Site of light perception and motor cells in a sun-tracking lupine (Lupinus succulentus)

From the results of shading experiments and irradiation of leaves from different directions, the pulvinus has been identified as the site of light perception in leaves of the sun-tracker Lupinus succulentus Dougl. Within the pulvinus, the light-sensitive region is located on the adaxial surface of the base of each leaflet. This is coincident with the site of the motor cells which are located in the peripheral four cell layers. Motor cells were visualized by microscopy and undergo dramatic changes in cell volume as leaflets incline to orient perpendicular to an oblique beam of light. Communication between leaflets appears to be minimal because each leaflet has the capacity to perceive and orient towards a light beam independently of the others. Hypotheses for the mechanism of directional light perception are discussed.     

Leaflet Movement of Robinia pseudoacacia in Response to a Changing Light Environment

Diurnal and nocturnal leaflet movement of black locust （Robinia pseudoacacia L.） was investigated under three light schemes： 100% natural irradiance, 50% shading, and 90% shading. Changes in leaf mid-vein angle were described by measurements of two planes： （i） β, the angle formed by the bottom of the petiolule and its relation to the horizontal plane; and （ii） θ, the angle between the petiolule and the main leaflet vein. The two highest light regimens had a significant effect on β. Variation in β tends to make the leaflet more erect, thereby minimizing any negative impact of high irradiance on leaf lamina. Light-dark rhythms induced variation in θ （termed nyctinastic movement）. Nyctinastic movement is important during the low light levels experienced by leaflets in early morning and late afternoon. At low light levels, the leaflet stopped nyctinastic movement and θ was fixed at an angle that may have enabled the leaf lamina to maximize light interception. After the light-dark cycle was reestablished, nyctinastic movement was restored. Taken together, our results suggest that irradiance induces variation in β leading to diurnal leaflet movement （diaheliotropism）, whereas the light-dark cycle influences θ, which results in nocturnal leaflet movement. Both angles are important for describing patterns of leaf movement in R. pseudocacia.     

Response to directional light by leaves of a sun-tracking lupine (Lupinus succulentus)

Experiments were done to examine the phototrophic response of sun-tracking leaves of Lupinus succulentus Dougl. to fixed beams of white and broad band light. Upon irradiation with 15 W m⁻² white light that struck the laminae at an angle of 45°, there was a 45–60 min lag period prior to leaf movement. The greatest rate of movement was 15° h⁻¹, and reorientation ceased when leaves attained a position within 15° of perpendicular to the light beam. Laminar movement was largely pulvinar, and a 60 min inductive light treatment was sufficient to activate a maximum pulvinar response in subsequent darkness. Light striking the lamina at angles between 20 and 70° induced similar maximum pulvinar responses and only light that struck the upper (adaxial) leaf surface was effective. Leaf tracking was fully activated by blue light but not by red or yellow light.     

Petiole twisting in the crowns of<Emphasis Type="Italic"> Psychotria limonensis:</Emphasis> implications for light interception and daily carbon gain

We used Y-plant, a computer-based model of crown architecture, to examine the implications of leaf reorientation resulting from petiole bending in Psychotria limonensis (Rubiaceae) seedlings. During this reorientation process, bending of the petioles of lower leaves that are potentially self-shaded by the upper leaves rotates the lamina around the stem's orthotropic axis so that self-shading is reduced. Simulations of daily light capture and assimilation revealed a 66% increase in daily C gain due to reorientation of the leaves as compared to simulations where the leaves remained in their characteristic opposite decussate pattern set by the phyllotaxy. This was due to enhanced carbon (C) gain of the lower leaves because of the reduction of shading by upper developing leaves in the same vertical plane. The light signal for this movement was experimentally examined by placing leaf-shaped filters above already fully expanded leaves and following the resulting shade-avoiding movements. The filters were either neutral density shade cloth that reduced the photon flux density (PFD) but did not alter the red to far red ratio (R:FR) or a film that reduced the PFD equivalently but also reduced the R:FR. Leaf reorientation was much more rapid and complete under the low R:FR as compared to the high R:FR indicating involvement of a phytochrome photosensory system that detected the presence of a shading leaf. Plants in gaps were found to lack a reorientation response indicating that the reorientation is specific to the shaded understory environment.     

Photobiological and Structural Studies of Light-driven Movements in the Solar-tracking Leaf of Lupinus palaestinus Bioss. (Fabaceae)

The compound, palmate lamina of Lupinus palaestinus reorients photonastically, as well as phototropically in response to non-directional and directional light signals, respectively, by structural deformations of pulvini. When the excitation provided by directional light is maintained constant (fluence rate, angle of incidence and azimuth, with respect to the leaflet laminae), the entire lamina reorients towards it at a constant angular velocity over a considerable time interval and displacement. The laminar pulvinules are considerably longer than the subtending common petiolar pulvinus and therefore contribute most to laminar reorientation. The pulvinar region is characterized by transverse folds around its circumference, and longitudinal rib-like thickenings on the external walls of its epidermis that facilitate axial and transverse deformations. Specialized “joints”, at the distal and proximal ends of each pulvinule, contribute most to its flexing. Anthocyanin is notable by its absence. Specialized “motor” tissues surrounding the central vascular core participate in pulvinar deformation by undergoing directional and differential volume changes. The bundle sheath is characterized by numerous starch grains. The multi-layered cortical parenchyma exhibits an abundance of transversely oriented primary pit fields and associated plasmodesmata. When the leaflet lamina rotates around its midrib, the pulvinus twists along its axis, exhibiting epidermal and cortical deformation. The functional significance of these specializations is discussed.     

Phototropic leaf movements and photosynthetic performance in an amphibious fern, Marsilea quadrifolia

Diurnal phototropism has not been reported in ferns. In this study we found that the four leaflets of the amphibious fern Marsilea quadrifolia are capable of adjusting their leaflet angle and leaflet azimuth in response to changes in the position of the sun’s direct beam, exhibiting more diaphototropic movements (orienting the plane of the lamina perpendicular to incident light) in the morning and late afternoon, and more paraphototropic movements (orienting the plane of the lamina parallel to incident light) at noon. In addition, by cutting off the leaflet lamina and covering portions of leaflets with black tape, the junction between the leaflet and petiole was found to be responsible for light reception. Among the light spectrum investigated, blue light was the most effective at inducing diaphototropism. The role of diurnal phototropism in enhancing carbon return and ameliorating photoinhibition was also evaluated. It was concluded that diurnal phototropic leaf movement represents one of the plastic responses enabling this amphibious fern to grow under terrestrial conditions.     

Photonastic Control of Leaflet Orientation in Melilotus indicus (Fabaceae)

Leaflet orientation in Melilotus indicus (L.) All. Is under photonastic control during the day and nyctinastic control during the night, but also exhibits a diaphototropic (solar-tracking) response. Detached leaves with the two lateral leaflets excised were used to study the solar-tracking capability of the terminal leaflet. Perception of the photonastic excitation is located in the pulvinule. The lower (abaxial) and upper (adaxial) surfaces perceive photonastic excitation, which results in concomitant contraction of the side exposed to light and/or expansion of the opposite side. Steady state laminar elevation is determined by the fluence rates of the light incident simultaneously on the opposite sides. Light sensitivity of the lower side exceeds that of the upper. Response to photonastic excitation of either side is affected by angle of incidence of the light, but angular dependence is restricted to a limited range of angle of incidence, which differs for the two sides. This may be accounted for by the different topography of the two pulvinar surfaces and the localization in them of the light-sensitive tissues.     

Auxin‐mediated lamina growth in tomato leaves is restricted by two parallel mechanisms

In the development of tomato compound leaves, local auxin maxima points, separated by the expression of the Aux/IAA protein SlIAA9/ENTIRE (E), direct the formation of discrete leaflets along the leaf margin. The local auxin maxima promote leaflet initiation, while E acts between leaflets to inhibit auxin response and lamina growth, enabling leaflet separation. Here, we show that a group of auxin response factors (ARFs), which are targeted by miR160, antagonizes auxin response and lamina growth in conjunction with E. In wild‐type leaf primordia, the miR160‐targeted ARFs SlARF10A and SlARF17 are expressed in leaflets, and SlmiR160 is expressed in provascular tissues. Leaf overexpression of the miR160‐targeted ARFs SlARF10A, SlARF10B or SlARF17, led to reduced lamina and increased leaf complexity, and suppressed auxin response in young leaves. In agreement, leaf overexpression of miR160 resulted in simplified leaves due to ectopic lamina growth between leaflets, reminiscent of e leaves. Genetic interactions suggest that E and miR160‐targeted ARFs act partially redundantly but are both required for local inhibition of lamina growth between initiating leaflets. These results show that different types of auxin signal antagonists act cooperatively to ensure leaflet separation in tomato leaf margins.     

Role of pulvinar chloroplasts in light-driven leaf movements of the trifoliate leaf of bean (Phaseolus vulgaris L.)

Laminar pulvini of bean (Phaseolus vulgaris L.) contain numerouschloroplasts in cells of their motor tissue. The quantitativerelationships of the chloroplast pigments, chlorophyll a andb, ß-carotene, lutein, neoxanthin as well as the xanthophyllcycle carotenoids (violaxanthin, antheraxanthin and zeaxanthin)were similar to those of mesophyll chloroplasts from leafletlaminae. Exposure of pulvinules to light caused deepoxidationof violaxanthin to zeaxanthin, showing that the xanthophyllcycle is functioning. Chlorophyll fluorescence analysis of pulvinulesconfirmed that their chloroplasts are capable of both photosyntheticelectron transport and non-photochemical fluorescence quenching,showing that they build up a considerable transthylakoid protongradient in the light. Application of DCMU to excised pulvinulesand laminar discs, as well as to pulvinules of intact, attachedterminal leaflets blocked electron transport and fluorescencequenching. Application of the uncoupler CCCP to intact pulvinulesalso prevented non-photochemical fluorescence quenching. Therate of movement of the low-light-adapted terminal leaflet inresponse to exposure of its pulvinule to overhead red light(500 µmol m^–2 s^–1) was reduced when the pulvinulewas pretreated with DCMU. The pulvinar response to overheadblue light (50 µmol ^–2 s^–1), which is morepronounced than to red light, was not affected by similar pretreatmentwith DCMU. Pretreatment with CCCP caused a short lag in theresponse to red light, but did not affect its subsequent rate.The results suggest that the pulvinar response to red, but notto blue light, requires non-cyclic electron transport and theresulting generation of ATP Key words: Leaf movements, light, non-cyclic electron transport, Phaseolus, pulvinar chloroplasts     

Rhythmic Leaflet Movement in Albizzia julibrissin: Effect of Electrolytes and Temperature Alteration

The rhythmic movement of darkened Albizzia leaflets is accompanied by K⁺ flux in pulvinule motor cells whose turgor changes control opening and closing. The azide-sensitive open phase is promoted by an increase in temperature from 16 to 33C (Q₁₀ = 3), implying active transport of K⁺ ions during this period. The azide-insensitive closed phase is less temperature-sensitive and has a Q₁₀ less than 1, implying diffusion or some other physical process as the predominant pathway of K⁺ flux at this time. Thus rhythmic leaflet movement is probably due to oscillation in active K⁺ transport or membrane permeability or both. External electrolytes (0. 1 n) alter leaflet angle during the open, but not the closed, phase of the rhythm. All chlorides except NH₄⁺ promote opening, with divalent more effective than monovalent ions. Some anions promote and others inhibit opening; activity is not correlated with charge. It is likely that electrolytes alter leaflet movement by altering K⁺ flux, accomplishing this by interacting with key macromolecules in motor cell membranes.     

PHYTOCHROME CONTROLLED NYCTINASTY IN ALBIZZIA JULIBRISSIN. II. POTASSIUM FLUX AS A BASIS FOR LEAFLET MOVEMENT

Excised Albizzia leaflet pairs exposed to red (R) light close within 30–90 min after transfer to darkness. Interruption of darkness by far-red (FR) light at any time after R inhibits closure within ca. 10 min. Similarly, irradiation with R at any time after prior FR promotes closure within ca. 10 min, and the increased rate of closure is independent of the time lapse between the FR and R irradiations. Closure in the dark is inhibited by NaN₃ and DNP (5 X 10^–4 m ), by anaerobic conditions and by externally applied salts of monovalent cations, especially K; it is also temperature sensitive. Pulvinule cells are very high in K. Electron microprobe analysis of cryostated, lyophilized pulvinules reveals that during closure, K is lost from ventral cells and enters dorsal cells. FR before darkness inhibits the former but not the latter process. Thus, K flux appears to control the changes in volume of the pulvinule cells that control leaflet movement. While leaflet closure normally requires a dark period, salts of organic acids such as sodium acetate, propionate, and butyrate cause closure in the light.     

PHYTOCHROME CONTROLLED NYCTINASTY IN ALBIZZIA JULIBRISSIN. I. ANATOMY AND FINE STRUCTURE OF THE PULVINULE

Under appropriate experimental conditions, phytochrome controls leaflet closure in Albizzia by regulating differential turgor changes in motor cells of the pulvinule. Closure occurs when subepidermal dorsal cells expand and ventral cells become compressed; reopening involves the reverse changes. The internal cells surrounding the vascular core remain relatively unchanged during leaflet movement. Fine structural studies revealed several unusual features of the motor cells including: (1) fibrils oriented parallel to one another in the cytoplasmic matrix; (2) numerous spherosomes that appear to coalesce and enlarge to form vacuoles; and (3) a multivacuolate condition.     

The function, action and adaptive significance of phytochrome in light-grown plants

Abstract. It has previously been proposed that the fundamental function of phytochrome in the natural environment is the perception of the relative proportions of red and far-red light, i.e. the red: far-red ratio. This paper re-evaluates this hypothesis, for vegetative green plants, in the light of recent findings. Essentially, three issues are considered: (a) the modulation of the response to red: far-red by fluence rate: (b) the anticipation of competition for light by perception of changes in red: far-red that precede actual shading: and (c) characteristics of phytochrome that may be important in the mechanism of photoperception (i.e. the accumulation of photoconversion intermediates, and the stability of Pfr). We conclude: (a) the red: far-red ratio provides a reliable signal of plant density, even before shading by neighbours occurs: (b) plants are able to perceive and respond to these signals, and that possible ambiguities due to low red: far-red at low solar angles may be avoided by modulation of the perception process by fluence-rate dependent mechanisms; (c) although direct experimental evidence does not yet exist, circumstantial evidence suggests that the perception of red: far-red may confer positive adaptive advantage; and (d) plants of certain species perceive and respond to fluence rate changes, mediated perhaps by a blue-light absorbing photoreceptor or by phytochrome, but that these responses do not necessarily lead to shade avoidance reactions and their ecological relevance is not fully understood.     

Action dichroism in perception of vectorial photo-excitation in the solar-tracking leaf ofLavatera cretica L.

The leaf lamina ofLavatera cretica L. exhibits a diaphototropic response that discriminates between two opposite, constant vectorial excitations by white light beams whose fluence rates differ by as little as 10% (50 versus 45 μmol·m⁻²·S⁻¹). The relationship between the response (angular velocity of laminar reorientation) and the fluence-rate ratio is linear. The lamina similarly discriminates between two such excitations by polarized light, one with the electrical vector transverse to the plane of the two beams (θ) and the opposite one with the vector parallel to that plane (⪙). When two such beams were of equal fluence rate, the lamina reoriented towards the ⪙ beam. When the fluence rate of the θ beam was maintained at 50 μmol·m⁻²·s⁻¹ and that of the ⪙ beam was reduced, the response to the latter (angular velocity of laminar reorientation) was reduced progressively. Further reduction in the fluence rate of the ⪙ polarized beam eventually resulted in reorientation in the opposite direction (towards the θ beam) and the response to the latter increased progressively with the reduction in fluence rate. The equilibrium was at a ⪙/θ ratio of 0.62. Measurements of reflectance of oblique beams of ⪙ and θ polarized light from the upper laminar surface, and of transmittance of such light ghrough the lamina, eliminated the possibility that optical dichroism of the lamina contributed significantly to these results. The implications of this action dichroism to the postulated mechanism of perception of vectorial excitation by these leaves is discussed. Dedicated to the 60thbirth day of Professor Hans Mohr     

Systemic regulation of leaf anatomical structure, photosynthetic performance, and high-light tolerance in sorghum

Leaf anatomy of C3 plants is mainly regulated by a systemic irradiance signal. Since the anatomical features of C4 plants are different from that of C3 plants, we investigated whether the systemic irradiance signal regulates leaf anatomical structure and photosynthetic performance in sorghum (Sorghum bicolor), a C4 plant. Compared with growth under ambient conditions (A), no significant changes in anatomical structure were observed in newly developed leaves by shading young leaves alone (YS). Shading mature leaves (MS) or whole plants (S), on the other hand, caused shade-leaf anatomy in newly developed leaves. By contrast, chloroplast ultrastructure in developing leaves depended only on their local light conditions. Functionally, shading young leaves alone had little effect on their net photosynthetic capacity and stomatal conductance, but shading mature leaves or whole plants significantly decreased these two parameters in newly developed leaves. Specifically, the net photosynthetic rate in newly developed leaves exhibited a positive linear correlation with that of mature leaves, as did stomatal conductance. In MS and S treatments, newly developed leaves exhibited severe photoinhibition under high light. By contrast, newly developed leaves in A and YS treatments were more resistant to high light relative to those in MS- and S-treated seedlings. We suggest that (1) leaf anatomical structure, photosynthetic capacity, and high-light tolerance in newly developed sorghum leaves were regulated by a systemic irradiance signal from mature leaves; and (2) chloroplast ultrastructure only weakly influenced the development of photosynthetic capacity and high-light tolerance. The potential significance of the regulation by a systemic irradiance signal is discussed.     

Why does Viola hondoensis (Violaceae) shed its winter leaves in spring?

? Premise of the study: Viola hondoensis is a perennial herb that inhabits the understory of temperate, deciduous forests. It is an evergreen plant with a leaf life span that is shorter than a year. Its summer leaves are produced in spring and shed in autumn; winter leaves are produced in autumn and shed in spring. Here we asked why the plant sheds its winter leaves in spring, though climate conditions improve from spring to summer. We proposed four hypotheses for the cause of shedding: (1) changes in seasonal environment such as day length or air temperature, (2) shading by canopy deciduous trees, (3) self-shading by taller summer leaves, and (4) competition for nutrients between summer and winter leaves. ? Methods: To test these hypotheses, we manipulated the environment of winter leaves: (1) plants were transplanted to the open site where there was no shading by canopy trees. (2) Petioles of summer leaves were anchored to the soil surface to avoid shading of winter leaves. (3) Sink organs were removed to eliminate nutrient competition. ? Key results: Longevity of winter leaves was extended when shading by summer leaves was eliminated and when sink organs were removed, but not when plants were transplanted to the open site. ? Conclusion: We conclude that the relative difference in light availability between summer and winter leaves is a critical factor for regulation of leaf shedding, consistent with the theory of maximization of the whole-plant photosynthesis.     

弱光胁迫影响夏玉米光合效率的生理机制初探

大田条件下, 以普通夏玉米(Zea mays) ‘泰玉2号’为材料, 于授粉后1-20天遮光55% (+S), 以大田自然光照条件下生长的玉米作为对照(-S), 研究了遮光及恢复过程中玉米植株的光合性能、叶绿体荧光参数、叶黄素循环以及光能分配的变化, 初步揭示夏玉米开花后弱光条件下光适应的生理机制, 为玉米高产稳产提供理论依据。结果表明, 遮光后玉米穗位叶叶绿素含量及可溶性蛋白含量均减少, RuBP羧化酶和PEP羧化酶活性显著降低, 导致穗位叶净光合速率(P_n)迅速下降, 光饱和点也明显降低; 恢复初期P_n迅速升高, 光合关键酶活性有所增强。遮光后植株的最大光化学效率(F_v/F_m)、实际光化学效率(Ф_PSII)显著降低, 非光化学淬灭(NPQ)则显著升高, 而恢复初期植株穗位叶Ф_PSII有所升高, 表明突然暴露在自然光下的光合电子传递速率明显加快, 这与其光合速率及光合酶活性的趋势保持一致; 遮光处理对穗位叶叶黄素循环库的大小(紫黄质+花药黄质+玉米黄质(V + A + Z))影响不显著, 但使叶黄素循环的脱环氧化状态(A + Z)/(V + A + Z)增加; 遮光后植株分配于光化学反应的光能明显减少, 天线耗散光能比率显著增加, 恢复过程中植株主要以过剩非光化学反应的形式耗散过剩的光能。遮光后及恢复初期, 玉米植株的PSII原初光化学活性明显下降, 限制了光合碳代谢的电子供应从而抑制了光合作用, 主要依赖叶黄素循环途径进行能量耗散, 而在光照转换后遮光的玉米叶片在适应自然光过程中的光保护机制不断完善, 光合能力逐渐得到 恢复。     

SUN-TRACKING AND RELATED LEAF MOVEMENTS IN A DESERT LUPINE (LUPINUS ARIZONICUS)

Both diaheliotropic (sun-tracking) and paraheliotropic (cupping) leaf movements are described for the Arizona Lupine [Lupinus arizonicus (Wats.) Wats.]. The leaf movements are shown to be non-circadian in nature. Evidence is presented that an active K⁺ ion transport mechanism is involved in these turgor-related leaf movements. Increasing concentrations of lanthanum, a known ion transport inhibitor, showed increasing inhibition of both leaf movements. Increasing concentrations of other salts did not inhibit either leaf movements, instead there was an increase in the cupping leaf movement (elevation of the leaflets) which is shown to be a water stress response.     

Nitrogen allocation in response to partial shading of a plant: Possible mechanisms

The significance of photosynthetic and transpiration rates for the perception by plants of light gradients in leaf canopies has been investigated with regard to nitrogen allocation and re-allocation. A gradient of photon flux density (PFD) over a plant's foliage was simulated by shading one leaf of a pair of primary leaves of bean ( Phaseolus vulgaris L. cv. Rentegever). Photosynthetic rate was manipulated independently of PFD and, to some extent, also of transpiration, by subjecting the leaf to different CO₂ concentrations. Transpiration rate was changed independently of PFD and photosynthetic rate by subjecting the leaf to different vapour pressure differences (VPD). A reduced partial pressure of CO₂ reduced specific leaf mass (SLM) as did a decreased PFD, but did not change leaf N per unit area (N_LA) and light saturated rate of photosynthesis (A_max). A reduced VPD caused several effects consistent with the effect of PFD. It decreased N_LA and A_max and increased the chlorophyll to N ratio in old and young leaves. Furthermore, it decreased the chlorophyll a to b ratio and inhibited leaf growth in young leaves. The transpiration stream is partitioned among the leaves of a plant according to their transpiration rates. The results suggest that relative rates of import of xylem sap into leaves of a plant play an important role in the perception of partial shading of a plant, a situation normally found in dense vegetations. The possible role of cytokinin influx into leaves as controlled by transpiration rate, is discussed.