Phytochrome modulation of blue-light-induced phototropism

Red light enhances hypocotyl phototropism toward unilateral blue light through a phytochrome‐mediated response. This study demonstrates how the phytochromes modulate blue‐light‐induced phototropism in the absence of a red light pre‐treatment. It was found that phytochromes A, B, and D have conditionally overlapping functions in the promotion of blue‐light‐induced phototropism. Under very low blue light intensities (0.01 µmol m^?2 s^?1) phyA activity is necessary for the progression of a normal phototropic response, whereas above 1.0 µmol m^?1 s^?2 phyB and phyD have functional redundancy with phyA to promote phototropism. PhyA also contributes to attenuation of phototropism under high fluence rates of unilateral blue light, which was previously shown to be dependent on the phototropins and cryptochromes. From these results, it appears that phytochromes are required to develop a robust phototropic response under low fluence rates, whereas under high irradiances where phototropism may be less important, phyA suppresses phototropism.     

A Unified Model of Shoot Tropism in Plants: Photo-, Gravi- and Propio-ception

Land plants rely mainly on gravitropism and phototropism to control their posture and spatial orientation. In natural conditions, these two major tropisms act concurrently to create a photogravitropic equilibrium in the responsive organ. Recently, a parsimonious model was developed that accurately predicted the complete gravitropic and proprioceptive control over the movement of different organs in different species in response to gravitational stimuli. Here we show that the framework of this unifying graviproprioceptive model can be readily extended to include phototropism. The interaction between gravitropism and phototropism results in an alignment of the apical part of the organ toward a photogravitropic set-point angle. This angle is determined by a combination of the two directional stimuli, gravity and light, weighted by the ratio between the gravi- and photo-sensitivities of the plant organ. In the model, two dimensionless numbers, the graviproprioceptive number B and the photograviceptive number M, control the dynamics and the shapes of the movement. The extended model agrees well with two sets of detailed quantitative data on photogravitropic equilibrium in oat coleoptiles. It is demonstrated that the influence of light intensity I can be included in the model in a power-law-dependent relationship M(I). The numbers B and M and the related photograviceptive number D are all quantitative genetic traits that can be measured in a straightforward manner, opening the way to the phenotyping of molecular and mechanical aspects of shoot tropism.     

Second positive- and negative phototropism in Vaucheria geminata

Second positive and negative phototropism are found in Vaucheriageminata when the light intensity is very high. The dose-responsecurve of this alga resembles those obtained with Avena coleoptiles,suggesting that the phototropism of both plants occurs througha common initial process. ¹ Present address: Institute for Agricultural Research, TohokuUniversity, Sendai 980, Japan. (Received October 13, 1976; )     

植物下胚轴向光弯曲机制研究进展

植物向光弯曲生长主要是由于其向光和背光面生长素的不对称分布引起。近年来研究发现,在不同强度的蓝光单侧照射下,植物可能存在不同的向光弯曲调节机制。目前,关于向光素PHOT1介导弱蓝光引起的下胚轴弯曲研究较为详细,即PHOT1感受蓝光后,与其下游的信号蛋白NPH3、RPT2和PKS1相互作用,调控生长素运输蛋白的活性及定位,诱导生长素的不对称分布引起向光弯曲。PHOT1和PHOT2以功能冗余方式调节强蓝光引起的植物下胚轴向光弯曲,NPH3可能作为共享调节因子,引发不同的信号转导通路实现功能互补。此外,其他光受体、激素、蛋白激酶、蛋白磷酸酶以及Ca2＋也参与了植物向光弯曲的调节。本文就近年来有关植物下胚轴向光弯曲信号组分及可能的网络关系进行总结,并对该研究领域存在的问题及今后可能的研究方向进行展望。     

Effect of constant light on the circadian rhythm in locomotor activity in intact or eyelids-removed rats

According to the Aschoff's role, exposure to continuous light (LL) results in the elongation of the free-running period of the rat circadian rhythm. However, the LL may not always mean the constant intensity of the light for the suprachiasmatic nucleus, since the rat may regulate the contrast of the illumination by their eyelids which are closed during the sleep phase. In this study, the surgical removal of the eyelids under the LL caused arrhythmicity of the locomotor activity in 7 of 10 rats. The remaining 3 rats maintained the free-running rhythm after the removal of the eyelids. These results suggest that constant light may affect the free-running rhythm of the rat with or without eyelids in the different manner.     

THE MECHANISMS OF TROPISTIC REACTIONS AND THE STRYCHNINE EFFECT IN DAPHNIA

1. Experiments with strychnine were performed to test two assumptions important in the development of a theory for the mechanisms involved in the tropisms exhibited by Daphnia. 2. After a brief interval in strychnine solution Daphnia exhibits a reversal of the primary sign (a) of phototropism, from negative to positive; and (b) of galvanotropism, from anodic to cathodic. In both cases the orientation of the body remains the same. 3. The mechanism responsible for the sign of phototropism and galvanotropism in Daphnia is therefore distinct from that underlying orientation. 4. Evidence is obtained indicating that changes in sign of tropism, produced by changes in illumination or by subjection to strychnine, involve the control of antagonistic muscles in the swimming appendages which are reciprocally innervated.     

STEREOTROPISM IN RATS AND MICE

Typical stereotropic orientation toward a lateral surface of contact is obtained in young rats and mice, and with adult mice congenitally blind. Removal of vibrissæ or tail or both does not essentially affect this response. Equal contact on both sides of the body prevents orientation toward either source of contact. Unequal contact areas on the two sides leads to orientation toward the more extensive contact. This behavior very exactly parallels the stereotropic conduct of arthropods, and thus provides a fairly complete instance of a tropism in mammals.     

Hypocotyl growth orientation in blue light is determined by phytochrome A inhibition of gravitropism and phototropin promotion of phototropism

How developing seedlings integrate gravitropic and phototropic stimuli to determine their direction of growth is poorly understood. In this study we tested whether blue light influences hypocotyl gravitropism in Arabidopsis. Phototropin1 (phot1) triggers phototropism under low fluence rates of blue light but, at least in the dark, has no effect on gravitropism. By analyzing the growth orientation of phototropism-deficient seedlings in response to gravitropic and phototropic stimulations we show that blue light not only triggers phototropism but also represses hypocotyl gravitropism. At low fluence rates of blue light phot1 mutants were agravitropic. In contrast, phyAphot1 double mutants grew exclusively according to gravity demonstrating that phytochrome A (phyA) is necessary to inhibit gravitropism. Analyses of phot1cry1cry2 triple mutants indicate that cryptochromes play a minor role in this response. Thus the optimal growth orientation of hypocotyls is determined by the action of phyA-suppressing gravitropism and the phototropin-triggering phototropism. It has long been known that phytochromes promote phototropism but the mechanism involved is still unknown. Our data show that by inhibiting gravitropism phyA acts as a positive regulator of phototropism.     

ON THE EQUILIBRATION OF GEOTROPIC AND PHOTOTROPIC EXCITATIONS IN THE RAT

The intensity of light required to just counterbalance geotropic orientation of young rats, with eyelids unopened, is so related to the angle of inclination (α) of the creeping plane that the ratio log I/log sin α is constant. This relationship, and the statistical variability of I as measured at each value of α, may be deduced from the known phototropic and the geotropic conduct as studied separately, and affords proof that in the compounding of the two kinds of excitation the rat is behaving as a machine.     

Growth Distribution during Phototropism of Arabidopsis thaliana Seedlings

The elongation rates of two opposite sides of hypocotyls of Arabidopsis thaliana seedlings were measured during phototropism by using an infrared imaging system. In first positive phototropism, second positive phototropism, and red light-enhanced first positive phototropism, curvature toward the light source was the result of an increase in the rate of elongation of the shaded side and a decrease in the rate of elongation of the lighted side of the seedlings. The phase of straightening that followed maximum curvature resulted from a decrease in the elongation rate of the shaded side and an increase in the elongation rate of the lighted side. These data for the three types of blue light-induced phototropism tested in this study and for the phase of straightening are all clearly consistent with the growth rate changes predicted by the Cholodny-Went theory.     

Stem phototropism toward blue and ultraviolet light

Positive phototropism is the process through which plants orient their organs toward a directional light source. While the blue light receptors phototropins (phot) play a major role in phototropism toward blue (B) and ultraviolet (UV) radiation, recent research showed that the UVB light receptor UVR8 also triggers phototropism toward UVB. In addition, new details of the molecular mechanisms underlying the activity of these receptors and interaction with other environmental signals have emerged in the past years. In this review, we summarize the current knowledge about hypocotyledoneous and inflorescence stem growth reorientation toward B and UVB, with a focus on the molecular mechanisms.     

Modeling bidirectional transport of quantum dot nanoparticles in membrane nanotubes

This paper develops a model of transport of quantum dot (QD) nanoparticles in membrane nanotubes (MNTs). It is assumed that QDs are transported inside intracellular organelles (called here nanoparticle-loaded vesicles, NLVs) that are propelled by either kinesin or dynein molecular motors while moving on microtubules (MTs). A vesicle may have both types of motors attached to it, but the motors are assumed to work in a cooperative fashion, meaning that at a given time the vesicle is moved by either kinesin or dynein motors. The motors are assumed not to work against each other, when one type of motors is pulling the vesicle, the other type is inactive. From time to time the motors may switch their roles: passive motors can become active motors and vice versa, resulting in the change of the vesicle’s direction of motion. It is further assumed that QDs can escape NLVs and become free QDs, which are then transported by diffusion. Free QDs can be internalized by NLVs. The effects of two possible types of MT orientation in MNTs are investigated: when all MTs have a uniform polarity orientation, with their plus-ends directed toward one of the cells connected by an MNT, and when MTs have a mixed polarity orientation, with half of MTs having their plus-ends directed toward one of the cells and the other half having their plus-ends directed toward the other cell. Computational results are presented for three cases. The first case is when organelles are as likely to be transported by kinesin motors as by dynein motors. The second case is when organelles are more likely to be transported by kinesin motors than by dynein motors, and the third case is when NLVs do not associate with dynein motors at all.     

Differential roles of auxin efflux carrier PIN proteins in hypocotyl phototropism of etiolated Arabidopsis seedlings depend on the direction of light stimulus

In a recent study, we demonstrated that although the auxin efflux carrier PIN-FORMED (PIN) proteins, such as PIN3 and PIN7, are required for the pulse-induced first positive phototropism in etiolated Arabidopsis hypocotyls, they are not necessary for the continuous-light-induced second positive phototropism when the seedlings are grown on the surface of agar medium, which causes the hypocotyls to separate from the agar surface. Previous reports have shown that hypocotyl phototropism is slightly impaired in pin3 single mutants when they are grown along the surface of agar medium, where the hypocotyls always contact the agar, producing some friction. To clarify the possible involvement of PIN3 and PIN7 in continuous-light-induced phototropism, we investigated hypocotyl phototropism in the pin3 pin7 double mutant grown along the surface of agar medium. Intriguingly, the phototropic curvature was slightly impaired in the double mutant when the phototropic stimulus was presented on the adaxial side of the hook, but was not impaired when the phototropic stimulus was presented on the abaxial side of the hook. These results indicate that PIN proteins are required for continuous-light-induced second positive phototropism, depending on the direction of the light stimulus, when the seedlings are in contact with agar medium.     

Target-directed Orientation in Displaced Honeybees

Under sunny weather conditions, displaced honeybees (Apis mellifera) usually fly into the celestial compass direction and thus may be misled from their goal, or they are disorientated. Under cloudy conditions, they may determine the celestial compass direction from prominent landmarks. They may also fly directly toward their goal from a release site. In two experiments, we investigated the orientation of displaced bees when a landmark (target) was close to the goal under different weather conditions. It is shown that in sunny conditions, the celestial compass will override target orientation under most conditions. Under 100% cloud cover, the celestial compass direction retrieved from landmarks modulates target-orientated behaviour but is not by itself a primary orientation factor. The bees will fly toward a previously encountered landmark that signals the target, and in case of several similar landmarks which are visible to the bees, they will choose the one in the direction nearest the celestial compass direction. The results indicate that honeybee orientation is the result of a set of context-specific interdependent orientation mechanisms.     

The Effect of the Apical Bud on the Growth of Lateral Buds on Subterranean Shoots

The influence of the apical bud on the growth of the lateral buds on subterranean shoots was studied in Stachys sieboldiiMig. and Helianthus rigidus(Gass.) Desv. Removing and damaging the apical parts of subterranean shoots or their treatment with 2% chlorocholine chloride shoot enhanced shoot branching. The response to light of the apical bud was invariably negative: the stolons, which came out or were extracted from the soil, grew back into the ground (negative phototropism). The response to light of lateral buds was autonomous and depended on the conditions of their initiation. The lateral buds developed in darkness manifested negative phototropism when withdrawn from the soil and exposed to the light, whereas the buds developed in the light showed positive phototropism. The author concludes that the concept of apical dominance, thoroughly studied in aboveground shoots, is also valid for subterranean shoots. However, in contrast to the former, in the latter case, the apical bud does not control the growth orientation of the lateral buds.     

Specific inhibition of phototropism in corn seedlings

Geotropism was used as a control for the specificity of potential inhibitors of phototropism by the coleoptiles of corn (Zea mays) seedlings. The compounds tested fall into three categories showing: (a) no inhibition of either phototropism or geotropism (KCl); (b) nonspecific inhibition of both phototropism and geotropism (KCN); and (c) specific inhibition of phototropism (KI, NaN₃, and phenylacetic acid). Simultaneous irradiation of coleoptiles with phototropically inert light in addition to the phototropically active blue light also results in an inhibition of phototropism. Since azide, iodide, and phenylacetic acid are known to interact with flavins while a simultaneous irradiation with a phototropically inert light may depopulate the first triplet state of flavins, these data support the hypothesis that the photoreceptor pigment for phototropism in corn is a flavin.     

Polarity induction versus phototropism in maize: Auxin cannot replace blue light

In a previous study (Nick and Schäfer 1991, Planta 185, 415–424), unilateral blue light had been shown, in maize coleoptiles, to induce phototropism and a stable transverse polarity, which became detectable as stable curvature if counteracting gravitropic stimulation was removed by rotation on a horizontal clinostat. This response was accompanied by a reorientation of cortical microtubules in the outer epidermis (Nick et al. 1990, Planta 181, 162–168). In the present study, this stable transverse polarity is shown to be correlated with stability of microtubule orientation against blue light and changes of auxin content. The role of auxin in this stabilisation was assessed. Although auxin can induce reorientation of microtubules it fails to induce the stabilisation of microtubule orientation induced by blue light. This was even true for gradients of auxin able to induce a bending response similar to that ellicited by phototropic stimulation. Experiments involving partial irradiation demonstrated different perception sites for phototropism and polarity induction. Phototropism starts from the very coleoptile tip and involves transmission of a signal (auxin) towards the subapical elongation zone. In contrast, polarity induction requires local action of blue light in the elongation zone itself. This blue-light response is independent of auxin.This work was supported by the Deutsche Forschungsgemeinschaft and two grants of the Studienstiftung des Deutschen Volkes and the Human Frontier Science Program Organization to P.N.     

Heliotropic leaf movements in common beans controlled by air temperature

Heliotropic leaf movements were examined in common beans (Phaseolus vulgaris cv Blue Lake Bush) under outdoor and laboratory conditions. Heliotropic leaf movements in well-watered plants were partly controlled by temperature, and appeared to be independent of atmospheric humidity and CO₂ concentration. When environmental conditions were held constant in the laboratory, increased air temperature caused bean leaves to orient more obliquely to a light source. Ambient CO₂, intercellular CO₂, and net photosynthesis were not correlated with the temperature-induced changes in heliotropic movements, nor did they significantly affect these movements directly. The effect of air temperature on leaf movements need not be mediated through a change in leaf water potential, transpiration, or leaf conductance. Air temperature modified laminar orientation in light through its effect on tissue temperature in the pulvinal region, not that of the lamina or petiole. However, under darkness the temperature effects on leaf movements were not expressed. Active heliotropic movements in response to air temperature allowed lamina temperature to remain close to the thermal optimum of photosynthesis. This temperature effect underlies a commonly observed pattern of leaf movements under well-watered conditions: a tendency for leaves to face the sun more obliquely on hot days than cool days.     

ON STEREOTROPISM IN TENEBRIO LARV?

Larvæ of Tenebrio while creeping show homostrophic responses, and stereotropic orientation to lateral contacts. Homostrophic orientation is inhibited by stereotropism. Both depend upon the anterior portion of the central nervous system. Stereotropic orientation due to unilateral contact, particularly at the anterior end, persists briefly after the cessation of the contact. Equal posterior bilateral contact of the body obliterates stereotropic bending. Unequal posterior bilateral contacts lead to orientation through an angle roughly proportional to the differences in contact areas. Functional symmetry in such responses is not disturbed by asymmetrical distribution of the body "hairs." The stereotropic orientation undergoes reversal of direction, central in origin, when the stimulation is sufficiently intense. Stereotropic response, leading to maintained lateral contact with a surface or to bending when the end of such a surface has been passed, is inhibited by a definite intensity of light. These findings (1) round out the demonstration that stereotropism is truly of a tropistic character, and (2) make possible the understanding of conduct in a case involving the participation of contact stimulation, phototropism, temperature, and homostrophy.