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

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

The interrelationship of growth and frost tolerance in winter rye

The reduction in growth of winter cereals that occurs in the fall is thought to be required for the development of frost resistance. In the present study, the interrelationship of freezing tolerance and growth was examined by raising winter rye ( Secale cereale cv. Puma) plants at 20/16°C (day/night) and at 5/3°C under 8-, 16- and 24-h daylengths to vary growth rates and frost tolerance. Temperature and irradiance were quantified as thermal time, photothermal time and photosynthetic photon flux and examined by multiple linear regression in order to determine their effects on growth and frost tolerance of rye shoots. At low temperature, both growth and frost tolerance were markedly influenced by daylength and irradiance. Plants grown at 5/3°C with a short daylength accumulated shoot dry weight and increased frost tolerance at a greater rate per unit photothermal time or photon flux than plants grown at longer daylengths. Moreover, 5/3°C plants grown with a 16-h day grew more slowly and were less frost tolerant than plants grown with a 24-h day. We conclude that the interrelationship between growth and frost tolerance is a quantitative one. Frost tolerance is induced only by low temperature, but the development of forst tolerance is dependent upon both irradiance, which affects the amount of photoassimilate available, and daylength, which may affect the partitioning of photoassimilates between growth and frost tolerance.     

Dual induction rather than intermediate daylength response of flowering in Echinacea purpurea

Echinacea purpurea cv. Bravado and Magnus have been reported to be intermediate daylength plants (IDP) which flower in response to photoperiods between 13 and 16 h. The present experiments with E. purpurea cv. Bravado show that E. purpurea is actually a dual induction short-long-day plant which flowers promptly and consistently when grown in short day (SD) followed by long day (LD) conditions, but not with the reverse sequence of photoperiods. The flowering response increased with increasing duration of both the SD and the LD treatments. A minimum of 4 weeks of SD followed by 12 LD was required for complete flowering. No flowering occurred in continuous SD or LD, whereas a high proportion of plants flowered in continuous 14-h daylength. However, flowering was more variable in intermediate daylength than after transition from SD to LD. Furthermore, photoperiods between 13 and 16 h could satisfy both the primary SD induction and the secondary LD induction requirements. As a number of dual induction plants, both short-long-day and long-short-day plants, have such an overlapping window of effective photoperiods that can trigger both the SD and LD responses, the rationale for maintaining IDP as a separate and genuine flowering response group is seriously challenged.     

Photoperiodic regulation of receptacle induction in Sargassum horneri (Phaeophyceae) using clonal thalli

We developed a clonal culture of Sargassum horneri to investigate the effect of photoperiod on reproduction in this species. Regenerated vegetative thalli were obtained using lateral branches excised from a thallus grown from a single embryo under short‐day conditions (SD = 10:14 h light : dark cycle). Lateral branches excised from the SD‐regenerated thallus became vegetative thalli that remained in that phase as long as they were cultured under SD. When an excised lateral branch was cultured under long‐day conditions (LD = 14:10 h light : dark cycle), it began to enter the reproductive phase while still less than 50 mm long. Induction of the reproductive phase was accompanied by a distinctive morphological change – suppression of blade formation at the apical region of the branch; elongation of branches without blades was then followed by differentiation of receptacles bearing conceptacles on their surface. Apices of receptacles were able to interconvert between reproductive and vegetative phases, as blades resprouted upon transfer from LD to SD. The critical day length for induction of receptacle formation was between 13 and 14 h; receptacle formation was also induced under SD conditions with night breaks (NBs). These results strongly suggest that reproductive regulation of S. horneri is a photoperiodic long‐day response. NBs with blue and green light were effective for reproductive induction but not with red light. This suggests that blue‐ and/or green‐light photoreceptors are involved in the photoperiodic reproductive response of S. horneri.     

The influence of light quality on the development of the brown algae Petalonia and Scytosiphon

Petalonia fascia (O. F. Müll.) Kuntze, P. zosterifolia (Reinke) Kuntze and Scytosiphon lomentaria (Lyngb.) J. Ag. have been cultured in white, blue and red light of equal quantum irradiance at 15°C. Hairs, knot-filaments and Ralfsia-like crusts were formed only in blue light, whereas the prostrate system of red-grown plants consisted entirely of sparingly-branched and mostly uniseriate filaments. The production of erect thalli from the prostrate system was controlled or stimulated by red light, but these erect thalli became fertile only in the presence of blue light. All three species exhibited all of these types of response, although specific differences in the degree of certain types of response were observed.     

Circadian Rhythms in Stomatal Responsiveness to Red and Blue Light

Stomata of many plants have circadian rhythms in responsiveness to environmental cues as well as circadian rhythms in aperture. Stomatal responses to red light and blue light are mediated by photosynthetic photoreceptors; responses to blue light are additionally controlled by a specific blue-light photoreceptor. This paper describes circadian rhythmic aspects of stomatal responsiveness to red and blue light in Vicia faba. Plants were exposed to a repeated light:dark regime of 1.5:2.5 h for a total of 48 h, and because the plants could not entrain to this short light:dark cycle, circadian rhythms were able to "free run" as if in continuous light. The rhythm in the stomatal conductance established during the 1.5-h light periods was caused both by a rhythm in sensitivity to light and by a rhythm in the stomatal conductance established during the preceding 2.5-h dark periods. Both rhythms peaked during the middle of the subjective day. Although the stomatal response to blue light is greater than the response to red light at all times of day, there was no discernible difference in period, phase, or amplitude of the rhythm in sensitivity to the two light qualities. We observed no circadian rhythmicity in net carbon assimilation with the 1.5:2.5 h light regime for either red or blue light. In continuous white light, small rhythmic changes in photosynthetic assimilation were observed, but at relatively high light levels, and these appeared to be attributable largely to changes in internal CO2 availability governed by stomatal conductance.     

Rhythmic changes in ribonuclease activity in relation to nucleic acid synthesis in cotyledons ofChenopodium rubrum L. and to floral induction of the plants

Ribonuclease (RNAse) activity was investigated in cotyledons ofChenopodium rubrum plants subjected to various conditions of illumination (photoperiodic induction, continuous light, induction cancelled by interrupting the dark period by a light-break). At the end of the dark period of the single inductive cycles RNAse activity of induced plants was inferior to that of plants grown in continuous light. At the end of the first two cycles the activity was lowest after the interruption of the dark period by light. The investigation of the enzyme in 6h intervals showed rhythmic changes in activity to occur in induced plants. Enzyme activity followed a pattern opposed to this of nucleic acid (NA) synthesis in the cotyledons. In plants from continuous light the enzyme activity did not show any rhythm and in plants having obtained a light-break during the inductive period the rhythm was less distinct than in the induced ones. The period length of the endogenous rhythm of NA synthesis in the cotyledons is about half as long as this of flowering and the peaks of flowering coincide with the throughs of NA synthesis.     

Circadian rhythms and the induction of flowering in the long-day grass Lolium temulentum L.

Plants of Lolium temulentum L. strain Ceres were grown in 8-h short day (SD) for 45 d before being exposed either to a single long day (LD) or to a single 8-h SD given during an extended dark period. For LD induction, the critical photoperiod was between 12 and 14 h, and more than 16 h were needed for a maximal flowering response. During exposure to a single 24-h LD, the translocation of the floral stimulus began between the fourteenth and the sixteenth hours after the start of the light period, and was completed by the twenty-fourth hour. Full flowering was also induced by one 8-h SD beginning 4 or 28 h after the start of a 40-h dark period, i.e. by shifting 12 h forward or beyond the usual SD. The effectiveness of a so-called ‘displaced short day’ (DSD) was not affected by light quality and light intensity. With a mixture of incandescent and fluorescent lights at a total photosynthetic photon flux density of 400 μmol m⁻² s⁻¹, a 4-h light exposure beginning 4 h after the start of a 40-h dark period was sufficient to induce 100% flowering. The flower-inducing effect of a single 8-h DSD was also assessed during a 64-h dark period. Results revealed two maxima at a 20-h interval. This fluctuation in light sensitivity suggests that a circadian rhythm is involved in the control of flowering of L. temulentum.     

Shoot Auxin and Light in the Mechanism Maintaining the Posture of the Cotyledons of Linum usitatissimum

The mean cotyledon angle (liminal day posture) of flax is linearlyrelated to light intensity, the cotyledons being raised withincreasing intensity; adjustments occur when plants are transferredto new conditions. Short days, low light intensity, and prolongeddarkness result in epinasty. Epinastic and hyponastic responsesare enhanced by excision of the epicotyl shoot. Applicationof indoleacetic acid to the cut shoot is an effective substitutefor the excised portion. Ringing the intact epicotyl with 2,3,5-tri-iodobenzoicacid has an effect comparable to excision. Given continuously or in a light/dark cycle, blue light induceshyponasty, and red light maintains the liminal posture for about48 h after which time epinasty occurs. Far-red light given continuouslyinduces epinasty to an extent comparable to the effect of darkness.Irradiation with far-red light even for brief periods priorto placing the plants in darkness enhances the epinastic response:the effect of far-red light is reversed by a subsequent shortexposure to red light. Interruption of a prolonged period of darkness by blue lightconsiderably lessens the subsequent epinastic reaction whetheror not any epinasty has occurred before the light-break. Redor white light given under these circumstances has less effect.     

Photoperiodism of diapause induction in Thyrassia penangae (Lepidoptera: Zygaenidae)

Thyrassia penangae enters winter diapause as a prepupa in a cocoon. Photoperiodism of diapause induction was systematically investigated in this moth. The photoperiodic response curves under 24-h light-dark cycles showed that this insect is a typical long-day species. The critical daylength was 13 h 30 min at 25 °C, 13 h at 30 °C and 12 h 20 min at 28 °C. Transferring experiments from a short day (LD 12:12) to a long day (LD 15:9) or vice versa indicated that photoperiodic sensitivity mainly occurs during the larval period. In experiments using non-24-h light-dark cycles, when the length of photophase exceeded the critical daylength (13.5 h), was diapause inhibited effectively, even when the length of scotophase exceeded the critical nightlength (10.5 h). Only when a long scotophase was combined with a short photophase, diapause was induced effectively. This result suggests that daylength measurement is more important than nightlength measurement in T. penangae. Night interruption experiments under 24-h light-dark cycles exhibited two points of apparent light sensitivity, but the photosensitive position was highly influenced by temperature and the length of scotophase. Nanda-Hamner experiments failed to reveal the involvement of a circadian system in this photoperiodic time measurement. All light-dark cycles from LD 12:12 to LD 12:72 resulted in a short day response, and all cycles from LD 14:4 to LD 14:72 resulted in a long day response, suggesting that photoperiodic time measurement in this moth is performed by a day-interval timer or an hourglass-like clock.     

Circadian and diurnal calcium oscillations encode photoperiodic information in Arabidopsis

We have tested the hypothesis that circadian oscillations in the concentration of cytosolic free calcium ([Ca2+]cyt) can encode information. We imaged oscillations of [Ca2+]cyt in the cotyledons and leaves of Arabidopsis (Arabidopsis thaliana) that have a 24-h period in light/dark cycles and also constant light. The amplitude, phase, and shape of the oscillations of [Ca2+]cyt and [Ca2+]cyt at critical daily time points were controlled by the light/dark regimes in which the plants were grown. These data provide evidence that 24-h oscillations in [Ca2+]cyt encode information concerning daylength and light intensity, which are two major regulators of plant growth and development.     

Light Quality and Irradiance Level Interaction in the Control of Expression of Light-Harvesting Complex of Photosystem II: Pigments, Pigment-Proteins, and mRNA Accumulation

Effects of red and blue light at irradiances from 1.6 to 28.3 micromolar per square meter per second on chloroplast pigments, light-harvesting pigment-proteins associated with photosystem II, and the corresponding mRNA were evaluated in maize (Zea mays L.) plants (OP Golden Bantum) grown for 14 days under 14 hours light/10 hours dark cycles. Accumulation of pigments, pigment-proteins, and mRNA was less in blue than in red light of equal irradiance. The difference between blue and red light, however, varied as a function of irradiance level, and the pattern of this variation suggests irradiance-controlled activation/deactivation (switching) of blue-light receptor. The maximum reduction in blue light of mRNA and proteins associated with light-harvesting complex occurs at lower irradiance levels than the maximum reduction of chlorophylls a and b.     

Seed set inCichorium intybus L. by pollination of flowers developedin vitro

Formation of viable seeds ofCichorium intybus L. was achieved in anin vitro system. Flower formation, pollination, fertilization, embryogenesis and seed development occurredin vitro on chicory root explants on culture medium lacking plant growth regulators. After flower induction under a 24-h daylength treatment, the explants were transferred to a 16-h daylength at 40 E m^-2s^-1 irradiance for pollination and further seed development. Negative results were obtained when root explants were maintained continuously under a 24-h daylength during the whole culture period. Lower seed set was obtained when the cultures were at low irradiance. The need of a dark period and adequate level of irradiance are suggested as important factors to obtain viable seeds. The developedin vitro system can be used as a model to study the factors controlling the reproductive processes, and for the study of self-incompatibility in chicory.     

Regulation of Transpiration in Avena. Responses to White Light Steps

The transient responses in transpiration rate of the primary leaf of 8-days-old oat plants were studied when step changes in the white light irradiance were administered to the plants. A light or dark step was achieved by an instant increase or decrease, respectively, of the irradiance on the leaf surface. Usually the light step responses consisted of one rapid response (rise time 1–2 min) and one slow response (rise time 5–10 min). The dark step response had a fall time of about 10 min. At a high irradiance level and in CO₂-free air the slow response disappeared. This could mean that the slow response is CO₂-controlled. The rapid response was still present under these conditions and is therefore probably caused by other processes than the slow response. In long time experiments several cycles of a circadian rhythm (period 26–28 h) could be followed in the light step responses. Thus the rapid response was small at that time of the day when the slow response was most pronounced and vice versa. The endogenous nature of the rhythm was established in phase-shift experiments. It is concluded that experiments on stomatal dynamics must take circadian variations into account.     

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.

     

Effects of Photoperiod on the Apparent Viscosity of Leaf Cytoplasm in Kalanchoe blossfeldiana: 2. Some Long- and Short-term Effects of Daylength and Spectral Composition of Light

The effects of a range of daylength treatments upon the apparentviscosity of the mesophyll cytoplasm were tested in Kalanchoeand some other daylength-sensitive species by means of a centrifugationtechnique. In Kalanchoe, apparent viscosity increases with increasingshort-day induction, the rise being rapid initially but fallingoff subsequently in the same way as the flowering response.Since the response to light-break treatment is equivalent tolong-day conditions, the effect is truly photoperiodic. Theshort-day increase appears to be only partially reversible uponreturn to long-days. Single leaves may be treated and the effect is not translocatedto the opposite member of a leaf pair. The immediate response to change from light to dark and viceversa is oscillatory; e.g. in the dark an initial fall is succeededby a steep rise and then a more gradual fall in apparent viscosity. Light-breaks with red light are equivalent to white light, andthose with far-red light to darkness. No reversal of red effectsby far-red light has yet been demonstrated. In other species tested, apparent viscosity was significantlyhigher under short-day in Chrysanthemum and Michaelmas Daisy,and under continuous light in Epilobium, i.e. in the daylengthswhich induce flowering. The results are discussed in relationto the present theories of photoperiodism.     

PRECONDITIONING OF GERMINATION IN LETTUCE AT TIME OF FRUIT RIPENING

Koller , D. (The Hebrew U., Jerusalem. ² ) Preconditioning of germination in lettuce at time of fruit ripening. Amer. Jour. Bot. 49(8): 841–844. 1962.—The effects of controlled environments during maturation on subsequent germination response of the seed were investigated with the light-sensitive lettuce ‘Grand Rapids.‘ The plants matured under a variety of temperature/photoperiod combinations, after completing vegetative growth and flower initiation under identical conditions. Good seed set was obtained only at night temperatures between 17 and 23 C, under short-day (8-hr) conditions. Seed germination was tested at 20, 23 and 26 C, in continuous dark and after a light-break, respectively. The conditions under which the seed had ripened brought about quantitative changes in the germination response. The stimulating effects of a light-break on germination at high temperatures did not change, but maturation under high temperatures or in continuous light increased the high-temperature tolerance of seed germination both in continuous dark and after a short light-break.     

Photoperiodic control of tetrasporangium formation in the red alga Rhodochorton purpureum

Six geographical isolates of Rhodochorton purpureum (Lightfoot) Rosenvinge (Rhodophyta, Nemalionales) formed tetrasporangia only in short days at 10°C. For most isolates, the critical day-length increased with latitude of origin from 9.5 h for an isolate from California to 14.5 h for one from Antarctica. Tetrasporangium production could be induced by 9–15 short-day cycles followed by a further 22–28 cycles in long days. A night-break consisting of 1 h of white light in the middle of a 16-h dark period inhibited the short-day response of isolates from low latitudes, but not those from higher latitudes. When a similar night-break was given in the middle of a 14-h dark period, however, the response of all isolates was at least partially inhibited. Night-breaks given at any time in the central 7 h of a 14-h dark period were equally inhibitory. Broad-band red light (0.3–0.4 mmol m^-2), given as a night-break, caused 50% inhibition of the short-day response. At a slightly higher photon exposure (0.6 mmol m^-2, given as 1 μmol m^-2 s^-1 for 10 min), narrow-band red (662 nm) and blue (448 nm) light caused similar inhibition, but green (547 nm) and far-red (731 nm) were ineffective as night-breaks. The inhibitory effect of a 10-min night-break with red light could not be reversed by subsequent exposure to an equal photon exposure of far-red light. These results add to the existing evidence that the pigments mediating photoperiodic responses among algae are more varied than those among flowering plants.     

Photoperiodic long-day control of sporophyll and hair formation in the brown alga Undaria pinnatifida

Two photoperiodic responses, the development of sporophylls and hairs, havebeen quantified in sporophytes of the brown alga Undaria pinnatifida. In a finalexperiment, the algae were cultivated in outdoor, 2000-L seawater tanks in agreenhouse for up to 12 weeks, and daylength was regulated by automatic blindsmounted on top of the tanks. Vegetative young sporophytes were treated undershort-day (SD; 8 h light per day) or long-day conditions (LD; 16 h light perday), at 12 h light per day or in a night-break regime (NB; 8 h light per day,7.5 h dark, 1 h light, 7.5 h dark). The earliest sporophyll development wasobserved 6, 7 or 9 weeks under LD, NB or SD conditions, respectively. After 12 weeksthe sporophylls were significantly longer and wider under LD or NB conditions than inthe SD regime, and only half of the experimental algae had formed sporophyllsunder SD conditions, but all algae under LD or NB conditions. In a foregoing 7-weekculture experiment performed in 300-lL indoor tanks, enhanced sporophyll formationhad also been observed under LD and not under SD conditions (NB omitted). In bothexperiments, blade elongation rates remained high until the end of theexperiments in SD, but declined during sporophyll initiation in LD, NB or at 12 hlight per day. Another difference caused by photoperiod was observed in regard to thedevelopment of surface hair spots which occurred in both experiments on the bladesin LD, NB or at 12 h light per day with identical densities, but were completelylacking under SD conditions. It is concluded that U. pinnatifida is afacultatative long-day plant in regard to reproduction forming vigorously sporophyllsin long days, and an obligate long-day plant in regard to hair formation.     

A CIRCADIAN RHYTHM IN THE LONG-DAY PHOTOPERIODIC INDUCTION OF ERECT AXIS DEVELOPMENT IN THE MARINE RED ALGA NEMALION HELMINTHOIDES1,2

Two isolates of Nemalion helminthoides from the west coast of Ireland showed a heteromorphic life history in which tetraspores formed under short-day conditions on uniaxial, prostrate tetrasporophytes gave rise to uniaxial, prostrate growths similar in morphology to the tetrasporophytes. The induction of multiaxial, erect axes on tetraspore-derived plants was a long-day photoperiodic response. At 11°C, 16:8 h LD, nitrate concentration in the enriched seawater medium had little effect on the numbers of erect exes formed. Induction of erect axes occurred only in daylengths ≥ 12 h, mainly between 7–13° C, and the critical daylength, which generally lies between 14 and 16 h, changed with temperature. Night-breaks (NB) of 1 h light in the middle of 16 h night were ineffective in the promotion of erect axis formation, and day-breaks (DB) of 1 h darkness in the middle of a 16 h day did not inhibit axis formation. Neither continuous light nor NB of 1–1.5 h given at various times during the dark period of an 8:16 h LD cycle promoted the formation of erect axes. At 9° C, equivalent photon exposures (0.69 and 0.34 mol·m^?2) at two different instantaneous photon flux densities resulted in erect axis formation only in the 14 h, 16 h and DB regimes. Photoregimes of 16 h light in combination with dark cycles of various lengths resulted in the maximal promotion of the formation of erect axes in diurnal (22–24 h) and bi-diurnal (44–48 h) cycles, a diminishing response in non-diurnal cycles greater than 24 and 48 h, and a minimal response at 32 h. These data show that the formation of erect axes is a long-day photoperiodic response and provide further evidence for a connection between endogenous circadian rhythms and long-day photoperiodic responses.