photoblastic;the upper one had a red-light (R) requirement and was positivelyphotoblastic. Germination of the lower seeds was inhibited bya prolonged single irradiation with R, blue (B) or far-red (FR)light applied during imbibition. The maximal inhibitory effectof a single irradiation occurred 9 h and 13 h after the startof soaking at 33 °C and 23 °C, respectively. However,the inhibitory effect of R differed from that of B and FR, byonly delaying germination. A single exposure to B or FR lightcould be replaced by intermittent B or FR irradiation, and theireffects were repeatedly reversible by the following R irradiation.If the upper seeds were not exposed to R during imbibition,they failed to germinate even at 33 °C which was optimalfor germination, and the promotive effect of R increased withdelay of its application time. The photoperceptive locus incocklebur seeds was the axial tissue for all B, R and FR. Lightreceived by the cotyledonary tissue had little effect. Germinationdimorphism in response to light is discussed with respect tothe phytochrome content and the ageing of axial tissues. Key words: Blue light, Dimorphism, Far red light, Germination, Red light, Xanthium seed     

Action Spectrum between 250 and 800 Nanometers for the Photoinduced Inhibition of Spore Germination in Pteris vittata

An action spectrum between 250 and 800 nm for the inhibitionof red-light-induced germination of spores in the fern Pterisvittata was determined on the Okazaki Large Spectrograph. Theresultant spectrum showed prominent peaks of effectiveness atabout 370, 440 and 730 nm and a minor peak in the neighborhoodof 260 nm. Next, a brief red light irradiation was given immediatelyafter the monochromatic irradiation to cancel the inhibitoryeffect caused by simultaneously formed P_R. This resulted ina complete disappearance of the peak at 730 nm and considerabledecrease of other peaks in the shorter wavelength region exceptat 260 nm. Further correction of the latter spectrum by consideringthe transmission spectrum of a spore coat revealed that 260nm light acted more effectively than lights of 370 and 440 nm.The inhibitory effect of UV light on spore germination was nullifiedby subsequent irradiation with red light for 24 h or darknessfor 48 h followed by a brief red irradiation, indicating thatthe inhibitory action of UV light was ascribable to a blue-ultraviolet light-absorbing pigment. ⁴Present address (KT) and permanent address (MF): Botany Department,Faculty of Science, University of Tokyo, Hongo, Tokyo 113, Japan. (Received July 30, 1983; Accepted November 21, 1983)     

Dormancy in Dioscorea: Gibberellin-Induced Inhibition or Promotion in Seed Germination of D. tokoro and D. tenuipes in Relation to Light Quality

Effects of light and gibberellic acid (GA₃) application on the germination of Dioscorea tokoro Makino and Dioscorea tenuipes Franch. et Savat. were observed. For complete germination, seeds of both species required prechilling in moist condition before incubation at a higher temperature. Red light irradiation during the incubation after the prechilling promoted germination; blue, green, or far red light markedly inhibited the germination of both species.     

Photocontrol of fungal spore germination

Germination of Puccinia graminis f. sp. tritici uredospores is inhibited by continuous irradiation. Prehydration of spores enhances both dark germination and photoinhibition. Simultaneous irradiation with ineffective red (653 nanometers) and inhibitory far red light (720 nanometers) results in partial nullification of the inhibition brought about by far red light alone. This result would be consistent with the involveent of a photoreversible pigment system similar to phytochrome, operating via the high irradiance reaction.     

Photocontrol of the germination of onoclea spores: I. Action spectrum

Light stimulates the germination of spores of the fern Onoclea sensibilis L. At high dosages, broad band red, far red, and blue light promote maximal germination. Maximal sensitivity to these spectral regions is attained from 6 to 48 hours of dark presoaking, and all induced rapid germination after a lag of 30 to 36 hours. Maximal germination is attained approximately 70 hours after irradiation. Dose response curves suggest log linearity. The action spectrum to cause 50% germination shows that spores are most sensitive to irradiation in the red region (620-680 nm) with an incident energy less than 1000 ergs cm⁻²; sensitivity decreases towards both shorter and longer wavelengths. Although the action spectrum is suggestive of phytochrome involvement, photoreversibility of germination between red and far red light has not been demonstrated with Onoclea spores. An absorption spectrum of the intact spores reveals the presence of chlorophylls and carotenoids. Since the presence of 3-(3,4-dichlorophenyl)-1,1-dimethylurea does not inhibit germination, it is concluded that photosynthesis does not play a role in the germination process.     

Stimulation of Empress Tree Seed Germination by Liquid Smoke

The germination of Empress tree (Paulownia tomentosa Steud.) seeds is phytochrome-controlled. Liquid smoke could not induce germination in darkness but red light irradiation of liquid smoke imbibed seeds induced a high percentage of germination. Maximum germination was achieved at liquid smoke concentration of 0.1% (v/v) when present during the imbibition phase or during the phase of phytochrome activity. The light requirement of these seeds could be completely substituted by exogenously applied gibberellins. In the presence of liquid smoke, optimal concentrations of GA₃, GA₄, and GA₉ necessary for inducing germination were several times lower than in the controls, while that of GA₇ was equally active when applied at a concentration one order of magnitude lower. The inhibitory effect of the applied growth retardants was strongly reduced and liquid smoke, in the presence of retardants, allowed light-induced germination, if applied simultaneously or after retardants treatment.     

The effect of SAN 9789 and light on the germination of seeds from Taraxacum vulgare

Photoblastic seeds (achenes) of Taraxacum vulgare coll. were treated with a water solution of SAN 9789, 4-chloro-5 (methylamino) -2- (α,α,α-trifluoro- m -tolyl) -3(2H) pyridazinone. SAN-treatment increased the germination in darkness from 0 to 12%. An irradiation for 5 min with red light, giving a germination of 12% for seeds in water only, gave together with SAN treatment a germination of 60%. In both water and SAN, the effect of red irradiation could be reversed by a short irradiation (15 min) of far-red light. If far-red light was repeatedly given (5 min per h) it had hardly any effect on germination in water (4% germination), but for seeds in SAN solution, intermittent far-red light had a stimulating effect (63% germination). If far-red light was given continuously for 96 h, the germination in water was 1% and in SAN solution 17%. The results in the present paper indicate that SAN may broaden the concentration interval of P_fr for which germination is high.