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.     

The effect of daylength on the transition to flowering in phytochrome-deficient, late-flowering and double mutants of Arabidopsis thaliana

The effect of daylength on flowering was investigated in the following mutants of Arabidopsis thaliana : phytochrome B deficient ( hy3=phyB ); phytochrome chromophore deficient ( hy2 ); late-flowering ( co, gi. fca and fwa ); the hy2 and hy3 , late-flowering double mutants and the hy2, hy3 , late-flowering triple mutants. The hy mutants flower with fewer rosette leaves than the Landsberg erecta wild type under both long day and short day conditions and express this effect to a different degree in all late-flowering mutant backgrounds and under both daylengths, with the exception of fca under short days. The number of cauline leaves and days to flowering is less affected by the hy genotype. The hy2, hy3 double mutants flower with even fewer rosette leaves than the hy2 and hy3 monogenic mutants, suggesting an inhibitory role for phytochrome B and other stable phytochromes on flowering. The complex interaction between phytochrome, daylength and the effect of the late-flowering genes on the various parameters that describe the transition to flowering in Arabidopsis is discussed.     

The Influence of Nicotinic Acid and Plant Hormones on Flowering in Lemna

Nicotinic acid induces flowering in Lemna paucicostata 151 and381 and Lemna gibba G3 when they are grown in one tenth-strengthM medium under continuous light. For L. paucicostata 151 and381, the simultaneous addition of IAA, GA₃ or ABA to the mediumleads to an inhibition of the flower-inducing effect of nicotinicacid, while zeatin leads to a further stimulation of floweringabove that obtained by nicotinic acid alone. By contrast, inL. gibba G3 all four plant hormones inhibit the nicotinic acid-inducedstimulation of flowering. The effect of nicotinic acid on flowering in all three plantsis strongly daylength dependent when the plants are grown inhalf-strength Hutner's medium. Thus, nicotinic acid causes floweringin L. gibba G3 on continuous light but not on 9L:15D or 10L:14Dregimes. In L. paucicostata 381 nicotinic acid has a small effecton 12L:12D regime, a large effect on a 13L:11D regime and noeffect with daylengths longer than 14 hours, and in L. paucicostata151 nicotinic acid is only effective on daylengths shorter thanabout 11 hours. However, in L. paucicostata 151 and 381 treatmentwith both nicotinic acid and zeatin results in flowering undercontinuous light on half-strength Hutner's medium. Nicotinic acid is present in different Lemna but its concentrationdoes not appear to be influenced by changes in daylength. Thus,flowering clearly cannot be controlled by nicotinic acid actingalone, but the results of this study indicate that nicotinicacid could interact with other factors, possibly including oneor more of the known plant hormones, to influence the floweringprocess in Lemna. (Received August 28, 1985; Accepted October 29, 1985)     

Influence of Ammonium on the Ability of Salicylic Acid to Induce Flowering in the Short-Day Plant Lemna paucicostata 6746

When the short-day plant Lemna paucicostata 6746 is grown inhalf-strength Hutner's medium, which contains 1.25 mM NH4NO3,salicylic acid does not induce flowering on daylengths of 16hr or longer. By contrast, in M (Hoagland-type medium), Pirson-Seidelor ammonium-free half-strength Hutner's media, none of whichcontain ammonium, salicylic acid is able to induce some floweringeven under continuous light. Neverthless, in each of these threemedia the effect of salicylic acid is strongly daylength dependentbecause there is a sharp drop in the flowering response to salicylicacid between the 12 and 16 hr daylengths, and the floweringresponse is nearly constant from the 16 hr daylength to continuouslight. Ammonium has the opposite effect and at 50 to 75 µMis able to overcome the salicylic acid effect completely andprevent any flowering on daylength of 16 hr or longer. (Received December 3, 1980; Accepted March 5, 1981)     

Crassulacean Acid Metabolism and Crassulacean Acid Metabolism Modifications in Peperomia camptotricha

Peperomia camptotricha, a tropical epiphyte from Mexico, shows variable forms of Crassulacean acid metabolism (CAM). Young leaves exhibit CAM-cycling, while mature leaves show an intermediate type of metabolism, between CAM and CAM-cycling, having approximately the same amount of nighttime gas exchange as daytime. Metabolism of young leaves appears independent of daylength, but mature leaves have a tendency toward more CAM-like metabolism under short days (8 hours). Large differences in the physical appearance of plants were found between those grown under short daylengths and those grown under long daylengths (14 hours). Some anatomical differences were also detected in the leaves. Water stress caused a switch to CAM in young and mature leaves, and as water stress increased, they shifted to CAM-idling.     

Role of Adenosine 3'5'-Cyclic Monophosphate in Flowering of a Short-Day Duckweed Lemna paucicostata 6746

The well-known second messenger in animal systems, adenosine3',5'-cyclic monophosphate (cAMP), stimulated flowering in theshort-day plant, Lemna paucicostata 6746, under both inductiveand non-inductive photoperiods, when grown on modified Bonnerand Devirian medium. Flowering could be achieved even in continuouslight in the presence of cAMP, although the intensity of theresponse was still daylength-dependent. Besides flower induction,cAMP also enhanced their development. 5'-AMP and 5'-ATP alsoinduced some flowering under non-inductive long days, but theresponse was weak as compared to that of cAMP. 5'-ADP, in contrast,had no effect whatsoever on flowering under long days, whereasa small effect was observed under inductive short days. Whatcould be the probable mechanism of action of cAMP and whetherits effect on flowering is mediated through the cAMP-adenylcyclasesystem, has been discussed. (Received May 11, 1988; Accepted June 23, 1988)     

Factors Controlling Flowering in the Hop (Humulus lupulus L.)

In experiments on the effects of daylength on the growth andflowering of the perennial hop it was shown that Humulus lupulusis a short-day plant. The absolute length of the short day isimportant since very short days (8h) induce dormancy beforeflowering can occur. Light-break treatment may therefore promoteor inhibit flowering according to the associated main photoperiod.A minimum node number must have been differentiated before thehop can be induced to initiate flowers, an effect analogousto the juvenile condition. Minimum leaf number and criticaldaylength for induction depend on variety. At low temperature,induction is possible with longer photoperiods. Promotion offlowers by growth retardants (B9 and CCC) in unfavourable daylengths,and delay of initiation by gibberellic acid treatment were alsoobserved.     

Effect of Low-intensity Red and Far-red Light and High-intensity White Light on the Flowering Response of the Long-day Plant Lemna gibba G3

The long-day plant Lemna gibba L., strain G3 exhibits a relatively low sensitivity to short, white-light interruptions given during the dark period of a short-day cycle. However, the plants are fairly sensitive to low-intensity red light treatments given during a 15-hour dark period on the third day of a 2LD-(9L:15D)-2LD-7SD schedule. Far-red light is almost as effective as red light, and attempts to reverse the red light response with subsequent far-red light treatments have not been successful. Blue light proved to be without effect. When plants were grown on a 48-hour cycle with 15 minutes of red light every 4 hours during the dark period, the critical daylength was reduced from about 32 hours to slightly less than 12 hours.

Continuous red light induced a fairly good flowering response. However, as little as 1 hour of white light each day gave a significant improvement in the flowering response over that of the continuous red light control. White light of 600 to 700 ft-c was more effective than white light of 60 to 70 ft-c. The white light was much more effective when divided into 2 equal exposures given 8 to 12 hours apart. These results suggest an increase in light sensitivity with regard to flower induction about 8 to 10 hours after the start of the light period.

     

Reproductive response of a tropical mammal, the musk shrew (Suncus murinus), to photoperiod

Musk shrews (Suncus murinus) were maintained for 8 weeks in long (16 h light:8 h darkness) or short (8 h light:16 h darkness) daylengths. Males housed in short daylength had significantly lighter androgen-dependent sex accessory organs than did males kept in long daylengths. This same trend was noted in male sexual behaviour. However, the weights of the testes and epididymides and sperm numbers did not differ. Females housed in short daylengths had significantly lighter cervices and were less likely to demonstrate sex behaviour than animals kept in under long daylengths. Ovarian and uterine weights did not differ. These results suggest that the ability to respond to photoperiod can exist in tropical mammals, even if it is not used as a cue to time seasonal breeding.     

Factors Affecting the Flowering of Andropogon gayanus Kunth. Responses to Photoperiod, Temperature and Growth Regulators

Andropogon gayanus is a short-day plant with a critical daylengthfor flowering between 12 and 14 h. Flowering is more intenseas the daylength is shortened from 12 h to 8 h, initiation isalso accelerated by increasing durations of short-day treatment.Plants flower more readily with increasing age. The maximumflowering response can be induced by short-day exposure of onlyone expanding leaf. A fixed number of short days distributedamong different groups of leaves on the plant gave less floweringthan the same number given to only one group. Approximately25°C was optimal for flowering. Root removal treatmentsin short day reduced both growth and flowering responses. Application of the growth regulators indol 3yl-acetic acid,abscisic acid, gibberellic acid or dimethylaminosuccinamic acid(B9) was inhibitory to flowering in all cases but gibberellicacid and B9 applied simultaneously had no inhibitory effect.Growth hormone treatments failed to stimulate flower developmentin long day. These results are discussed in relation to the internal mechanismscontrolling growth and flowering.     

Comparison of photoperiodic sensitivity of green-leafed and red-leafed perilla

The flowering responses of Perilla frutescens (L.) Britt. (green-leafed Perilla) and P. crispa (Thunb.) Tanaka (red-leafed Perilla) to different 24-hour photoperiodic regimes were compared quantitatively. Although both taxa are short-day plants, P. frutescens was less strict in its short-day requirement, forming macroscopically visible inflorescence primordia and open flowers even in continuous light after 77 to 85 and 132 days, respectively, compared to 12 and 22 days in 8-hour days. P. crispa took only 4 days longer to reach the same stages in 8-hour short days, but exhibited a sharply defined critical day-length between 14 and 16 hours. Two-hour light-breaks in the middle of a 16-hour night were more effective in inhibiting floral development in P. crispa than in P. frutescens, for which they had a small effect compared to true long-days, particularly on the formation of inflorescence primordia. The number of short days needed to obtain the flowering in P. frutescens plants raised in continuous light decreased in a linear manner with the age of the plants upon transfer to short days.     

Photoperiodic control of apical senescence in a genetic line of peas

An early flowering genetic line of peas (Pisum sativum L.), designated G2, has dominant genes at two different loci, both of which function in short days to greatly extend the reproductive phase and thus to delay apical senescence. Long days (18 hours) promote senescence in this line, but the effect is reversible by reinstatement of short days (9 hours) until 3 to 4 days before the apex senesces. The response to photoperiod was quantitative. Increasing the photoperiod from 14 to 18 hours led to a progressive decrease in the number of nodes formed prior to death of the apex. Induction of senescence was determined by the total number of hours of light and darkness rather than by the length of the dark period. Senescence required flower and fruit development as well as long days.     

Influence of day length and temperature on number of main stem leaves and time to flowering in lupin

A growth chamber experiment was carried out to investigate the influence of day length and temperature on the development of flowering in eight varieties of the three grain lupin species Lupinus albus (Wat and C3396), L. angustifolius (Gungurru, Polonez and W26) and L. luteus, (Juno, Radames and Teo). The plants were grown at two temperatures, 10°C and 18°C, in combination with five daylength regimes: 10, 14, 18, 24 h day at full light intensity and 10 h full light extended with 8 h low intensity light. Increased daylength decreased days from sowing to flowering in all varieties, but had little effect on thermal time to flowering in most varieties. However, C3396, W26 and Radames had a significantly longer thermal time to flowering at high, non‐vernalising temperature (18°C) at short daylengths. Low light intensity daylength extension did not significantly influence thermal time to flowering. For flower initiation, measured as number of leaves on the main stem three types of response were found. All varieties formed fewer leaves on the main stem at 10°C than at 18°C, although the two thermo‐neutral varieties of L. luteus, Juno and Teo, gave only a small response to temperature and daylength. In Polonez, Gungurru and Wat, low temperature decreased leaf number, but there was only a small response to changes in daylength. Three varieties, C3396, W26 and Radames, showed longer thermal time to flowering at 18°C with short daylengths. This could be explained by a greater number of main stem leaves formed at short daylength at non‐vernalising temperatures. Increased daylength decreased leaf number in these varieties, but never to a smaller number than for plants grown at 10°C. In these varieties, low intensity extension of the daylength had a similar (W26, Radames) or decreased (C3396) effect compared to full light extension. The hastening of time to flowering by long days could be separated into two effects: a high light energy effect hastened development by increasing the rate of leaf appearance in all varieties, while low light energy in thermo‐sensitive varieties was able to substitute for vernalisation by decreasing leaf number.     

Studies on the Photoperiodic Responses in Short-Day Plant Helianthus tubevotus

Plants of Helianthus tuberosus, variety white tuber, were treated with various daylengths of 4, 6, 8, 10, 11, 12, 13, 14, 15, 18 h. for 25 days as soon as six leaves on a plant appeard Irradiation for 6–13 h per day induced the plants to form flower buds and flowering, daylength with 14 h or longer kept the plants in vegetative growth. The experiments showed that this variety of Helianthus tuberosus required short days for flowering and the critical daylength was about 13 h. The plants were treated with short days for different durations. At least 17 days were required, Formation of flower buds and flowering had positive correlation with the number of short days over 17 days. After short-day induction, the shorter the daylength is, the more the flower buds inverted. Long-day treatment after an appropriate period of short days wouid reduce the number of flower and induce new vegetative branches from flowering granehes.     

Identification of the Flower-inducing Factor Isolated from Aphid Honeydew as being Salicylic Acid

Honeydew produced by the aphid Dactynotus ambrosiae when feeding on flowering or vegetative plants of the short day plant Xanthium strumarium contains an active substance capable of inducing flowering in the long day plant Lemna gibba G3. In the present study, this active material has been identified as salicylic acid through the use of gas-liquid chromatography and mass, infrared, and ultraviolet spectrometry. Authentic salicylic acid induces flowering in L. gibba G3 under strict short day conditions with an optimal response at about 5.6 μm. The possible significance of salicylic acid for the control of flowering in Xanthium or L. gibba G3 is discussed.     

Daylength influences pelage and plasma prolactin concentrations but not reproduction in the prairie vole, Microtus ochrogaster

Short daylengths did not affect testes weight or spermatogenic index in male voles or uterine weight in female voles. Short daylengths did stimulate the growth of a winter pelage in both sexes; short-day voles had longer underhairs and guard hairs and a thicker, more dense pelage than did long-day voles. Plasma prolactin concentrations were five times higher in long-day than in short-day females and 25% higher in long-day males than in short-day males. The effect of short daylength on pelage was prevented by pinealectomy. We suggest that the growth of a winter coat is an obligate adaptation for winter survival, stimulated by exposure to short daylengths, but that changes in breeding activity are facultative and dependent to a greater extent on other cues for seasonal synchronization.     

Photophosphorylation during Chloroplast Development in Red Kidney Bean: II. Photophosphorylation and Photoreduction Appear Concomitantly but Initially are Uncoupled

Cyclic phosphorylation with phenazine methosulfate and noncyclic phosphorylation and reduction with ferricyanide were detected in isolated chloroplasts from greening bean leaves after 3 to 4 hours of illumination. Activity commenced when rapid synthesis of chlorophyll was initiated. Rates of photophosphorylation were comparable to mature levels by 15 to 18 hours of development. Photoreduction of ferricyanide attained a peak value by 12 hours of illumination and subsequently fell to normal levels by 15 to 18 hours. With ferricyanide, the P/e₂ ratios were initially less than 0.1 but were close to 1.0 after 18 hours of illumination. The data suggested that photosystems I and II appeared concomitantly in the chloroplast but were not fully operative until later in development. Proplastids and immature chloroplasts exhibited high capacities to reduce ferricyanide in the dark. The rates of dark reduction rapidly diminished to low levels by 15 hours of illumination when normal rates of photochemical activity were observed. After a 2-to 3-day lag, a rapid increase in leaf fresh weight was noted at the time total chlorophyll content reached steady state values on a fresh weight basis. With fresh weight as an index of growth, primary leaves completed their development after 6 to 7 days of illumination.     

Temporal and intraclonal variation of flowering and pseudovivipary in Poa bulbosa

Background and Aims

Versatility in the reproductive development of pseudoviviparous grasses in response to growth conditions is an intriguing reproduction strategy. To better understand this strategy, this study examined variation in flowering and pseudovivipary among populations, co-occurring clones within populations, and among tillers in individual clones of Poa bulbosa, a summer-dormant geophytic grass that reproduces sexually by seed, and asexually by basal tiller bulbs and bulbils formed in proliferated panicles.

Methods

Clones were collected from 17 populations across a rainfall gradient. Patterns of reproduction were monitored for 11 years in a common garden experiment and related to interannual differences in climatic conditions. Intraclonal variation in flowering and pseudovivipary was studied in a phytotron, under daylengths marginal for flowering induction.

Key Results

Clones showed large temporal variability in their reproductive behaviour. They flowered in some years but not in others, produced normal or proliferated panicles in different years, or became dormant without flowering. Proliferating clones did not show a distinct time sequence of flowering and proliferation across years. Populations differed in incidence of flowering and proliferation. The proportion of flowering clones increased with decreasing rainfall at the site of population origin, but no consistent relationship was found between flowering and precipitation in the common garden experiment across years. In contrast, flowering decreased at higher temperatures during early growth stages after bulb sprouting. Pulses of soil fertilization greatly increased the proportion of flowering clones and panicle production. High intraclonal tiller heterogeneity was observed, as shown by the divergent developmental fates of daughter plants arising from bulbs from the same parent clone and grown under similar conditions. Panicle proliferation was enhanced by non-inductive 8 h short days, while marginally inductive 12 h days promoted normal panicles.

Conclusions

Interannual variation in flowering and proliferation in P. bulbosa clones was attributed to differences in the onset of the rainy season, resulting in different daylength and temperature conditions during the early stages of growth, during which induction of flowering and dormancy occurs.