CONSTANS delays flowering and affects tuber yield in potato

CONSTANS (CO) has a central role in the photoperiodic regulation of flowering in Arabidopsis thaliana. We show here that potato (Solanum tuberosum ssp. andigena) plants constitutively expressing Arabidopsis CO (pACo plants) flower late under all photoperiodic conditions tested. Exogenous application of gibberellic acid to pACo plants corrected their short stem phenotype but not their late flowering. To further understand the effect of CO in potato, we used three photoperiodic conditions: short days (SD), which strongly induce tuberisation of wild type plants, SD supplemented with a night break (SD+NB), which are moderately inductive, and tuberisation-inhibiting long days. Tuberisation of pACo plants was delayed under SD and very strongly delayed or completely inhibited under SD+NB, suggesting that CO affects an autonomous pathway controlling potato tuberisation. In addition, tuber yield, a trait of high agronomic relevance, was significantly increased in pACo plants expressing moderate CO levels. Our results indicate that CO affects flowering and stem elongation through distinct mechanisms and suggest that its effects on flowering and tuberisation in potato are photoperiod-independent.     

GENETIC AND HORMONAL REGULATION OF GROWTH,FLOWERING, AND SEX EXPRESSION IN PLANTS

The interaction of the genetic and hormonal regulation of growth, flowering, and sex expression in plants is discussed. The genetic control of these processes is characterized, and data on their hormonal regulation are supplied. The interaction of genetic and hormonal regulation is considered with reference to tall-growing and genetic dwarf forms of the pea and wheat plants. It is shown that in the dwarf forms of the pea plant and in many other varieties, growth stimulation in response to treatment with the phytohormone gibberellic acid is clearly manifested and the expression of genetic dwarfism is eliminated, whereas in dwarf wheats it is expressed only slightly, if at all. At the same time both tall-growing and dwarf forms of both pea and wheat show a clearly defined growth retardation response to treatment with the growth inhibitor, abscisic acid, which causes the expression of physiological dwarfism. The short- and long-day characteristics of the photoperiodic response of plants are described as genetically controlled features, and data are given on the induction of flowering of a long-day variety coneflower grown under short-day conditions with the aid of gibberellins extracted from leaves of long-day vegetative plants of short-day Mammoth tobacco. Data are also supplied on the induction of flowering of a short-day variety, red-leaved goosefoot, grown under continuous light with the aid of metabolites extracted from leaves of the same Mammoth tobacco plants flowering under short-day conditions. This demonstrates the possibility of hormonal regulation of the genetically controlled long-day and short-day characteristics in photoperiodically sensitive plants. Genetic and hormonal regulation of sex expression in two dioecious plants, hemp and spinach, is discussed. It is shown that sex expression in these plants is regulated by gibberellins which are synthesized in leaves and cause male sex expression and by cytokinins which are synthesized in the roots and cause female sex expression. These data indicate that sex expression in dioecious plants is the result of interaction between the genetic apparatus and phytohormones.     

What is going on with the hormonal control of flowering in plants?

Molecular genetic studies using Arabidopsis thaliana as a model system have overwhelmingly revealed many important molecular mechanisms underlying the control of various biological events, including floral induction in plants. The major genetic pathways of flowering have been characterized in-depth, and include the photoperiod, vernalization, autonomous and gibberellin pathways. In recent years, novel flowering pathways are increasingly being identified. These include age, thermosensory, sugar, stress and hormonal signals to control floral transition. Among them, hormonal control of flowering except the gibberellin pathway is not formally considered a major flowering pathway per se, due to relatively weak and often pleiotropic genetic effects, complex phenotypic variations, including some controversial ones. However, a number of recent studies have suggested that various stress signals may be mediated by hormonal regulation of flowering. In view of molecular diversity in plant kingdoms, this review begins with an assessment of photoperiodic flowering, not in A. thaliana, but in rice (Oryza sativa); rice is a staple crop for human consumption worldwide, and is a model system of short-day plants, cereals and breeding crops. The rice flowering pathway is then compared with that of A. thaliana. This review then aims to update our knowledge on hormonal control of flowering, and integrate it into the entire flowering gene network.     

Effects of diurnal and bidiurnal photoperiodic treatments on mitosis in Pharbitis

Abstract

Photoperiodic effect on mitotic activity of buds from dwarf Pharbitis has been analyzed. No significant differences in mitotic activity were found in plants grown under long days or diurnal (24 h) light break photoperiodic treatments. Differences in both mitotic activity and flowering were seen in plants subjected to diurnal short days, bidiurnal (48 h) short days, or bidiurnal short days with light breaks. An elevation of mitotic activity occurs in plants grown in bidiurnal photoperiodic treatments compared to diurnal treatments. The differences in mitotic activity of buds, both vegetative and floral, seem to indicate that both phytochrome and light effect on an endogenous rhythm influence meristematic activity. Also, the extended dark period of a bidiurnal short day enhances both mitosis and flowering.     

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.     

Substances of plant flowering

The investigation of the hormonal nature of plant flowering in connection with their photoperiodic reaction has shown that flowering depends on a bicomponental system of hormones, gibberellins regulating stem formation and growth and substances of the anthesin type regulating flower formation. In agreement with the division of the photoperiodic reaction into a leaf and a stem phase the study of the internal factors acting on plant flowering was carried out by means of leaf and stem (apex, bud and callus) models. The results obtained from work with leaf models proved the presence of two groups of hormones of flowering in plants. The data obtained from the application of stem models pointed to the localization of the action of gibberellin and anthesin in different zones of the shoot apices and characterized the potential capacity for flower formation of isolated callus tissue of neutral and photoperiodically sensitive species.     

Polyamine translocation following photoperiodic flowering induction in soybean

Radioactivity translocation after [¹⁴C]-spermidine application over the third trifoliate leaf of soybean plants ( Glycine max . [L.] Merr, cv. Williams) was checked during the first 72 h of short day (SD) treatment to study the involvement of polyamines (PAs) in photoperiodic flowering induction. PAs and/or their metabolites were translocated from the supplied leaf to all parts of the plant. Radioactivity reached its highest concentration in the upper portion of the stem, i.e. the apical bud and the youngest leaf. After the beginning of the first inductive night, the detected radioactivity showed two peaks of maximal concentration. The first arose after the first inductive night, coinciding with the proper flowering induction process; the second one arose after the third inductive night, coinciding with the first morphological symptoms of the transition of vegetative meristems to the reproductive condition. Soluble free PAs showed a different balance in the apical bud of SD-induced plants compared with LD-non induced control plants. Soluble conjugated PAs were detected as traces. It is suggested that under flowering inductive conditions, PAs play a different role according to the stage of the flowering process. Thus, their translocation from the leaves to the axillary and apical buds might be related, in a first step, to the fact that they were part of the complex mechanism of the flowering signal, and in a second step, to the flower transition of vegetative buds.     

Transition from absolute to quantitative short-day control in Nicotiana tabacum cv. Maryland Mammoth

The response in vitro of thin cell layers, excised from different stem regions of Nicotiana tabacum cv. Maryland Mammoth plants at various developmental stages, was studied under different photoperiodic treatments. The aim was to determine at which stage of plant development, and in which region of the stem, the absolute short-day requirement, indispensable for the induction of the flowering process in this genotype, becomes quantitative and whether it remains short-day. The explants were cultured on a medium suitable for flower neoformation, and were exposed for 30 days to the following treatments: continuous darkness, 8 h light/16 h dark per day, 16 h light/8 h dark per day, and continuous light. The first flowers on explants were observed from plants that were still in the vegetative state, but whose apex showed an accelerated production of axillary vegetative buds, as observed histologically. These explants were excised from the first 10 internodes below the first node with a leaf ≥ 5 cm in length (apical site), and produced flowers only under short-day treatment. When the apical dome initiated the organization of the terminal flower, the apical site explants developed flowers under both short-day and long-day treatments. At the same stage, explants from the 15th to the 20th internode below the first leaf ≥ 5 cm in length also formed flowers, but only under short-day. When the plant showed a complete inflorescence, flowers were also present on explants from the most basal stem internodes and from the inflorescence branches. At this stage, flower neoformation occurred under all treatments; however, under short-day the number of explants showing flowers not associated with vegetative buds on the same sample greatly exceeded that observed under other treatments, as did the mean number of flowers per explant (except the basal regions). In conclusion, in the post-inductive phases of the flowering process, the photoperiodic requirement of this genotype is always short-day. The superficial tissues of the stem require either absolute or quantitative short-day treatment, depending on their position on the stem and the stage of evolution of the flowering process in the terminal apex.     

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.     

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.     

钙在植物花发育过程中的作用

对于园林观赏植物,开花是一个非常重要的发育阶段,它直接影响花卉的品质。近年来,植物花发育的分子生物学研究进展迅速,并取得了一些突破性成果。钙作为第二信使在植物信号转导中起着非常重要的作用,大量研究显示,钙有可能参与开花控制。本文总结了钙信号与植物花发育这一领域的最新研究进展,包括以下几个方面的内容：钙在植物成花诱导(包括光周期诱导和低温诱导)中的作用;花芽分化时期钙在植物叶芽和花芽中的动态分布及组织培养条件下不同钙浓度对花芽分化的影响;钙与花衰老的关系。     

钙在植物花发育过程中的作用

对于园林观赏植物,开花是一个非常重要的发育阶段,它直接影响花卉的品质。近年 来,植物花发育的分子生物学研究进展迅速,并取得了一些突破性成果。钙作为第二信使在植物信号转导中起着非常重要的作用,大量研究显示,钙有可能参与开花控制。本文总结了钙信号与植物花发育这一领域的最新研究进展,包括以下几个方面的内容：钙在植物成花诱 导（包括光周期诱导和低温诱导）中的作用;花芽分化时期钙在植物叶芽和花芽中的动态分布及组织培养条件下不同钙浓度对花芽分化的影响;钙与花衰老的关系。      

Photoperiodism and enzyme rhythms: Kinetic characteristics of the photoperiodic induction of Crassulacean acid metabolism

Summary The effect of photoperiod on Crassulacean acid metabolism (CAM) in Kalanchoe blossfeldiana Poellniz, cv. Tom Thumb, has characteristics similar to its effect on flowering in this plant (although these two phenomena are not causally related). The photoperiodic control of CAM is based on (a) dependance on phytochrome, (b) an endogenous circadian rhythm of sensitivity to photoperiodic signals, (c) a balance between specific positive (increase in enzyme capacity) and negative (inhibitory substances) effects of the photoperiod. Variations in malate content, capacity of phosphoenolpyruvate (PEP) carboxylase, and capacity of CAM inhibitors in young leaves were measured under photoperiodic conditions noninductive for CAM and after transfer into photoperiodic conditions inductive for CAM. Essential characteristics of the photoperiodic induction of CAM are: 1) lag time for malate accumulation; 2) after-effect of the inductive photoperiod on the malate accumulation, on the increase in PEP carboxylase capacity, and on the decrease in the level of long-day produced inhibitors; final levels of malate, enzyme capacity and inhibitor are proportional to the number of inductive day-night cycles; 3) cireadian rhythm in PEP carboxylase capacity with a fixed phase under noninductive photoperiods and a continuously shifting phase under inductive photoperiods, after complex advancing and delaying transients. Kinetic similarities indicate that photoperiodic control of different physiological functions, namely, CAM and flowering, may be achieved through similar mechanisms. Preliminary results with species of Bryophyllum and Sedum support this hypothesis. Phase relationships suggest different degrees of coupling between endogenous enzymic rhythm and photoperiod, depending on whether the plants are under long days or short days.     

Photoperiodic control of dormancy in Sedum telephium and some other herbaceous perennial plants

The environmental control of dormancy and flowering of the herbaceous perennial Sedum telephium was studied in controlled environments. Short photoperiods induced growth cessation and the formation of resting buds in both seedlings and mature plants, whereas long photoperiods resulted in immediate growth activation of dormant buds. No chilling was required for dormancy release, even in plants induced to dormancy and maintained at high temperature (21°C) for more than 3 months. The critical photoperiod for dormancy release was about 15 h, a minimum of four long-day (LD) cycles (24 h) being required. The true photoperiodic nature of this response was ascertained by night interruption experiments. Flowering of S. telephium was found to have an obligatory LD requirement, with no requirement for vernalization. The critical photoperiod and minimum number of inductive cycles for floral induction were the same as for dormancy release. Dormancy release by long days was also obtained in preliminary experiments with three other herbaceous perennials. The eco-physiological significance of photoperiodic control of dormancy is discussed, and it is concluded that the mechanism ensures stability of winter dormancy, even under conditions of climatic warming.     

FTIP1 is an essential regulator required for florigen transport

The capacity to respond to day length, photoperiodism, is crucial for flowering plants to adapt to seasonal change. The photoperiodic control of flowering in plants is mediated by a long-distance mobile floral stimulus called florigen that moves from leaves to the shoot apex. Although the proteins encoded by FLOWERING LOCUS T (FT) in Arabidopsis and its orthologs in other plants are identified as the long-sought florigen, whether their transport is a simple diffusion process or under regulation remains elusive. Here we show that an endoplasmic reticulum (ER) membrane protein, FT-INTERACTING PROTEIN 1 (FTIP1), is an essential regulator required for FT protein transport in Arabidopsis. Loss of function of FTIP1 exhibits late flowering under long days, which is partly due to the compromised FT movement to the shoot apex. FTIP1 and FT share similar mRNA expression patterns and subcellular localization, and they interact specifically in phloem companion cells. FTIP1 is required for FT export from companion cells to sieve elements, thus affecting FT transport through the phloem to the SAM. Our results provide a mechanistic understanding of florigen transport, demonstrating that FT moves in a regulated manner and that FTIP1 mediates FT transport to induce flowering.     

A Set of Three Genes Regulates Photoperiodic Responses of Flowering in Rice (Oryza sativa)

Differentiation in photoperiodic response of flowering has been key to the evolution and wide geographic distribution of rice, an essentially short-day plant. Crosses were made such that the hybrid F1 plants flower later than the late-flowering parents to investigate the genetic basis underlying this differentiation. From initial experiments, three major genes for flowering time were identified from four naturally occurring variants under natural long-day conditions. An F2-derived trigenic mutant line bred-true for a day-neutral response was selected and used as the recipient to synchronize the genetic background for the major genes. Experiments conducted under various daylengths indicated that these genes are responsible for photoperiodic sensitivity and the trihybrid has a critical daylength between 13.5 and 14 h. The three genes regulate photoperiodic responses qualitatively and quantitatively through complementary and other epistatic effects, respectively. The complementation suggests that the three genes act in a linear manner to repress the transition from the vegetative to reproductive phases under long daylengths. This set of genes also provides a model to understand the genetic mechanism underlying the elongated vegetative growth period in the F1 generation, which is usually an obstacle to the use of heterosis, and the selection for early maturation in rice breeding.