5-Fluorodeoxyuridine inhibition of photoperiodically induced flowering inChenopodium rubrum L.

Flowering in the short day plantChenopodium rubrum was inhibited by 5-fluoro-deoxyuridine (FDU) at a concentration of 4×10^?6 M and higher when applied during photoperiodic induction or immediately afterwards. This inhibition is always accompanied by a general reduction of growth (e.g. a decrease in the first leaf length). The mitotic activity within the shoot apex is completely blocked by FDU application during the photoperiodic treatment. The floral induction (evocationsensu Evans) was not cancelled in this situation as was revealed when reversing the FDU effect by thymidine application. One day after the end of the photoperiodic treatment (the plants were transferred to continuous light again) the FDU inhibition becomes irreparable. The results indicate that DNA synthesis and hence the mitotic activity are not obligatory prerequisites for photoperiodic floral induction inChenopodium. Low concentrations of FDU may promote flowering under suboptimal floral induction.     

Growth effects of 2-thiouracil and possibility of selective inhibition of floral differentiation inChenopodium rubrum L.

The effect of 2-thiouracil on vegetative growth and floral differentiation was investigated inChenopodium rubrum plants grown in water cultures. Between the low concentrations of the agent, stimulating vegetative growth and floral differentiation, and those inhibiting both these processes, a narrow concentration range was found (1.10^?5 m to 2.10^?5 m), where growth was inhibited selectively. At a concentration of 1.10^?4 m a selective inhibition of development was found when 2-thiouracil was applied at the beginning of photoperiodic induction. Inhibition of development was strong regardless of whether 2-thiouracil was applied before, during or closely after 4 days of photoperiodic induction; the degree of growth inhibition, however, changed in dependence on photoperiodic induction. The strongest relative inhibition of development, calculated as a ratio between development and growth, was observed always at the beginning of photoperiodic induction. Investigation of plant growth as well as the anatomical and autoradiographic study after the application of 2-thiouracil indicate that the inhibition becomes evident at the end of 4 days of application by an overall growth inhibition and a decrease of mitotic activity. Reversal by uracil was possible after simultaneous application of 2-thiouracil. The nature of the selective inhibition is discussed and two possible interpretations of the data obtained are analyzed: a) different response of growth processes in apices and young vegetative organs respectively with regard to different participation of cell division and elongation, b) specific inhibition of floral differentiation.     

Methyl jasmonate inhibits growth and flowering inChenopodium rubrum

C. rubrum plants of different age were treated with methyl jasmonate (JA-Me), in some cases in combination with photoperiodic flower induction. Plants treated with JA-Me (3×10^?4, 3×10^?5 and 5×10^?7M) showed inhibition of growth and flowering. No effect of JA-Me application on ethylene formation was observed.     

Response of Lemma perpusilla to Periodic Transfer to Distilled Water

The flowering of Lemma perpusilla grown on half-strength Hutner's medium with sucrose under inductive photo-periods is inhibited in a periodic manner by daily transfers to water for short periods of time. The phase of maximal inhibition of flowering caused by water treatment is about 1 to 2 hours after the time of maximal sensitivity to light pulses. The rhythm of sensitivity to water treatments does not persist under continuous blue light. Supplementing the water with either Ca(NO₃)₂ or K₂HPO₄ partially reverses the inhibition of flowering, with the first salt being more effective. Supplementation with NH₄NO₃ or MgSO₄ increases the inhibition. The water effect on flowering is not observed in plants grown on half-strength Hutner's medium without sucrose. The water treatments may act by removing or destroying a crucial precursor for photoperiodic induction, with the other conditions modifying permeability. The system provides a new technique for investigating the mechanism of photoperiodic induction.     

Effects of exogenous cytokinins on flowering of the short-day plantChenopodium rubrum L.

Kinetin at a concentration from 3.10^-6 M to 1.10^-3 M was applied to the plumule ofChenopodium rubrum plants during photoperiodic induction. Different levels of induction were compared (one and three short days). The higher concentrations of kinetin applied to induced plants inhibited flower formation. The rate of leaf initiation was increased under these treatments. Lower concentrations of kinetin (from 3.10^-6 M to 1.10^-5 M) usually promoted lateral bud formation and flowering. The step-wise application of kinetin revealed that the inhibitory effect on flowering had been restricted to the inductive period. The effects of kinetin, benzyladenine and trans-zeatin were compared in plants partially induced by two short days. High concentrations always inhibited flowering. Benzyladenine was the most effective in this respect. Root removal diminished the inhibitory effects of cytokinins on flowering as was stated with benzyladenine. It is assumed that endogenous cytokinins play a role in the regulation of organogenetic activity of the stem apical meristem. Depending on the photoperiodic conditions, they presumably exert their activity by maintaining the vegetative functions of the apex.     

The incorporation of uridine-3H into the shoot apices of photoperiodically induced and non-induced plants ofChenopodium rubrum L.

The influence of photoperiodic induction on the incorporation of uridine-³H into the shoot apices ofChenopodium rubrum was studied using the technique of autoradiography. No increase in uridine incorporation was detected either during induction lasting three days or immediately after its termination. Pyroninophylia likewise did not rise. However, changes in uridine incorporation related to morphogenetic activity during leaf formation and later during differentiation of inflorescences were well marked. The distribution of label in the nucleus immediately after three inductive cycles shows the ratio of extranucleolar to nucleolar incorporation to be higher in non-induced control plants than in induced ones. Data from literature pointing to an activation of RNA synthesis during transition to flowering are discussed and compared with other systems where ontogenetic changes are accompanied by marked changes in RNA synthesis. It is assumed that the activation of RNA synthesis after induction is connected mainly with the activation of growth. However, inChenopodium rubrum photoperiodic induction proceeds together with limited growth and without activation of RNA synthesis.     

Ethylene and IAA interactions in the inhibition of photoperiodic flower induction of <Emphasis Type="Italic">Pharbitis nil</Emphasis>

It has been shown that both IAA and ethylene application inhibit flower induction in the short-day plant Pharbitis nil. However application of IAA has elevated ethylene production in this plant, as well. Strong enhancement of ethylene production is also correlated with the night-break effect, which completely inhibits flowering. In order to determine what the role of IAA and ethylene is in the photoperiodic flower induction in Pharbitis nil, we measured changes in their levels during inductive and non-inductive photoperiods, and the effects of ethylene biosynthesis and action inhibitors on inhibition of flowering by IAA. Our results have shown that the inhibitory effect of IAA on Pharbitis nil flowering is not physiological but is connected with its effect on ethylene biosynthesis.     

大豆开花后光周期反应的研究

利用中国大豆主要生态区的生育期不同的代表品种研究了大豆（Ｇｌｙｃｉｎｅ ｍ ａｘ （Ｌ．）Ｍｅｒｒ．）开花后对光照长度的反应． 结果表明,不同成熟期的大豆品种开花后普遍存在着对光照长度的反应．这种反应属于典型的光周期现象,而不是由温度的替代作用、光合时间的改变或前期短日后效应引起的．开花后光周期反应不仅存在于大豆的花荚期,而且存在于鼓粒期．研究认为∶大豆开花结实对光周期的需求是一个连续过程;光周期对大豆生育期的调控作用存在于出苗至成熟的全过程;光周期诱导开花和促进成熟的作用有一定的共同性;光周期诱导效果具有持效性和可逆性     

Ethylene and ABA interactions in the regulation of flower induction in Pharbitis nil

Hormones are included in the essential elements that control the induction of flowering. Ethylene is thought to be a strong inhibitor of flowering in short day plants (SDPs), whereas the involvement of abscisic acid (ABA) in the regulation of flowering of plants is not well understood. The dual role of ABA in the photoperiodic flower induction of the SDP Pharbitis nil and the interaction between ABA and ethylene were examined in the present experiments. Application of ABA on the cotyledons during the inductive 16-h-long night inhibited flowering. However, ABA application on the cotyledons or the shoot apices during the subinductive 12-h-long night resulted in slight stimulation of flowering. Application of ABA also resulted in enhanced ethylene production. Whereas nordihydroguaiaretic acid (NDGA) - an ABA biosynthesis inhibitor - applied on the cotyledons of 5-d-old seedlings during the inductive night inhibited both the formation of axillary and of terminal flower buds, application of 2-aminoethoxyvinylglycine (AVG) and 2,5-norbornadiene (NBD) - inhibitors of ethylene action - reversed the inhibitory effect of ABA on flowering. ABA levels in the cotyledons of seedlings exposed to a 16-h-long inductive night markedly increased. Such an effect was not observed when the inductive night was interrupted with a 15-min-long red light pulse or when seedlings were treated at the same time with gaseous ethylene during the dark period. Lower levels of ABA were observed in seedlings treated with NDGA during the inductive night. These results may suggest that ABA plays an important role in the photoperiodic induction of flowering in P. nil seedlings, and that the inhibitory effect of ethylene on P. nil flowering inhibition may depend on its influence on the ABA level. A reversal of the inhibitory effect of ethylene on flower induction through a simultaneous treatment of induced seedlings with both ethylene and ABA strongly supports this hypothesis.     

Indirect Action of Benzyladenine and Other Chemicals on Flowering of Pharbitis nil Chois: Action by Interference with Assimilate Translocation from Induced Cotyledons

Benzyladenine (BA) brushed on the cotyledons of 4-day-old seedlings of Pharbitis nil Chois. markedly stimulates flowering. Greates response is obtained for concentrations between 44 and 440 micromolar. The action of BA is on processes in the cotyledon as shown by the response to its site of application, to the dosage applied and to the requirement for its application prior to the dark period. There was little or no effect of BA treatment on either the time measurement processes of photoperiodic induction or on the generation of floral stimulus. Transport of photosynthetic assimilate from the cotyledons to the shoot apex was altered.     

Investigation of the endogenous rhythm of flowering inChenopodium rubrum L.

Under the conditions applied in our laboratory 4 1/2 days old plants ofChenopodium rubrum require 2–3 photoperiodic cycles for maximal flowering response, whereas 2 1/2 days old plants are able to flower after having obtained a single inductive cycle. The period length of the free-running rhythm of flowering observed in 2 1/2 days old plants after a single transfer from light to darkness is 30h and the first peak of flowering occurs at about hour 12 in darkness. When a cycle consisting of 16h darkness and 8h light or of 8h darkness and 8h light precedes the long dark period the rhythm is rephased. Rephasing is greater when the light commenced to act on the positive slope of the first peak of the free running rhythm than when it impinged on the negative slope. With an 8h interruption of darkness by light rhythm phase is controlled by the light-on, as well as by the light-off signal. Feeding 0.4 M glucose during the long period of darkness enhanced the amplitude of the flowering response and, moreover, substituted for one photoperiodic cycle.     

Treatments that Enhance Flowering in the Post-inductive Period of a Short-day Plant, Chenopodium rubrum L.

Flowering was promoted in young plants of Chenopodium rubrumL. by application of growth inhibitors such as 5-fluorodeoxyuridine(FUDR) and (2-chloroethyl) trimethylammonium chloride (CCC),growth substances (indol-3yl-acetic acid, IAA), by the removalof roots and by drought. All the treatments were effective onlyduring the post-inductive period and at the threshold levelof photoperiodic induction. The response of plants was strictlytime-dependent. The experimental data indicate that the stimulationof flowering is usually accompanied by inhibition of leaf initiationand growth. The treatments probably produced variation in thequantitative expression of flowering by causing a shift in emphasisin the development of leaf and bud primordia at the shoot apex.The dynamic analysis of differentiation of the shoot apex indicatesa correlation between the morphological stage of the shoot apexand its responsiveness to the treatments.     

Changes in organ growth ofChenopodium rubrum due to suboptimal and multiple photoperiodic cycles with and without flowering effect

The growth changes of cotyledons, leaves, hypocotyls and roots due to photoperiodic induction in short day plantChenopodium rubrum were investigated in relation to flowering. Six-day old plants were induced by photoperiods with a different number of dark hours. We found that the degree of inhibition which occurred during induction in the growth of leaves, cotyledons and roots similarly as the stimulation of hypocotyl is proportional to the length of dark period. The photoperiods with 12, 16 and 20 dark hours bring about marked inhibition of growth and at the same time induce flowering in terminal and axillary meristems. The inhibitory effect of critical period for flowering,i.e. 8 dark hours, is not apparent in all criteria used and even the flower differentiation is retarded. The photoperiods of 4 and 6 dark hours did not affect growth and were ineffective in inducing flowering even if their number has been increased. The experiments with inductive photoperiod interrupted by light break have clearly shown that growth pattern characteristic for induced plants can be evoked in purely vegetative ones. Such statement did not exclude the possible importance of growth inhibition as a modifying factor of flower differentiation. We demonstrated that the early events of flower bud differentiation are accompanied by stimulation of leaf growth. The evaluation of growth and development of axillary buds at different nodes of insertion enabled us to quantify the photoperiodic effect and to detect the effects due to differences in dark period length not exceeding 2 hours.     

Root-shoot correlation linked with photoperiodic floral induction inChenopodium rubrum L.

Inhibition of root growth was observed inChenopodium rubrum under photoperiodic conditions inducing flowering. That this inhibition is mediated by the cotyledons was shown directly by the effect of their excision, which changes the responsiveness of the roots to photoperiodic treatment. On the other hand, decapitation did not lead to such an effect. Some evidence is put forward suggesting that changes in IAA may be involved in these correlations. The existence of two different mechanisms of photoperiodic action in flowering and in root growth is proposed to explain these differences.     

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

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

Gibberellins and the floral transition in Sinapis alba

The putative role of gibberellins in the transition to flowering was investigated in Sinapis alba , a caulescent long-day (LD) plant. It was observed that: (1) physiological doses of exogenous gibberellins (GA₁, GA₃, GA₉) do not cause the floral shift of the meristem when applied to plants grown in short days but have some positive effect on the flowering response to a suboptimal LD; no inhibition was observed in any case; (2) GA-biosynthesis inhibitors (prohexadione-Ca and paclobutrazol) considerably inhibit stem growth but have some negative effect on flowering only when a suboptimal LD is given; and (3) the floral transition induced by one 22-h LD does not correlate with any detectable change in GA content of the apical bud, of the leaves, and of the phloem exudate reaching the apex. Taken together, these results suggest that GAs do not act as a major signal for photoperiodic flower induction in Sinapis .     

Sixty-Five Years of Searching for the Signals That Trigger Flowering

This review summarizes the long-term research of photoperiodic flower induction in two Chenopodium species, one of which, C. rubrum, is a short-day plant, while the other, C. murale, is a long-day plant. In the course of purification of florigenic extracts inducing flowering in C. rubrum plants kept under noninductive photoperiodic conditions, we did not succeed in identifying its active component(s). During the inductive treatment, the phloem transport of cytokinins was enhanced and their content in the shoot apex greatly increased for some time. The review also summarizes the results of the application of direct electric current to the leaves of C. rubrum plants. This treatment interferes with the transport of some active signal(s) from leaves to the apex, resulting in the inhibition of flowering induction. The problem of the rhythmicity of flowering in C. rubrum plants was considered, particularly, the possibility of melatonin (N-acetyl-5-methoxytryptamin) participation in photoperiodic induction. Melatonin accumulation in darkness appears to determine the amplitude of the flowering rhythm but not its phase or period.     

Chemical control of flowering in the long-day plantLemna gibba G3

Lemna gibba G3 is an ideal system for studying the chemical control of flowering in a photoperiodic plant due to its small size and aquatic growth habit which allow substances to be taken up continuously and rapidly distributed throughout the plant. Each of the known plant growth regulators has been tested onL. gibba G3 and only the gibberellins appear to be important for flowering, although they are not the limiting factor for flowering on short days. Salicylic acid (SA) and ferricyanide will both induce flowering inL. gibba G3 with ferricyanide being most effective on short days where flowering is daylength limited and SA most effective where flowering is limited by factors other than daylength. The ferricyanide action is probably due to HCN and it may act during photoperception or photoinduction. SA is most effective when reversing the inhibition caused by various parameters including copper and agar, and its effect is always strongly daylength dependent. It is postulated that SA may interact with the flowering stimulus to promote flowering and thus that SA acts at some point following photoinduction and the formation of the flowering stimulus.     

Changes in the content of endogenous auxins in apical buds ofchenopodium rubrum L. Induced with respect to the endogenous rhythm in capacity to flower

The content of endogenous auxins was examined in apical buds ofChenopodium rubrum plants induced by a photoperiodic cycle of 16h darkness and 8h light followed by a dark period of various duration so as to correspond with either maximal or minimal flowering response in the endogenous rhythm in capacity to flower initiated by the photoperiodic treatment. Apical buds of potentially generative plants contained less auxins than apical buds of plants which remained in the vegetative state. Apical buds from plants treated with kinetin (1. 10^-3 M) and therefore remaining in the vegetative state showed an auxin level comparable to that of untreated plants exhibiting minimal flowering response irrespective of the duration of the second dark period. Plants cultivated on a sucrose solution (0.6 M) during the second dark period became generative even at the normal minimum of flowering. The auxin content of the apical buds was low, similarly as in untreated plants induced for a period leading to maximal flowering response. On the other hand, apical buds from plants grown on sucrose solution during a dark period leading to the manifestation of maximal flowering response showed a relatively high auxin content comparable to that found in untreated plants which had obtained a more extended induction by three photoperiodic cycles. The results are discussed with respect to the possible role of endogenous auxins in the regulation of the changes in growth correlations occurring in the shoot apex during photoperiodic induction and in the expression of the competence to flower.