Fluence Rates Of Red Light Research Articles

Comparative functional analysis of full‐length and N‐terminal fragments of phytochrome C, D and E in red light‐induced signaling

Phytochromes (phy) C, D and E are involved in the regulation of red/far-red light-induced photomorphogenesis of Arabidopsis thaliana, but only limited data are available on the mode of action and biological function of these lesser studied phytochrome species. We fused N-terminal fragments or full-length PHYC, D and E to YELLOW FLUORESCENT PROTEIN (YFP), and analyzed the function, stability and intracellular distribution of these fusion proteins inplanta. The activity of the constitutively nuclear-localized homodimers of N-terminal fragments was comparable with that of full-length PHYC, D, E-YFP, and resulted in the regulation of various red light-induced photomorphogenic responses in the studied genetic backgrounds. PHYE-YFP was active in the absence of phyB and phyD, and PHYE-YFP controlled responses, as well as accumulation, of the fusion protein in the nuclei, was saturated at low fluence rates of red light and did not require functional FAR-RED ELONGATED HYPOCOTYL1 (FHY-1) and FHY-1-like proteins. Our data suggest that PHYC-YFP, PHYD-YFP and PHYE-YFP fusion proteins, as well as their truncated N-terminal derivatives, are biologically active in the modulation of red light-regulated photomorphogenesis. We propose that PHYE-YFP can function as a homodimer and that low-fluence red light-induced translocation of phyE and phyA into the nuclei is mediated by different molecular mechanisms.

Genetic Linkages Between Circadian Clock-Associated Components and Phytochrome-Dependent Red Light Signal Transduction in Arabidopsis thaliana

The current best candidates for Arabidopsis thaliana clock components are CCA1 (CIRCADIAN CLOCK-ASSOCIATED 1) and its homolog LHY (LATE ELONGATED HYPOCOTYL). In addition, five members of a small family, PSEUDO-RESPONSE REGULATORS (including PRR1, PRR3, PRR5, PRR7 and PRR9), are believed to be another type of clock component. The originally described member of PRRs is TOC1 (or PRR1) (TIMING OF CAB EXPRESSION 1). Interestingly, seedlings of A. thaliana carrying a certain lesion (i.e. loss-of-function or misexpression) of a given clock-associated gene commonly display a characteristic phenotype of light response during early photomorphogenesis. For instance, cca1 lhy double mutant seedlings show a shorter hypocotyl length than the wild type under a given fluence rate of red light (i.e. hypersensitivity to red light). In contrast, both toc1 single and prr7 prr5 double mutant seedlings with longer hypocotyls are hyposensitive under the same conditions. These phenotypes are indicative of linkage between the circadian clock and red light signal transduction mechanisms. Here this issue was addressed by conducting combinatorial genetic and epistasis analyses with a large number of mutants and transgenic lines carrying lesions in clock-associated genes, including a cca1 lhy toc1 triple mutant and a cca1 lhy prr7 prr5 quadruple mutant. Taking these results together, we propose a genetic model for clock-associated red light signaling, in which CCA1 and LHY function upstream of TOC1 (PRR1) in a negative manner, in turn, TOC1 (PRR1) serves as a positive regulator. PRR7 and PRR5 also act as positive regulators, but independently from TOC1 (PRR1). It is further suggested that these signaling pathways are coordinately integrated into the phytochrome-mediated red light signal transduction pathway, in which PIF3 (PHYTOCHROME-INTERACTING FACTOR 3) functions as a negative regulator immediately downstream of phyB.

Overexpression of homologous phytochrome genes in tomato: exploring the limits in photoperception

Transgenic tomato [Lycopersicon esculentum (=Solanum lycopersicum)] lines overexpressing tomato PHYA, PHYB1, or PHYB2, under control of the constitutive double-35S promoter from cauliflower mosaic virus (CaMV) have been generated to test the level of saturation in individual phytochrome-signalling pathways in tomato. Western blot analysis confirmed the elevated phytochrome protein levels in dark-grown seedlings of the respective PHY overexpressing (PHYOE) lines. Exposure to 4 h of red light resulted in a decrease in phytochrome A protein level in the PHYAOE lines, indicating that the chromophore availability is not limiting for assembly into holoprotein and that the excess of phytochrome A protein is also targeted for light-regulated destruction. The elongation and anthocyanin accumulation responses of plants grown under white light, red light, far-red light, and end-of-day far-red light were used for characterization of selected PHYOE lines. In addition, the anthocyanin accumulation response to different fluence rates of red light of 4-d-old dark-grown seedlings was studied. The elevated levels of phyA in the PHYAOE lines had little effect on seedling and adult plant phenotype. Both PHYAOE in the phyA mutant background and PHYB2OE in the double-mutant background rescued the mutant phenotype, proving that expression of the transgene results in biologically active phytochrome. The PHYB1OE lines showed mild effects on the inhibition of stem elongation and anthocyanin accumulation and little or no effect on the red light high irradiance response. By contrast, the PHYB2OE lines showed a strong inhibition of elongation, enhancement of anthocyanin accumulation, and a strong amplification of the red light high irradiance response.

The importance of being red when young: anthocyanins and the protection of young leaves of Quercus coccifera from insect herbivory and excess light

Young leaves of many plants are transiently red because of the accumulation of anthocyanins, with the redness disappearing as leaves mature. Among the many hypothetical functions of foliar anthocyanins, two are tested in this field study: the sunscreen photoprotective function against excess visible light and the handicap signal against herbivory. We took advantage of intraspecies variation in anthocyanin concentrations of young leaves of Quercus coccifera L. to compare in vivo chlorophyll fluorescence parameters, reflectance spectra, total phenolics and the extent of herbivory of leaves of red- and green-leaved phenotypes occupying the same habitat. Photosystem II (PSII) photochemical efficiencies obtained at various photon fluence rates of red light were similar in green and red leaves. In white light, PSII efficiencies were slightly higher in red leaves than in green leaves, indicating a slight photoprotective role of anthocyanins in the field. However, compared with red phenotypes, green phenotypes suffered greater herbivore damage, as judged by the number of leaves attacked and the area lost to herbivory. In addition, there was a positive correlation between the concentrations of anthocyanins and total phenolics. We suggest that the importance of a photoprotective anthocyanic screen is low in thin, young leaves with low chlorophyll concentrations because the green light attenuated by anthocyanins is less significant for chlorophyll excitation. However, the decreased reflectance in the green spectral band and the concomitant leveling of reflectance throughout the 400-570 nm spectral range may either make red leaves less discernible to some insect herbivores or make insect herbivores more discernible to predators, or both. Moreover, excessive herbivory may be additionally discouraged by the high phenolic concentrations in red leaves.

Light modulation of the gravitropic set-point angle (GSA).

A study has been made of the means by which light influences the gravitropic set-point angle (GSA) of the nodes of Tradescantia and the hypocotyls of the lazy-2 mutant of tomato. In light-grown Tradescantia there is a light-regulated developmental change in the GSA with the magnitude of this change being dependent on the photon flux density of white light. The photosynthetic inhibitor DCMU abolished the effect of white light. Low fluence rates of red light had no significant effect on the GSA of Tradescantia: It was concluded that there is an interaction between photosynthesis and the GSA in Tradescantia: The light-induced reduction of the GSA of the hypocotyl of lazy-2 tomato has previously been assumed to be solely an action of light acting via phytochrome. However, it can be shown that the GSA of hypocotyls of lazy-2 seedlings grown in white light is sensitive to DCMU and norflurazon treatment, hence the light effects on the GSA of an organ can be mediated via both phytochrome and photosynthesis. The implication of these findings to the study of gravitropism is discussed.

Irradiance-dependent regulation of gravitropism by red light in protonemata of the moss Ceratodon purpureus.

Apical cells of protonemata of the moss Ceratodon purpureus (Hedw.) Brid. are negatively gravitropic in the dark and positively phototropic in red light. Various fluence rates of unilateral red light were tested to determine whether both tropisms operate simultaneously. At irradiances > or = 140 nmol m-2 s-1 no gravitropism could be detected and phototropism predominated, despite the presence of amyloplast sedimentation. Gravitropism occurred at irradiances lower than 140 nmol m-1 s-1 with most cells oriented above the horizontal but not upright. At these low fluence rates, phototropism was indistinct at 1 g but apparent in microgravity, indicating that gravitropism and phototropism compete at 1 g. The frequency of protonemata that were negatively phototropic varied with the fluence rate and the duration of illumination, as well as with the position of the apical cell before illumination. These data show that the fluence rate of red light regulates whether gravitropism is allowed or completely repressed, and that it influences the polarity of phototropism and the extent to which apical cells are aligned in the light path.

Kinetics of Photobleaching of Protoporphyrin IX in the Skin of Nude Mice Exposed to Different Fluence Rates of Red Light

The purpose of the present study was to determine the kinetics and the fluence rate dependency of the photobleaching of protoporphyrin IX (PpIX) in normal skin of Balb/c nude mice after systemic and topical application of 5-aminolevulinic acid (ALA). ALA was administered systemically (200 mg/kg body weight, i.p.) and topically (20% w/w ALA cream) to the mice. Fluences of up to 40 J/cm2 were delivered by a dye laser (636 nm) at fluence rates of 37.5, 75, 150, 300 and 500 mW/cm2. The photo-bleaching rate was constant within this range of fluence rates. This result suggests that there is no oxygen effect for PpIX photobleaching in this region for the skin of Balb/c nude mice. During light exposure the fluorescence decay followed neither first- nor second-order kinetics. The decay rate was slightly faster after systemic application than after topical application of ALA, but did not depend on the time (1-8 h) between application and analysis.

Kinetics of Photobleaching of Protoporphyrin IX in the Skin of Nude Mice Exposed to Different Fluence Rates of Red Light

Kinetics of Photobleaching of Protoporphyrin IX in the Skin of Nude Mice Exposed to Different Fluence Rates of Red Light

Photocontrol of anthocyanin biosynthesis in tomato.

Juvenile anthocyanin biosynthesis has been studied in dark-grown seedlings of tomato (Lycopersicon esculentum Mill.) wild types (WTs) and photomorphogenic mutants. During a subsequent 24-hr period of monochromatic irradiation at different fluence rates of red light (R) the fluence-rate response relationships for induction of anthocyanin in all the WTs are similar, yet complex, showing a response at low fluence rates (LFRR) followed by a fluence rate-dependent high irradiance response (HIR). In the hypocotyl this response is restricted to the sub-epidermal layer of cells. The high-pigment-1 (hp-1) mutant exhibits a strong amplification of both response components. Theatroviolacea (atv) mutant shows strongest amplification of the HIR component. In contrast, a transgenic line overexpressing an oat phytochrome A gene (PHYA3 (+)) shows a most dramatic amplification of the LFRR component. The far-red light (FR)-insensitive (fri) mutant, deficient in phytochrome A (phyA), lacks the LFRR component whilst retaining a normal HIR. The temporarily R-insensitive (tri) mutant, deficient in phytochrome B1 (phyB1) retains the LFRR, but lacks the HIR. Thehp-1,fri andhp-1,tri double mutant, exhibit amplified, yet qualitatively similar responses to the monogenicfri andtri mutants. Thefri,tri double mutant lacks both response components in R, but a residual response to blue light (B) remains. Similarly, theaurea (au) mutant deficient in phytochrome chromophore biosynthesis and presumably all phytochromes, lacks both response components in the R and FR regions of the spectrum. Experiments at other wavelengths demonstrate that while there is only a small response in the FR spectral region (729 nm) in tomato, there is an appreciable HIR response in the near FR at 704 nm, which is retained in thetri mutant. This suggests that the labile phyA pool participates in the HIR at this wavelength. The intense pigmentation (Ip) mutant appears to be specifically deficient in the B1 induced anthocyanin biosynthesis. Adult plants, grown under fluorescent light/dark cycles, show a reduction of anthocyanin content of young developing leaves upon application of supplemtary or end-of-day FR. The involvement of different phytochrome species in anthocyanin biosynthesis based on micro-injection studies into theau mutant and studies using type specific phytochrome mutants is discussed.

Calcium-dependent membrane depolarisation activated by phytochrome in the moss Physcomitrella patens

In caulonemal filaments of Physcomitrella patens which had been preincubated in the dark for 24 h, irradiation with red light (640 nm, fluence rate 85 μmol · m−2 · s−1) evoked (i) the development of side branch initials and (ii) a rapid, but transient, depolarisation of the plasma membrane by 90 ± 13 mV from a resting potential of -178 ± 13 mV. This was followed by a transient hyperpolarisation to a value 21± 8 mV more negative than the original membrane potential. The refractory period for the transient depolarisation was between 12 and 15 min. The fluence rate of red light required to evoke maximal depolarisation was about 80 μmol · m−2 · s−1 for a 1-min pulse. At this fluence rate, a depolarising response could be recorded for pulse lengths as small as 7 s. The transient depolarisation was insensitive to 3-(3′,4′dichlorophenyl)-1,1-dimethyl urea (DCMU) and was unchanged in plants bleached by growth on norflurazon (SAN 9789). Furthermore, the electrical response could be blocked by simultaneous application of far-red light. These results suggest the involvement of the photoreceptor phytochrome in the response. Removing Ca2+ from the external medium or replacing Ca2+ with Mg2+ blocked the depolarisation. The depolarisation could also be blocked by the K+ channel-blocker tetraethylammonium (10 mM) and the Cl− channel-blocker niflumic acid (1 μM). Conversely, although calcium channel-antagonists such as nifedipine and lanthanides, applied at a concentration of 100 μM, also altered the response, they did not block it. A possible ionic mechanism for the membrane potential transient is advanced, and the physiological significance discussed in the context of early events in the phytochrome signalling pathway.

Close correspondence between the action spectra for the blue light responses of the guard cell and coleoptile chloroplasts, and the spectra for blue light-dependent stomatal opening and coleoptile phototropism.

Fluorescence spectroscopy was used to characterize blue light responses from chloroplasts of adaxial guard cells from Pima cotton (Gossypium barbadense) and coleoptile tips from corn (Zea mays). The chloroplast response to blue light was quantified by measurements of the blue light-induced enhancement of a red light-stimulated quenching of chlorophyll a fluorescence. In adaxial (upper) guard cells, low fluence rates of blue light applied under saturating fluence rates of red light enhanced the red light-stimulated fluorescence quenching by up to 50%. In contrast, added blue light did not alter the red light-stimulated quenching from abaxial (lower) guard cells. This response pattern paralleled the blue light sensitivity of stomatal opening in the two leaf surfaces. An action spectrum for the blue light-induced enhancement of the red light-stimulated quenching showed a major peak at 450 nm and two minor peaks at 420 and 470 nm. This spectrum matched closely an action spectrum for blue light-stimulated stomatal opening. Coleoptile chloroplasts also showed an enhancement by blue light of red light-stimulated quenching. The action spectrum of this response, showing a major peak at 450 nm, a minor peak at 470 nm, and a shoulder at 430 nm, closely matched an action spectrum for blue light-stimulated coleoptile phototropism. Both action spectra match the absorption spectrum of zeaxanthin, a chloroplastic carotenoid recently implicated in blue light photoreception of both guard cells and coleoptiles. The remarkable similarity between the action spectra for the blue light responses of guard cells and coleoptile chloroplasts and the spectra for blue light-stimulated stomatal opening and phototropism, coupled to the recently reported evidence on a role of zeaxanthin in blue light photoreception, indicates that the guard cell and coleoptile chloroplasts specialize in sensory transduction.

The inhibitor of zeaxanthin formation, dithiothreitol, inhibits blue-light-stimulated stomatal opening in Vicia faba

Dithiothreitol (DTT), an inhibitor of violaxanthin de-epoxidation and zeaxanthin formation in chloroplasts, inhibited blue-light-stimulated stomatal opening in epidermal peels of Vicia faba L. in a concentration-dependent fashion. Complete inhibition was observed at 3 mM DTT. The DTT effect was specific for the stomatal response to blue light, and the red-light-stimulated opening, which depends on photosynthetic reactions in the guard cells, was unaffected. Preirradiation of stomata in epidermal peels with increasing photon fluence rates of red light, prior to an incubation in 10 μmol·m-2·s-1 of blue light and 100 μmol·m-2·s-1 red light, resulted in a DTT-sensitive, blue-light-stimulated opening that was proportional to the fluence rate of the red light pre-treatment. Guard cells in epidermal peels and guard-cell protoplasts irradiated with red light showed increases in their zeaxanthin content that depended on the fluence rate of red light, or on the incubation time. The increases in zeaxanthin concentration were inhibited by DTT. The obtained results indicate that zeaxanthin could function as a photoreceptor mediating the stomatal responses to blue light.

Phytochrome-mediated light signal transmission to the phosphorylation of proteins in the plasma membrane and the soluble fraction of etiolated pea stem sections

The excised third internodes of etiolated pea seedlings were irradiated with red light (660 nm) for 40 s changing the intensity from 0, 10, 50 and 100 μmol m −2 s −1. The tissue was ground in a buffer and the plasma membrane was purified. The phosphorylation of plasma membrane proteins with molecular masses of 15, 50, 70 and 120 kDa from adenosine 5′[γ- 32P]triphosphate ([γ- 32P]ATP) at 0 °C for 7 s increased with increasing fluence rate of red light from 10 to 50 μmol m −2 s −1. However, the phosphorylation of these proteins was less enhanced at red-light fluence rate of 100 μmol m −2 s −1. Successive irradiation of the third internode by red (100 μmol m −2 s −1) for 20 s and far-red light (730 nm; 4μmol m −2 s −1) for 30 s reversed the phosphorylation of the 15 kDa protein. The 32P phosphoryl group of the 15 kDa protein which was received from [γ- 32P]ATP at 0 °C for 7 s was removed efficiently by subsequent addition at 0 °C for 5 s of cold 10 −6 M adenosine 5′-diphosphate or guanosine 5′-diphosphate and also 10 −5 M ATP or guanosine 5′-triphosphate. The phosphorylation of the 15 kDa protein in the soluble fraction showed a similar light response to that associated in the plasma membrane.

Light-Stimulated Cotyledon Expansion in the blu3 and hy4 Mutants of Arabidopsis thaliana

Cotyledon expansion in response to blue light was compared for wild-type Arabidopsis thaliana (L.) Heynh. and the mutants blu3 and hy4, which show reduced inhibition of hypocotyl growth in blue light. White, blue, and red light stimulated cotyledon expansion in both intact and excised cotyledons of wild-type seedlings (ecotypes No-0, WS, Co-0, La-er). Cotyledons on intact blu3 and hy4 seedlings did not grow as well as those on the wild type in response to blue light, but pretreatment of blu3 seedlings with low fluence rates of red light increased their responsiveness to blue light. Excision of cotyledons alleviated the mutant phenotype so that both mutant and wild-type cotyledons grew equally well in blue light. The loss of the mutant cotyledon phenotype upon excision indicates that the blu3 and hy4 lesions affect cotyledon expansion indirectly via a whole-plant response to light. Furthermore, the ability of excised, mutant cotyledons to grow normally in blue light shows that this growth response to blue light is mediated by a photosystem other than the ones impaired by the blu3 and hy4 lesions.

Sugar and Organic Acid Accumulation in Guard Cells of Vicia faba in Response to Red and Blue Light.

Changes in neutral sugar and organic acid content of guard cells were quantitated by high-performance liquid chromatography during stomatal opening in different light qualities. Sonicated Vicia faba epidermal peels were irradiated with 10 [mu]mol m-2 s-1 of blue light, a fluence rate insufficient for the activation of guard cell photosynthesis, or 125 [mu]mol m-2 s-1 of red light, in the presence of 1 mM KCl, 0.1 mM CaCl2. The low-fluence-rate blue light stimulated an average net stomatal opening of 4.7 [mu]m in 2 h, whereas the saturating fluence rate of red light stimulated an average net opening of 3.8 [mu]m in 2 h. Under blue light, the malate content of guard cells increased to 173% of the initial level during the first 30 min of opening and declined as opening continued. Sucrose levels continuously rose throughout the blue light-stimulated opening, reaching 215% of the initial level after 2 h. The starch hydrolysis products maltose and maltotriose remained elevated at all times. Under red light, guard cells showed very little increase in organic acid or maltose levels, whereas sucrose levels increased to 208% of the initial level after 2 h. Total measured organic metabolite concentrations were correlated with stomatal apertures in all cases except where substantial malate increases occurred. These results support the hypothesis that light quality modulates alternative mechanisms of osmotic accumulation in guard cells, including potassium uptake, photosynthetic sugar production, and starch breakdown.

PHOTORECEPTORS CONTROLLING TRANSCRIPTION OF rbcS GENES IN GREEN LEAF TISSUE OF Pisum sativum

We have investigated the photoreceptors controlling transcription of genes encoding the small subunit (rbcS) of ribulose 1,5-bisphosphate carboxylase-oxygenase in green leaf tissue of pea (Pisum sativum). RbcS transcription was measured by hybridising labelled transcripts of isolated nuclei to rbcS cDNA clones. Transfer of green Pisum leaf tissue to darkness for 5 h causes a substantial decrease in the rate of rbcS transcription and the rate is restored rapidly when the plants are returned to white light. Low fluence rates of red light are ineffective in restoring the rate of rbcS transcription, suggesting that phytochrome alone does not fully mediate the response. Blue light is similarly effective to white light of an equal fluence rate (120 mumol m-2 s-1) in restoring the rate of rbcS transcription in the dark-treated plants, indicating that a blue light photoreceptor is involved. However, red light at the same fluence rate produces about 65% of the effect of blue or white light, showing that the blue light photoreceptor is not the only photoreceptor controlling rbcS transcription in the green leaf tissue. The identity of the photoreceptor responsible for the red light effect is discussed. Similar effects of blue and red light are observed at the level of transcript abundance in dark-grown pea leaf tissue given a brief illumination with red light, which potentiates the tissue for rapid transcript accumulation in white light.

THE BLUE LIGHT RESPONSE OF STOMATA: PULSE KINETICS AND SOME MECHANISTIC IMPLICATIONS*

Abstract— Intact leaves of Commelina communis irradiated with high fluence rates of red light, showed discrete increases in stomatal conductance in response to pulses (1‐100 s) of blue light (250 μmol m−2 s−1). Red light pulses were ineffective, indicating that the conductance increases were not mediated by photosynthesis and that they constitute a specific stomatal response to blue light. The response peaked 15 min after the pulse and was completed within50–60 min. Conductance increases were proportional to pulse duration up to about 30 s and saturated at longer exposures. The relationship between stomatal responses and pulse duration approximately fitted an exponential function, with a t 9s. Pulse responses at two different fluences indicated that reciprocity held. Responses to two consecutive pulses varied with time between pulses. A saturating pulse applied immediately after a preceding one induced no additional response; two saturating pulses 50 min apart caused two identical, consecutive responses. Total increases in conductance induced by two pulses separated by intermediate time intervals increased with time between pulses with a = 9 min. These results point to a blue light‐dependent photoconversion of a molecular form, with the activity of the photoconversion product decaying in a thermal reaction. Under continuous blue light, prevailing fluence rates and rates of the light and thermal reactions are postulated to determine steady‐state activities of the photoconversion product and proportional increases in conductance levels. These findings have implications for the environmental and metabolic roles of the stomatal response to blue light.

Herbicides which Interfere with the Biosynthesis of Carotenoids and Their Effect on Pigment Excitation, Chlorophyll Fluorescence and Pigment Composition of the Thylakoid Membrane

Plants grown in the presence of the herbicides assayed synthesized chlorophylls during growth at low fluence rates. Subsequent irradiation with higher fluence rates of red light induced a strong chlorosis with SAN 6706 being a much stronger herbicide than J 852 or amino-triazole. All herbicides assayed also changed the content and composition of chlorophylls, carotenoids and pigment-protein-complexes of the thylakoid membrane and therefore the pigment excitation and chlorophyll fluorescence emission spectra of the plastid. With increasing herbicide toxicity the main characteristic emission bands at 690 and 730 nm disappeared and new emission bands at 715 (J 852) and 700 nm (SAN 6706) appeared. Such “artificial” membranes with a changed pigment composition were very susceptible to light. Presented data may be taken as evidence, that the lack of photoprotective cyclic carotenoids caused by the specific action of a bleaching herbicide is the primary event that may lead to a disturbed formation of the thylakoid membrane and its destruction by light and oxygen.

Phototropism and polarotropism of primary chloronemata of the moss Physcomitrella patens: responses of the wild-type

Primary chloronemata growing from germinated spores of the moss Physcomitrella patens adopt one of two preferred polarotropic orientations depending on the wavelength and photon fluence rate of monochromatic light. Growth is mainly parallel to the electrical vector of plane polarised light in blue light and higher fluence rates of red light, and perpendicular to the electrical vector in the green and far-red regions of the spectrum and in low fluence rates of red light. The transition between the two polarotropic orientations, at wavelengths where it can be observed, usually occurs over a narrow range of fluence rates, and at this point the filaments do not grow randomly but tend to adopt in approximately equal numbers one of the preferred directions of growth. The primary chloronemata are positively phototropic in far-red light and in red light of low fluence rates, but tend to grow at right angles to the incident light in high fluence rates of red light. Simultaneous illumination with a high fluence rate of red light and a low fluence rate of far-red light causes a marked increase in the percentage of filaments growing towards the red light source at the expense of those growing at right angles to it, supporting the hypothesis that in red and far-red light, at least, the responses are controlled by the photoequilibrium of a phytochrome pool.

Phototropism and polarotropism of primary chloronemata of the moss Physcomitrella patens: responses of mutant strains

The phototropic and polarotropic responses of primary chloronemata grown from germinated minated spores of three mutant strains of the moss, Physcomitrella patens, have been studied and compared with those of the wild-type. The mutants and wild-type show the same qualitative tropic responses but differ with respect to the light conditions under which they are expressed. In both the wild-type and mutants the responses are controlled by phytochrome. In monochromatic red light, at low fluence rates, wild-type primary chloronemata grow positively phototropically in unidirectional light or perpendicular to the electrical vector (E) in polarised light; at high fluence rates growth in unidirectional light is lateral to the incident light or, in polarised light, parallel to E. The mutants, however, show only the lateral phototropic or parallel polarotropic responses at all fluence rates of red light tested. In far-red light, the wild-type primary chloronemata adopt a positive phototropic or a perpendicular polarotropic response; the mutants show the same responses but in a lower percentage of filaments. These results and those at other wavelengths indicate either that the mutants are impaired in their ability to adopt the positive phototropic and perpendicular polarotropic responses or that in the mutants the transition between the "low light" (positive phototropic-perpendicular polarotropic) and the "high light" (lateral phototropic-parallel polarotropic) responses is shifted to a lower photon fluence rate. Possible explanations of this phenotypic difference are discussed.