PHYTOCHROME ACTION IN ORYZA SATIVA L.: I. GROWTH RESPONSES OF ETIOLATED COLEOPTILES TO RED, FAR-RED AND BLUE LIGHT

1. Under continuous irradiation, the growth of intact rice coleoptilewas strongly inhibited by red light, and somewhat preventedby blue and far-red light. The inhibitory effect of red lighton coleoptile elongation was caused by a low-energy brief irradiation,and a single exposure of 1.5 kiloergs cm^–2 incidentenergy of red light brought about the 50% inhibition. This photoinhibitionof growth was observed only after the coleoptile had elongatedto about 10 mm or longer. The red light-induced effect was reversedby an immediately following brief exposure to far-red light,and the photoresponses to red and far-red light were repeatedlyreversible. The escape reaction of red lightinduced effect tookplace at a rate so that 50% of the initial reversibility waslost within 9 hr in darkness at 27. The inhibition by bluelight and reversal by far-red irradiation was also achievedrepeatedly with successive treatments of the coleoptiles. Theevidence for a low intensity red far-red reversible controlof coleoptile growth, indicative of control by phytochrome,seems clearly established in etiolated intact seedlings.
2. Incontrast, the elongation of apically excised rice coleoptilesegments was promoted by a brief exposure to red light in 0.02M phosphate buffer, pH 7, and the effect was almost completelynullified by an immediately subsequent exposure to far-red light.It becomes evident that the growth of intact coleoptiles wasinhibited by a exposure to red light, while that of excisedsegments in a buffer was rather promoted by red irradiation.The direction of red light induced responses, either promotiveor inhibitory, depends upon the method of bioassay using intactcoleoptiles or their excised segments.

(Received July 24, 1967; )     

Phytochrome action inOryza sativa L. VI. Red far-red reversible effect on early development of coleoptiles

When 3–4 mm long coleoptiles of etiolated rice seedlings (cv. Koshijiwase) were irradiated with continuous red light their growth was seriously inhibited. If a brief exposure of red light (4×10³ ergs cm⁻²) was given to the short coleoptiles, the growth rate dropped immediately after the irradiation, but the growth did not stop till the coleoptile reached some calculated length. If another brief red irradaition of the same order was given 24 hr after the first, the growth rate and the final length dropped further. The effect of red light was reversed by successively given far-red light, and this response was repeatedly red and far-red reversible. The escape reaction was rather slow so that photoreversibility was not lost at all by 8th hr, and 50% of the initial reversibility was lost within ca. 16 hr at 25±0.5 C. Blue light also induced the inhibition of coleoptile elongation, the effect was reversed by subsequent far-red irradiation, and this could be obtained repeatedly. Thus, the photoinhibition of the young coleoptile can be concluded to be under the control of phytochrome, and the mode of action appeared quite different from the previously reported results with longer coleoptiles.     

The Effects of Red, Far-red, and Blue Light on the Geotropic Response of Coleoptiles of Zea mays

The effects of red, far-red, and blue light on the geotropicresponse of excised coleoptiles of Zea mays have been investigated.Seedlings were grown in darkness for 5 or 6 days, exposed tovarious light treatments, and then returned to darkness fordetermination of the geotropic response. The rate of response of the coleoptiles is decreased after theyhave been exposed to red light (620–700 mµ, 560ergs cm^–2sec^–1 for the 24 hrs, but not for the 4hrs, preceding stimulation by gravity. Furthermore, their rateof response is greatly reduced if they are exposed to red lightfor 10 min and then returned to darkness for 20 hrs before geotropicstimulation. At 25° C an interval of 6 to 8 hrs elapses between a 10-minexposure to red light and the first detectable decrease in thegeotropic response of the coleoptile. This interval can be lengthenedby exposing the seedlings to low temperatures (0° to 2°C) after the light treatment but cannot be greatly shortenedby increasing the duration of exposure to red light. Using a standard procedure of exposing 5-day-old etiolated seedlingsto light for various times, replacing them in darkness for 20hrs and then determining the response of the coleoptiles to4 hrs geotropic stimulation, it has been found that: (a) Exposureto red light for 15 sec significantly decreases the geotropiccurvature of the coleoptiles and that further reduction occurson increasing the length of the light treatment to 2 and 5 min.(b) Far-red light has no effect on the geotropic response ofthe coleoptiles but it can completely reverse the effect ofred light. After repeated alternate exposure to red and far-redlight the geotropic response of the coleoptile is determinedby the nature of the last exposure, (c) Complete reversal ofthe effect of red light by far-red radiation only occurs whenexposure to far-red follows immediately after exposure to red.The reversing effect of far-red radiation is reduced if a periodof darkness intervenes between the red and far-red light treatments,and is lost after a dark interval of approximately 2 hrs. The effect of red light on the rate of geotropic response ofthe coleoptiles is independent of their age and length at thetime of excision. Blue light acts in a similar way to red light, but the seedlingsare less sensitive to blue than to red light. Coleoptiles grown throughout in a mixture of continuous, weak,red, and far-red light have a lower rate of geotropic responsethan etiolated coleoptiles.     

Photobiology of phytochrome-mediated growth responses in sections of stem tissue from etiolated oats and corn

The far-red reversibility of the phytochrome-controlled stimulation of elongation of coleoptile sections by low fluence red light has been characterized in subapical coleoptile sections from dark-grown Avena sativa L., cv Lodi seedlings. The fluence dependence of the far-red reversal was the same whether or not the very low fluence response is also expressed. The capacity of far-red light to reverse the red light-induced response began to decline if the far-red light was given more than 90 minutes after the red irradiation. Escape was complete if the far red irradiation was given more than 240 minutes after the red irradiation. Sections consisting of both mesocotyl and coleoptile tissue from dark-grown Avena seedlings were found to have physiological regulation of the very low fluence response by indole 3-acetic acid and low external pH similar to that seen for sections consisting entirely of coleoptile tissue. The fluence-dependence of the red light-induced inhibition of mesocotyl elongation was studied in mesocotyl sections from dark grown Zea mays L. hybrid T-929 seedlings. Ten micromolar indole 3-acetic acid stimulates the control elongation of the sections, while at the same time increasing the sensitivity of the tissue for the light-induced inhibition of growth by a factor of 100.     

High-irradiance responses induced by far-red light in grass seedlings of the wild type or overexpressing phytochrome A

The occurrence of phytochrome-mediated highirradiance responses (HIR), previously characterised largely in dicotyledonous plants, was investigated in Triticum aestivum L., Zea mays L., Lolium multiflorum Lam. and in both wild-type Oryza sativa L. and in transgenic plants overexpressing oat phytochrome A under the control of a 35S promoter. Coleoptile growth was promoted (maize, ryegrass) or inhibited (wild-type rice) by continuous far-red light (FRc). However, at equal fluences, hourly pulses of far-red light (FRp) were equally effective, indicating that the growth responses to FRc were not true HIR. In contrast, in maize and rice, FRc increased anthocyanin content in the coleoptile in a fluence-rate dependent manner. This response was a true HIR as FRp had reduced effects. In maize, anthocyanin levels were significantly higher under FRc than under continuous red light. In rice, overexpression of phytochrome A increased the inhibition of coleoptile growth and the levels of anthocyanin under FRc but not under FRp or under continuous red light. The effect of FRc was fluence-rate dependent. In light-grown rice, overexpression of phytochrome A reduced leaf-sheath length, impaired the response to supplementary far-red light, but did not affect the response to canopy shade-light. In grasses, typical HIR, i.e. fluence-rate dependent responses showing reciprocity failure, can be induced by FRc. Under FRc, overexpressed phytochrome A operates through this action mode in transgenic rice.Abbreviations FR far-red light - FRc continuous far-red light - FRp pulses of far-red light - HIR high-irradiance responses - LFR low-fluence responses - OPHYA transgenic rice overexpressing oat phytochrome A - Pfr far-red light-absorbing form of phytochrome - phyA phytochrome A - R red light - Rc continuous red light - VLFR very low-fluence responses - WT wildtype We thank Marcelo J. Yanovsky for his help with the photographs and Professor Rodolfo A. Sanchez for providing a reprint of the paper by P.J.A.L. de Lint. This work was supported by grants from UBA (AG041) and Fundacion Antorchas (A-13218/1-15) to J.J.C.     

Ethylene and the Responses to Light of Rice Seedlings

The growth of rice seedlings (Oryza satira L.) in the presence of ethylene caused a change in the response to light of coleoptile elongation. In plants grown in air without added ethylene coleoptile elongation was promoted by red, far-red and yellow-green light only in very young seedlings; in older plants irradiation inhibited the growth of the coleoptile. The effect of growing plants in the presence of ethylene was to prolong the period during which light promoted coleoptile growth. Elongation of the first internode was inhibited by light whether or not the seedlings were grown in the presence of ethylene. A correlation existed between the growth effect of an irradiation and the initial decay rate of phytochrome which was established by the treatment. Regardless of wave length, light sources whose intensities were adjusted to produce a decay rate of about 10% per hour or less induced a moderate rate of coleoptile elongation which persisted for a relatively long period. Irradiation with red or yellow-green light of higher intensity which produced a higher rate of phytochrome decay induced a higher rate of coleoptile elongation, but growth stopped after several hours. Other observations, however, showed that one cannot establish a general simple correlation between the rate of elongation of rice coleoptiles under light and the status of measurable phytochrome in the plant.     

An unusual effect of the far-red absorbing form of phytochrome: Photoinhibition of seed germination inBromus sterilis L.

Seeds ofBromus sterilis L. germinated between 80–100% in darkness at 15° C but were inhibited by exposure to white or red light for 8 h per day. Exposure to far-red light resulted in germination similar to, or less than, that of seeds maintained in darkness. Germination is not permanently inhibited by light as seeds attain maximal germination when transferred back to darkness. Germination can be markedly delayed by exposure to a single pulse of red light following 4 h inhibition in darkness. The effect of the red light can be reversed by a single pulse of far-red light indicating that the photoreversible pigment phytochrome is involved in the response. The response ofB. sterilis seeds to light appears to be unique; the far-red-absorbing form of phytochrome (Pfr) actually inhibiting germination.Abbreviations Pr red absorbing form of phytochrome - Pfr far-red absorbing form of phytochrome     

Effects of red light on the growth of intact wheat and barley coleoptiles

The final lengths of intact dark-grown coleoptiles vary with species and cultivar. The growth distribution pattern in the apical 25-mm growing zone and the absolute amount of growth in each zone depend on the age and species of the coleoptile. A comparative study of several cultivars of wheat, Triticum vulgare, and barley, Hordeum vulgare, indicates that the growth distribution pattern in 30- to 38-mm coleoptiles varies with the species and cultivar. In barley, there are two patterns of growth distribution among the several cultivars, whereas in wheat, all cultivars exhibit a common zonal growth pattern. The total growth of coleoptiles, initially 30 to 38 mm in length, during a 24-hour dark incubation period is the same in dark-grown coleoptiles as in those irradiated with 3 minutes of red (660 nm) light prior to the incubation period. The growth distribution pattern in the growing zone of this 30- to 38-mm coleoptile is, however, altered by red light. Growth of the apical 5-mm zone is stimulated by red light and the zonal growth 5 to 10 mm below the apex is only slightly affected, whereas growth in the zones 10 to 15 to 20, and 20 to 25 mm below the apex is inhibited. This growth distribution pattern in irradiated coleoptiles changes as the coleoptile increases in length. The response of a zone following exposure to red light is dependent upon the age of the seedlings irradiated. The over-all effect of red light on growth of the intact coleoptile varies with the length of the coleoptile. In young seedling 20 to 29 mm in length, the cells of the coleoptile can compensate for the effects of red light, with the over-all growth of the dark-grown and irradiated coleoptile about the same. As the seedling grows older, the cells of the coleoptile can no longer make up for the effects of red light, and the over-all effect changes from compensation to pronounced inhibition.     

Biphasic effect of red light on the growth of coleoptiles in etiolated barley seedlings

InHordeum vulgare cultivar “Kirin-choku No. 1”, the final length of intact coleoptiles of totally etiolated seedlings was approximately twice as long as that of those grown under continuous red light. The fluence response curve of the latter was biphasic; the low-energy effect was saturated by red light of ca. 50 J m⁻² which gave rise to about 40% of the maximum inhibition by continuous irradiation with red light of 1.2 W m⁻², whereas the high-energy effect was induced by irradiation for 1 hr or longer. Coleoptiles of 3-day-old seedlings were most sensitive to light causing the low-energy effect, which was repeatedly red/far-red reversible. The growth inhibition was correlated to the photometrically measured percentage of Pfr so that the maximum effect was induced by red light of 50 J m⁻² which transformed 70% of phytochrome to Pfr in the coleoptile tip. Wavelength dependence of the high-energy effect showed that monochromatic light of 400, 600 and 650 nm greatly inhibited the coleoptile growth, whereas light of 700 and 750 nm promoted it instead. The effect was also induced by intermittent irradiation with red light, and the more frequently the intermittent treatment was given, the more the growth was inhibited.     

Phytochrome-controlled extension growth of Avena sativa L. seedlings

The effects of continuous red and far-red light and of brief light pulses on the growth kinetics of the mesocotyl, coleoptile, and primary leaf of intact oat (Avena sativa L.) seedlings were investigated. Mesocotyl lengthening is strongly inhibited, even by very small amounts of P_fr, the far-red light absorbing form of phytochrome (e.g., by [P_fr]0.1% of total phytochrome, established by a 756-nm light pulse). Coleoptile growth is at first promoted by P_fr, but apparently inhibited later. This inhibition is correlated in time with the rupturing of the coleoptile tip by the primary leaf, the growth of which is also promoted by phytochrome. The growth responses of all three seedling organs are fully reversible by far-red light. The apparent lack of photoreversibility observed by some previous investigators of the mesocotyl inhibition can be explained by an extremely high sensitivity to P_fr. Experiments with different seedling parts failed to demonstrate any further obvious interorgan relationship in the light-mediated growth responses of the mesocotyl and coleoptile. The organspecific growth kinetics, don't appear to be influenced by P_fr destruction. Following an irradiation, the growth responses are quantitatively determined by the level of P_fr established at the onset of darkness rather than by the actual P_fr level present during the growth period.Abbreviation P_fr far-red light absorbing form of phytochrome     

Photocontrol of stem elongation in light-grown plants of Fuchsia hybrida

Stems of the caulescent long-day plant, Fuchsia hybrida cv Lord Byron, showed 2 types of response to light. In one, internode length was increased by far-red irradiation given at the end of an 8 h photoperiod: the response was no greater with prolonged exposure and was less when the start of far-red was delayed. The effect of far-red was reversible by a subsequent exposure to red light. Internode length was inversely proportional to the P_fr/P ratio established before entry to darkness and there was no evidence for loss of P_fr during a 16 h dark period. The inhibitory effect of P_fr acted at a relatively late stage of internode growth. With the development of successive internodes a second response appeared in which stems lengthened following prolonged daily exposures to red or far-red light, or mixtures of the two, or to brief breaks with red or white light. In these later internodes, a short exposure to far-red near the middle of the night was not reversible by red because red alone promoted elongation at this time. Internode length increased with increase in the daily duration of light and, when light was given throughout an otherwise dark period of 16 h, with increase in illuminance to a saturation value of 200 lx from tungsten lamps. Elongation increased as a linear function of decrease in photostationary state of phytochrome down to P_fr/P0.3; however, internodes were shorter in far-red light than in 25% red/red+far-red. It was concluded that stem length is a net response to two modes of phytochrome action. An inductive effect of P_fr inhibits a late stage in internode expansion, and a phytochrome reaction which operates only in light (and may involve pigment cycling) promotes an early stage of internode development. Stem elongation is thus a function both of the daily duration of light and its red/red+far-red content. The outgrowth of axillary buds was controlled by the first type of phytochrome action only.Abbreviations and symbols FR far red light - R red light - P phytochrome - P_fr phytochrome in the far-red light absorbing form - SD 8 h short days - LDP long-day plant - SDP short-day plant     

Photocontrol of petiole elongation in light-grown strawberry plants

Summary The mode of phytochrome control of elongation growth was studied in fully-green strawberry (Fragaria x Ananassa Duch.) plants. Petiole growth showed two distinct types of response to light. In one, the end-of-day response, petioles were lengthened by low-intensity far-red irradiation for 1 h immediately following the 8 h photoperiod. The response was little or no greater with prolonged exposure and less when the start of far-red was delayed. It was already evident in the first leaf to emerge after treatment began. With the development of successive leaves a second, photoperiodic, type of response appeared, in which petioles lengthened following only prolonged exposure to red, far-red, mixtures of the two, or tungsten lighting, all at low levels of intensity. As with the inhibition of flowering in previous experiments, irradiation with red light during the second half of the otherwise long dark period gave the greatest response.Abbreviations and Symbols FR far-red light - HIR high irradiance response - R red light - P_r phytochrome in the red light absorbing form - P_fr phytochrome in the far-red light absorbing form - SDP short-day plant - LDP long-day plant - PAR photosynthetically active radiation     

Phytochrome inhibits the effectiveness of gibberellins to induce cell elongation in rice

Red light controls cell elongation in seedlings of rice (Oryza sativa L.) in a far-red-reversible manner (Nick and Furuya, 1993, Plant Growth Regul. 12, 195–206). The role of gibberellins and microtubules in the transduction of this response was investigated in the rice cultivars Nihon Masari (japonica type) and Kasarath (indica type). The dose dependence of mesocotyl elongation on applied gibberellic acid (GA₃) was shifted by red light, and this shift was reversed by far-red light. In contrast, coleoptile elongation was found to be independent of exogenous GA₃. Nevertheless, it was inhibited by red light, and this inhibition was reversed by far-red light. The content of the active gibberellin species GA₁ and GA₄ was estimated by radio-immunoassay. In the mesocotyl, the gibberellin content per cell was found to increase after irradiation with red light, and this increase was far-red reversible. Conversely, the cellular gibberellin content in japonica-type coleoptiles did not exhibit any significant light response. Microtubules reoriented from transverse to longitudinal arrays in response to red light and this reorientation could be reversed by subsequent far-red light in both the coleoptile and the mesocotyl. This movement was accompanied by changes in cell-wall birefringence, indicating parallel reorientations of cellulose deposition. The data indicate that phytochrome regulates the sensitivity of the tissue towards gibberellins, that gibberellin synthesis is controlled in a negative-feedback loop dependent on gibberellin effectiveness, and that at least two hormone-triggered signal chains are linked to the cytoskeleton in rice.Abbreviations D darkness - FR far-red light - GA₃ gibberellic acid - GC-SIM gas chromatography-selected ion monitoring - R red light This work was supported by a grant of the Human Frontier Science Organization to P.N. Advice and organizational support by Prof. M. Furuya (Hitachi Advanced Research Laboratory, Hatoyama, Japan) and Prof. N. Murofushi (Department of Agricultural Chemistry, University of Tokyo, Japan) is gratefully acknowledged. Seeds of both rice cultivars were kindly provided by Dr. O. Yatou (Institute for Radiation Breeding, Hitachi-Ohmiya, Japan), and the antiGA₁ Me-antiserum for the radio-immunoassays by Dr. I. Yamaguchi (Department of Agricultural Chemistry, University of Tokyo, Japan).     

Interactions of microtubule disorganizers, plant hormones, and red light in wheat coleoptile segment growth

Growth response of coleoptile segments excised from 3-day-old seedlings of wheat (Triticum vulgare cv. Baart) to gibberellic acid, indoleacetic acid, and 2,4-dichlorophenoxyacetic acid, to red light, and to several microtubule disorganizers depends on the initial position of the excised segment in the intact coleoptile. Red light, 660 nm, stimulates the growth of the apical cells, but inhibits markedly the growth of the cells in the basal region of the coleoptile. The effects of red light are independent of sucrose, gibberellic acid, indoleacetic acid, and 2,4-dichlorophenoxyacetic acid, even though these substances themselves markedly affect the growth of the coleoptile segments. Concentractions of the microtubule disorganizers, vinblastine sulfate, cupric chloride, urea, and colchicine, which do not alter significantly the growth of the dark control apical segments, substantially repress the promotive effects of red light or auxin on the increase in length of the apical cells of the coleoptile. This suggests that stimulation by red light and by auxin involves microtubule production. Microtubule disorganizers repress the growth of elongating cells of the coleoptile, yet on the other hand, auxin and irradiation do not alter significantly the response of basal cells to the microtubule disorganizing agents. We hypothesized that light and growth regulators induce polymerization of nonaggregated microtubule subunits, resulting in faster growth.     

Inhibitory action of red light on the growth of the maize mesocotyl: evaluation of the auxin hypothesis

Brief irradiation of 3-d-old maize (Zea mays L.) seedlings with red light (R; 180 J m^-2) inhibits elongation of the mesocotyl (70–80% inhibition in 8 h) and reduces its indole-3-acetic acid (IAA) content. The reduction in IAA content, apparent within a few hours, is the result of a reduction in the supply of IAA from the coleoptile unit (which includes the shoot apex and primary leaves). The fluence-response relationship for the inhibition of mesocotyl growth by R and far-red light closely resemble those for the reduction of the IAA supply from the coleoptile. The relationship between the concentration of IAA (1–10 M) supplied to the cut surface of the mesocotyl of seedlings with their coleoptile removed and the growth increment of the mesocotyl, measured after 4 h, is linear. The hypothesis that R inhibits mesocotyl growth mainly by reducing the IAA supply from the coleoptile is supported. However, mesocotyl growth in seedlings from which the coleoptiles have been removed is also inhibited by R (about 25% inhibition in 8 h). This inhibition is not related to changes in the IAA level, and not relieved by applied IAA. In intact seedlings, this effect may also participate in the inhibition of mesocotyl growth by R. Inhibition of cell division by R, whose mechanism is not known, will also result in reduced mesocotyl elongation especially in the long term (e.g. 24 h).Abbreviations FR far-red light - IAA indole-3-acetic acid - P_fr phytochrome in the far-red-absorbing form - P_r phytochrome in the red-absorbing form - R red light     

Effects of red and far-red light on cell division in the shoot apex ofSilene coeli-rosa L.

Summary 28-day-old plant ofSilene coeli-rosa were exposed at 1,700 hours to 5 or 10 minutes red light, 5 or 10 minutes far-red light, red followed by far-red, far-red followed by red or maintained in darkness. Measurements of the proportions of cells with the 2 C and 4 C amounts of DNA in the shoot apex of the plants, sampled at 2,000 hours, showed that far-red light promoted an increase in the G2 proportion whereas red light resulted in an increase in the G1 proportion of the cell cycle, relative to the dark controls. Moreover these changes were red, far-red reversible. All light treatments resulted in increases in the mitotic index in the apex compared with the dark controls, suggesting increases in the growth rate. The data implicate phytochrome in a low energy response and suggest that, in the shoot apex, G1 is shortened markedly following exposure to farred light, whilst G2 is shortened the most following exposure to red light. The results are discussed in relation to flower-initiation.     

Phytochrome control of electrical potentials and intercellular coupling in oat-coleoptile tissue

Summary Microelectrodes were used to demonstrate two electrical responses which occur in oat (Avena sativa L.) coleoptile parenchyma-cells during exposure to red light. The membrane potential of these cells depolarized 5–10 mV in several seconds in red light and repolarized more slowly in far-red light. By pulsing current through the cells, it was found that cellular coupling along the longitudinal axis of the coleoptile increased about 2-fold in red light, but that coupling along the lateral axis was not affected. The rapid changes in membrane potential are consistent with the idea of a membrane locale for early phytochrome action. The coupling experiments suggest that phytochrome may also affect plasmodesmata in this system.     

Phytochrome Action in Oryza sativa L: IV. Red and Far Red Reversible Effect on the Production of Ethylene in Excised Coleoptiles

Excised apical segments of etiolated rice (Oryza sativa L.) coleoptiles produced ethylene. Increasing the number of cut sites per coleoptile increased the rate of ethylene formation. Ethylene produced by an etiolated-intact seedling in the dark was about a half of that by the excised coleoptile segment. Red light of low energy as well as of continuous irradiation inhibited the production of ethylene. The inhibition by a low energy dose of red light was partly relieved, if the red light was followed immediately by a small dose of far red light. The effect of red and far red light was repeatedly reversible, indicating that ethylene production was regulated by a phytochrome system. If the exposure to far red light was preceded by a period of darkness, this photoreversibility disappeared; 50% of the initial reversibility was lost within 5 hours. Applied ethylene (10 microliters per liter) significantly promoted the growth of intact coleoptiles of either totally etiolated or red light-treated seedlings, but had no effect on the excised apical segment of coleoptile.     

Effect of red light on coleoptile growth

The effects of red light in reducing the growth of the oat (Avena sativa L.) coleoptile and the synthesis of auxin in the coleoptile tip are detectable 2 hours after treatment and become more pronounced with time. When the coleoptile tip is supplied with additional tryptophan the synthesis of auxin is doubled both in darkness and when exposed to red light. Treatment of the tip with gibberellic acid or pyridoxal phosphate overcomes the reduction of auxin synthesis caused by red light. The uptake of exogenous indoleacetic acid, at pH 6.5, by coleoptile tissue is doubled by exposure to red light. The effect of red light on coleoptile growth appears to be mediated by phytochrome in the cell membrane which delocalizes the tryptophan utilized for auxin synthesis.     

Phytochrome Modifies Blue-light-induced Electrical Changes in Corn Coleoptiles

Unilateral blue light administered to corn coleoptile segments produces no alteration of transmembrane potential on the light side, and only a small and slow hyperpolarization on the dark side. Red light causes a 5-15 millivolt depolarization in cells on the light side causes and somewhat smaller effects on the dark side. Blue given after red causes a rapid hyperpolarization on both sides of the coleoptile. The effect of the potentiating red preirradiation is probably due to phytochrome, being largely abolished by far-red given after red, but before the blue light. The effect of prior red irradiation decays in the dark, showing a half-time of about 45 minutes at room temperature. This rapid cooperativity between phytochrome and the phototropic pigment may indicate a common locale, possibly in a membrane.