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; )     

THE EFFECT OF LIGHT ON THE CELLULAR COMPONENTS OF POLARIZED GROWTH IN BEAN INTERNODES

First internodes of light-grown bean seedlings exposed to supplementary red and far-red light and those of dark-grown seedlings were sectioned and studied to determine the effects of irradiation on the cellular components of polarized growth. Cell counts and measurements of epidermis, cortex, and pith are given. Increased length of internodes of far-red-treated plants was caused by both increased rate and increased duration of cell elongation. The effect of far-red light is interpreted as a reversal of the accelerating effect of light upon cell maturation. It is suggested that investigations of the mechanism of the red, far-red response of stems be concerned with the processes involved in cell elongation. In darkness, rate and duration of cell division as well as rate and duration of cell elongation were greater than in any of the irradiated plants, indicating that only part of the photocontrol of stem elongation is mediated through the red, far-red system.     

Light Quality Regulates Lateral Root Development in Tobacco Seedlings by Shifting Auxin Distributions

Light is an important environmental regulator of diverse growth and developmental processes in plants. However, the mechanisms by which light quality regulates root growth are poorly understood. We analyzed lateral root (LR) growth of tobacco seedlings in response to three kinds of light qualities (red, white, and blue). Primary (1°) LR number and secondary (2°) LR density were elevated under red light (on days 9 and 12 of treatment) in comparison with white and blue lights. Higher IAA concentrations measured in roots and lower in leaves of plants treated with red light suggest that red light accelerated auxin transport from the leaves to roots (in comparison with other light qualities). Corroborative evidence for this suggestion was provided by elevated DR5::GUS expression levels at the shoot/root junction and in the 2° LR region. Applications of N-1-naphthylphthalamic acid (NPA) to red light-treated seedlings reduced both 1° LR number and 2° LR density to levels similar to those measured under white light; DR5::GUS expression levels were also similar between these light qualities after NPA application. Results were similar following exogenous auxin (NAA) application to blue light-treated seedlings. Direct [³H]IAA transport measurement indicated that the polar auxin transport from shoot to root was increased by red light. Red light promoted PIN3 expression levels and blue light reduced PIN1, 3–4 expression levels in the shoot/root junction and in the root, indicating that these genes play key roles in auxin transport regulation by red and blue lights. Overall, our findings suggest that three kinds of light qualities regulate LR formation in tobacco seedlings through modification of auxin polar transport.     

INFLUENCE OF THE TONIC EFFECT OF GRAVITATION AND AUXIN ON CELL ELONGATION AND POLARITY IN ROOTS

The influence of a longitudinal (tonic) gravitational force and of auxin on the pattern of growth and cell polarity has been studied on intact roots of wheat seedlings. A klinostat technique was used for controlling gravitation. Growth in length was evaluated as cell division activity, rate of cell elongation (μ/h) and duration of elongation (h). Exogenous auxin (1-NAA) increases the rate of cell elongation in all concentrations tested (10⁻⁸ — 3 × 10⁻⁷m ) and shortens the time of elongation with increasing concentration. It promotes rate of cell elongation in roots as it does in shoots. It also accentuates the polar insertion of root hairs and their growth. The tonic effect of gravitation resembles that of an increase in auxin both in light and darkness. The results are discussed in relation to plagiotropic growth of roots, root growth promotions by auxin, and the difference between root and shoot growth.     

EFFECTS OF RED AND FAR-RED ILLUMINATION ON THE ELONGATION OF FERN PROTONEMATA AND RHIZOIDS

The elongation of fern protonemata is controlled by red andfar-red light in an atypical fashion. Red light promotes theelongation of young plants but inhibits the elongation of olderplants. Far-red light promotes elongation regardless of filamentage, and the maximum promotion by far-red is greater than thepromotion which red light causes in young filaments. The elongationof rhizoids is under typical red, far-red control. Red lightpromotes elongation, and a period of far-red illumination followingred light treatment negates the promotive effects of red light. ¹ Present address of the authors: Dept. of Bacteriology andBotany, Syracuse University, Syracuse, New York, U. S. A. (Received November 5, 1962; )     

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.     

STIMULATIVE EFFECT OF HALOGENATED TYROSINE ON ROOT GROWTH OF RICE SEEDLING

The effect of iodotyrosine and its related compounds on theroot elongation was studied using intact roots of rice seedlingsgrown on an agar medium and in a culture solution. MIT (3-monoiodotyrosine), DIT (3,5-diiodotyrosine) markedlystimulated the growth of seminal root, but tyrosine itself wasnot active. Both iodinated amino acids doubled the length ofseminal root in comparison with the control when used at theconcentration of 10^–5M, where the growth of shoot andcrown root was not affected. And above 10^–5M, the developmentof lateral root and root hairs was inhibited. DBT (3,5-dibromotyrosine) also notably increased the lengthof seminal root. Potassium iodide did not accelerate the growthof root. (Received September 18, 1964; )     

Light-mediated bioregulation of tillering and photosynthate partitioning in wheat

The influence of plant population density on spectral distribution of light received by wheat ( Triticum aestivum L. cv. Coker 797) seedlings was measured under field conditions, and effects of red and far-red light on tillering and photosynthate partitioning were studied in controlled environments. Spectral distribution of light was measured in sunflecks at soil level in close-, intermediate-, and wide-spaced field populations during the tillering stage. Close-spaced seedlings received higher far-red/red light ratios than wide-spaced plants because of the larger amount of far-red reflected from green leaves of the more numerous nearby plants. The far-red/red light ratios in all population densities were higher in late afternoon than at noon. Close-spaced plants developed fewer tillers, less roots and longer leaves than wide-spaced seedlings under field conditions. In controlled environments, a higher far-red/red ratio during photosynthetic periods resulted in fewer tillers and longer leaves; whereas, brief red or far-red exposures at the end of each day had a more pronounced effect. Wheat seedlings that received 5-min exposures to far-red light at the end of the photosynthetic period each day for 20 consecutive days developed fewer tillers, longer leaves, less roots, and a higher shoot/root biomass ratio. The effects of far-red light were reversed by red light. The light spectral shifts associated with field plant population densities and the responses to red and far-red treatments under controlled environments suggest that phytochrome serves as a sensing mechanism that detects the amount of competition from other plants, and regulates the development of tillers and the partitioning of photosynthate between shoots and roots.     

Growth of a mangrove (Rhizophora apiculata) seedlings as influenced by GA3, light and salinity

The growth performance of Rhizophora apiculata Blume (mangrove) seedlings in the presence and absence of exogenous gibberellic acid (GA3) under different combinations of salinity and light was analyzed. Root and shoot growth responses of 75-day old seedlings in liquid-culture, were measured. It was concluded that light exhibited a significant inhibitory effect on all the growth parameters-number of primary roots, primary root length, shoot elongation, number of leaves, total leaf area; and, the GA3 treatment singly or in combinations with light, showed a significant influence on the total leaf area and primary root length.     

Non-photosynthetic effects of red and far-red light on root-nodule formation by leguminous plants

Summary Nodulation of pea and broad bean plants grown in the light was found to be reduced when the roots were exposed to far-red light for 5–15 minutes daily during 5 consecutive days following inoculation with nodule bacteria. Similar results were obtained following a single exposure to far-red light during a period of 15 minutes at the 3rd or 4th day after inoculation. When the roots were exposed to far-red light either before inoculation or during the first two days afterwards there were either no effects or only slight effects on nodulation The inhibitory effect of far-red light on nodulation was partly reduced by subsequent exposure to red light, provided that the same part of the plant was exposed to both red and far-red light,viz either the root or the shoot. When different parts of the plant were exposed to red and far-red light respectively, there was no interaction between the two kinds of light on nodulation. Plants whose roots were exposed to far-red light did not subsequently show stem elongation.Nodules were found to develop on the roots of pea plants grown in the dark, provided that the plants were kept at or below 22°C. At 25°C nodulation was almost absent. Nodulation was decreased by addition of kinetin and IAA. In contrast to plants grown in the light pea plants grown in the dark, inoculated with either an effective or ineffective strain of Rhizobium, developed equal numbers of nodules. Exposure to red light slightly increased the percentage of nodulated plants but decreased the number of nodules per plant. Exposure to far-red light slightly decreased both the percentage of nodulated plants and the number of nodules per plant. The effect of far-red light was counteracted by red light andvice versa.     

A synergistic effect of 2-ethyl-l-isopropoxycarbonyl-3-(4-tolylcarbamoyl) isourea with gibberellic acid on growth of rice seedlings

2-Ethyl-1-isopropoxycarbonyl-3-(4-tolylcarbamoyl)isourea [EITIU]stimulated both shoot and root growth of rice seedlings. Gibberellicacid [GA]-induced shoot elongation was further stimulated byEITIU, and the combined application of both compounds was shownto be distinctly synergistic. A similar synergistic action wasobserved in the growth of rice mesocotyls in the dark. The inhibitionof root growth caused by GA was overcome by combination withEITIU. The growth-stimulating activity of EITIU was not observedin lettuce hypocotyls. ¹ This paper is Part I in the series "Plant growth-regulatingactivities of isourea derivatives and related compounds." (Received February 7, 1976; )     

Root growth dynamics of Aleppo pine (Pinus halepensis Mill.) seedlings in relation to shoot elongation, plant size and tissue nitrogen concentration

Large and high nitrogen (N) concentration seedlings frequently have higher survival and growth in Mediterranean forest plantations than seedlings with the opposite traits, which has been linked to the production of deeper and larger root systems in the former type of seedlings. This study assessed the influence of seedling size and N concentration on root growth dynamics and its relation to shoot elongation in Aleppo pine (Pinus halepensis Mill.) seedlings. We cultivated seedlings that differed in size and tissue N concentration that were subsequently transplanted into transparent methacrylate tubes in the field. The number of roots, root depth, and the root and shoot elongation rate (length increase per unit time) were periodically measured for 10 weeks. At the end of the study, we also measured the twig water potential (ψ) and the mass of plant organs. New root mass at the end of the study increased with seedling size, which was linked to the production of a greater number of new roots of lower specific length rather than to higher elongation rate of individual roots. Neither plant size nor N concentration affected root depth. New root mass per leaf mass unit, shoot elongation rate, and pre-dawn ψ were reduced with reduction in seedling size, while mid-day ψ and the root relative growth rate were not affected by seedling size. N concentration had an additive effect on plant size on root growth but its overall effect was less important than seedling size. Shoot and roots had an antagonistic elongation pattern through time in small seedlings, indicating that the growth of both organs depressed each other and that they competed for the same resources. Antagonism between shoot and root elongation decreased with plant size, disappearing in large and medium seedlings, and it was independent of seedling N concentration. We conclude that root and shoot growth but not rooting depth increased with plant size and tissue N concentration in Aleppo pine seedlings. Since production of new roots is critical for the establishment of planted seedlings, higher absolute root growth in large seedlings may increase their transplanting performance relative to small seedlings. The lack of antagonism between root and shoot growth in large seedlings suggests that these plants can provide resources to sustain simultaneous growth of both organs.     

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.     

Photoreversible and photoirreversible absorbance changes in the red and far-red spectral regions in Zea primary roots

When 2-mm apical segments of the primary roots of Zea mays L.(cv. Golden Cross Bantam 70) were irradiated successively withred and far-red light, a photoirreversible absorbance decreasewas separated from the red far-red reversible absorbance changetypical of phytochrome. The difference spectrum of the reversiblechange showed maximum absorbance changes at 666 and 730 nm,while the photoirreversible change induced by red light showeda maximum decrease at 640 nm. The photoreversible absorbancechange was linearly proportional to the fluence of red lightbetween 1 and 6 J m^–2, while the photoirreversible absorbancechange was proportional to its logarithm. Red light of approximately6 J m^–2 induced 50% of the maximum photoirreversible absorbancechange at 640 nm but only about 25% of the maximum photoreversibleabsorbance change. Moreover, no effect of ascorbate on the twoabsorbance changes was observed. ¹Faculty of Education, University of Yamagata, Yamagata 990,Japan. (Received November 2, 1980; )     

Photoreversible and photoirreversible absorbance changes in the red and far-red spectral regions in Zea primary roots

When 2-mm apical segments of the primary roots of Zea mays L.(cv. Golden Cross Bantam 70) were irradiated successively withred and far-red light, a photoirreversible absorbance decreasewas separated from the red far-red reversible absorbance changetypical of phytochrome. The difference spectrum of the reversiblechange showed maximum absorbance changes at 666 and 730 nm,while the photoirreversible change induced by red light showeda maximum decrease at 640 nm. The photoreversible absorbancechange was linearly proportional to the fluence of red lightbetween 1 and 6 J m^–2, while the photoirreversible absorbancechange was proportional to its logarithm. Red light of approximately6 J m^–2 induced 50% of the maximum photoirreversible absorbancechange at 640 nm but only about 25% of the maximum photoreversibleabsorbance change. Moreover, no effect of ascorbate on the twoabsorbance changes was observed. ¹Faculty of Education, University of Yamagata, Yamagata 990,Japan. (Received November 2, 1980; )     

Effects of Colchicine and Light on Cell Form in Fern Gametophytes. Implications for a Mechanism of Light-induced Cell Elongation

Spores of the fern, Onoclea sensihilis L., suffer a disruption of normal development when they are cultured on media containing colchicine. Cell division is inhibited, and the spores develop into giant spherical cells under continuous white fluorescent light. In darkness only slight cell expansion occurs. Spherical cell expansion in the light requires continuous irradiation. Photosynthesis does not seem to be involved, since variations in light intensity do not affect the final cell diameter; the addition of sucrose to the medium does not permit cell expansion in darkness; and the inhibitor DCMU does not block the light-induced cell expansion. Continuous irradiation of colchicine-treated spores with blue, red or far-red light produces different patterns of cell expansion. Blue light permits spherical growth, similar to that found under white light, whereas red and far-red light promote the reestablishment of polarized filamentous growth. Although ethylene is unable to induce polarized cell expansion in colchicine-treated spores in darkness or white and blue light, it enhances filamentous growth which already is established by red or far-red irradiation. Both red and far-red light increase the elongation of normal filaments (untreated with colchicine) above that of dark-grown plants, but under all 3 conditions the rates of volume growth are identical. Light, however, does cause a decrease in the cell diameters of irradiated filaments. These data are used to construct an hypothesis to explain the promotion of cell elongation in fern protonemata by red and far-red light. The model proposes light-mediated changes in microtubular orientation and cell wall structure which lead to restriction of lateral cell expansion and enhanced elongation growth.     

Phytochromabhängiger Ionentransport in Erbsensämlingen

Summary Five to 6 day old dark-adapted dwarf and tall pea seedlings grown in water culture were illuminated for ten minutes with red light and/or ten minutes with far-red light, and 90 to 170 minutes later their roots were immersed in a 0.2 mM K⁺ solution containing labeled ⁸⁶Rb⁺. After two hours uptake the fresh-weights and radioactivities of the shoot organs were determined. It was found that red light inhibits K⁺uptake into internodes and promotes uptake into the plumula. The red-light effect on K⁺transport precedes the red-light induced growth inhibition of internodes and growth promotion of leaves and is abolished by far-red light given immediately after red. The red-light effect on K⁺transport is independent of the concentration of K⁺ given to the roots in the range between 0.2 to 125 mM.     

Primary and Lateral Root Development of Dark- and Light-Grown Cotton Seedlings under Salinity Stress*

Primary root growth dynamics and lateral root development of dark- and light grown cotton seedlings (Gossypium hirsutum L., cv. Acala SJ-2) were studied under control and salinity stress conditions. The seedlings were grown by two methods: A) in paper-lined, vermiculite-filled beakers with the plants growing between the paper and the glass wall (Gladish and Rost, 1993), and B) in hydroponics after germination and initial growth in germination paper rolls saturated with the treatment solutions (Kent and Läuchli, 1985). After germination, daily primary root elongation rate gradually incrased to a maximum, then gradually declined to close to zero for dark-grown seedlings, or to sustained rates of about 10 mm per day for light-grown control plants. Salinity stress delayed primary root growth and reduced peak elongation rates, without changing the general primary root growth pattern. These results suggest that salinity changed the time-scale, but did not modify the normal developmental sequence. Lateral root growth was more inhibited by salinity than primary root growth. In addition, elongation of lateral roots was more inhibited by salinity than their initiation and emergence. Light exposure of the shoot favored both sustained primary root growth from 7 days after planting, and lateral root emergence and growth. Salinity effects were more severe on seedlings germinated and grown in hydroponics (method B) than on vermiculite-grown plants (method A). These results emphasize the importance of growing conditions for the NaCl-induced effects on cotton root development. In addition, the differential effects of salinity on primary and lateral roots became evident, pointing to diverse control mechanisms for the development of these root types.