Effects of vanadate, N2 and light on the membrane potential of motor cells and the lateral leaflet movements of Desmodium motorium

The lateral leaflets of Desmodium motorium (Houtt.) Merr. exhibit ultradian up- and down movements, which are paralleled by oscillations of the membrane potential of motor cells in the pulvinus. By different treatments we have tested the hypothesis that both that both oscillation-types are causally related. The reactions of the leaflet movement and the membrane potential were evaluated by the following approaches. (1) Application of vanadate. an inhibitor of the proton pump in the plasmalemma. and N₂ suppressed leaflet movements and finally arrested the leaflet in the lower position. Before the oscillations damped out, a strong lengthening in period was found. This indicates that the pump is part of the ultradian clock. A period lenthening and a final suppression of the rhythm by vanadate was also seen in the extracellular electric potential of the pulvinus. Intracellular recordings in situ showed that vanadate application depolarized the motor cells. (2) Light of high fluence rates diminished the amplitude of the oscillations of the membrane potential of single motor cells and shortened the period. The same effects were observed when monitoring the lateral leaflet movement. The leaflet always moved towards the direction of the light. whether it was applied from the abaxial or from the adaxial part of the pulvinus. (3) When light was applied to the pulvinus of lateral leaflets. which had spontancously stopped moving in an upper position. oscillations were induced transiently. This effect was also found for the membrane potential of motor cells in the pulvinus. - Our results thus provide further evidence that the membrane potential controls the volume state of the motor cells in the pulvinus of lateral leaflets of Desmodium motorium .     

Studies on leaflet abscission of blue lupin leaves

Summary In blue lupin leaves, each leaflet abscises at an abscission zone situated in the pulvinus at its base. The time to abscission of leaflets of detached leaves is proportional to leaf age. Light accelerates abscission; within certain limits the acceleration is the greater the younger the leaf. At a given concentration, kinetin applied to a single leaflet accelerates leaflet abscission in young leaves kept in darkness, delays it in older ones. There is an interaction between kinetin and light which is dependent also on leaf age and kinetin concentration. The leaf can be considered as consisting of three regions, the petiole, the pulvinar region and the leaflets. The effects of kinetin and of light as well as their interactions depent on the regions of the leaf treated with these agents. Kinetin applied to a leaflet of a young leaf kept in darkness accelerates abscission, but kinetin applied to the pulvinar region of a similar leaf kept in darkness delays abscission. When any part of a leaf is illuminated, abscission is accelerated. The most light-sensitive region of the leaf is the pulvinar region, despite its relatively small area. Acceleration of abscission by light is greatest when illumination of the pulvinar region is combined with illumination of either the leaflets or the petiole. The interaction of light with kinetin is complex. Where the illuminated area includes the pulvinar region, kinetin delays abscission. This effect is most marked in the case where the pulvinar region alone is illuminated and kinetin is applied to a leaflet.Intrafoliar abscission as found in lupin leaves permits study of complex interactions of both distal and proximal stimuli involved in abscission.     

Phase response curve for the ultradian rhythm of the lateral leaflets of Desmodium gyrans using DC current pulses

In the present study the leaf movement rhythm was perturbed by the application of DC current pulses (15 microA, 10 seconds, voltage applied: 10 V) to the upper part of the pulvinus, passing through the pulvinus and its stalk. The pulses were applied at four different positions of the leaflets: when the leaves were at the lowermost position, when moving up, at the uppermost position and when moving down. The pre-perturbed and the post-perturbed rhythms were compared. We found that the rhythms were shifted in phase and the phase shifts observed at the four different positions of the leaflets were significantly different in magnitude as well as direction. Furthermore, we could also observe phase advances, which is in contrast to an earlier finding. A phase response curve (PRC) was constructed to illustrate the sensitivity of the oscillating leaflet system to DC pulses. Substantial delays of about 50 s (as compared to the period of about 200 s) were obtained when pulses were administered at the lowermost position and when leaflet were moving upwards, while advances or no phase shifts were recorded in the uppermost position and when leaflet were moving down respectively.     

Light perception for sun-tracking is on the lamina in Crotalaria pallida (Fabaceae)

Trifoliolate leaves of Crotalaria pallida Aiton (Fabaceae) exhibit sun-tracking behavior in simulated days and in response to a fixed, oblique light. The site of light perception for sun-tracking is on the lamina, necessitating transport of a signal from the lamina to the site of response (the pulvinule at the base of the leaflet). Evidence for the site of light perception on the lamina includes the following: (1) leaflet movement in response to oblique light is not affected by shading the pulvinule or by illuminating the pulvinule with vertical light; (2) leaflet movement is stopped by shading the lamina while illuminating the pulvinule with oblique light. The proximal end of the leaflet is the most sensitive region for light perception. Light reaching the midveins is not necessary for leaflet reorientation. Presentation times of as little as 10 min followed by darkness resulted in partial leaflet movement in young leaves. This indicates that the signal was not inductive in nature. Estimates of the rate of signal transport range from 3 to 12 cm/h, within the range of phloem transport.     

Phytochrome and Circadian Clocks in Samanea: Rhythmic Redistribution of Potassium and Chloride within the Pulvinus during Long Dark Periods

Previous investigations with the electron microprobe reveal that the movements of Samanea leaflets are correlated with massive redistribution of K within the pulvinus. Evidence is now presented that Cl moves with K, whether plants are in white light or darkness, whether or not the amplitude of free running oscillations has damped, and whether or not the rhythm has been rephased by phytochrome photoconversion. The mid-extensor to mid-flexor ratio of K + Cl is correlated with leaflet angle under all conditions. Total Cl in both inner cortex and motor region is approximately 0.6 as high as K. The stoichiometry between Cl and the migratory fraction of K is close to, but not precisely 1:1 in all regions of the pulvinus, suggesting that other ions or systems may also be involved in the balancing of electrical charges.     

HELIOTROPIC LEAF MOVEMENT RESPONSE TO H+/ATPase activation,H+/ATPase inhibition,AND K+ CHANNEL INHIBITION IN VIVO

The pulvinus, located at the base of soybean leaflets, is both the light perception and motor organ for heliotropic leaf movements. Our objective was to investigate the role of plasma membrane H⁺/ATPase and TEA-sensitive K⁺ channels in mediating pulvinar response to light. The plasma membrane H+/ATPase activator, fusicoccin, plasma membrane H⁺/ATPase inhibitors, vanadate and erythrosin-B, and the K⁺ channel blocker TEA were introduced to the intact pulvinus through the transpiration stream. The pulvinus was illuminated by a vertical light beam of 1,400 μmol m^-2 s^-1 to stimulate leaf movement. Leaf orientation was measured every 5 min for 60 min of illumination. All compounds tested inhibited pulvinar bending, but concentration and uptake time required for inhibition varied: 12.5 μM fusicoccin reduced leaf movement after 3 hr uptake, 2 mM vanadate reduced leaf movement after 6 hr uptake, 100 μM erythrosin-B reduced leaf movement after 3 hr uptake, and 15 mM TEA reduced leaf movement after 6 hr uptake. In all cases final leaf angle was reduced by higher concentrations and/or increased time for uptake of the chemical into the pulvinus. Results support the hypothesis that the proximal mechanism of heliotropic movement is similar to that of nyctinastic movements.     

Oscillations of the Membrane Potential of Pulvinar Motor Cells In Situ in Relation to Leaflet Movements of Desmodium motorium

The ultradian rhythmic movement of the lateral leaflets of Desmodiummotorium is accompanied by rhythmic changes of the extra- andintracellular electrical potentials in the pulvinus, which aremeasured in situ in the pulvinus against the bathing solutionof the petiole. Extra- and intracellular potentials oscillatewith 180'b0 phase difference to each other, as shown by simultaneousmeasurements of both types of potentials in the abaxial partof the pulvinus. Light-induced changes of these potentials movein opposite directions. The in situ membrane potential of themotor cells of the pulvinus was calculated from the differencebetween the extra- and intracellular potentials. It was foundto oscillate between –136 and –36 mV, in phase withthe intracellular and inverse to the extracellular potential.The phase relationship between the leaflet movement rhythm andthe in situ membrane potential rhythm was as follows: downwardmovement is preceded and accompanied by a strong depolarization,upward movement by hyperpolarization. Our results suggest that membrane depolarization in pulvinarmotor cells of Desmodium motorium drives and controls potassiumefflux and hyperpolarization potassium influx via potassiumchannels. Key words: Desmodium pulvinus, leaf movement, pulvinar motor cells, electrical potential     

Characterization of X-ray induced increase of mitotic cross-overs in Glycine max

Summary Three types of spots can be identified on the leaves of heterozygous light green, Y/y, Glycine max (L.) Merrill: dark green (D) and aurea (A) single spots (resembling the phenotypes of the homozygotes) and double (Db) spots consisting of adjacent D and A tissue. X-irradiation increased the frequency of each type of spot on simple and first compound leaves. The Db spots, indicative of mitotic crossing-over (MCO), increase linearly with increasing dosage. Moisture content of the seeds was independent of the rate of spot increase. At high dosages morphological alterations were observed, including spots on homozygotes, leaf area reduction, smaller seedlings, and abnormal leaf shapes. The frequency of light green spots on normal dark green, Y/Y, seedlings was tabulated and, as with all other spot types, increased with increasing X-ray dosage. Dormant soybean seeds contain leaf primordia of both simple and first compound leaves. Mature simple leaves contained more spots, reflecting a larger primordial cell number, while first compound leaves had larger spots, since each affected cell underwent more mitoses prior to leaf maturation. Within first compound leaves, the terminal leaflets developed asynchronously in relation to the lateral leaflets. Terminal leaflets were shown to be initiated first, have a larger percentage of the leaflet area covered with spots, and have larger mature leaflet area. The spontaneous rate of MCO, 3.39×10^–5 MCO events per mitosis, was increased 282-fold by 1600 R. We also ascertained that Mitomycin C is more specific for Db spot induction than X-rays. These results are compared with our similar irradiation experiments on tobacco shoot apices.     

Relationships between structure and motility ofAlbizzia motor organs: Changes in ultrastructure of cortical cells during dark-induced closure

Summary Paired leaflets ofAlbizzia julibrissin spread apart (open) in the daytime and fold together (close) at night. We examined the structure of cells in open and closedAlbizzia motor organs (pulvini) to identify reversible changes in structure associated with motility. Pulvini were fixed in glutaraldehyde and stained using conventional methods. The pulvinus has a central vascular cylinder bordered by thick-walled collenchyma cells, in turn surrounded by an endodermis and many layers of cortical parenchyma. Cortical cells in the extensor undergo large changes in shape during leaflet closure linked with: formation of wall infoldings, development of a large periplasmic space filled with fibrils and membranes, development of lobes on the nucleus, evagination of the nuclear outer envelope membrane, break-up of the large central vacuole to form many small vacuoles, and linking of the plasmalemma to inner regions of the cytoplasm by microfilaments. Cortical cells in the flexor, by contrast, remain relatively stable during leaflet movement. Microtubules are present near the plasmalemma in both extensor and flexor cells; in the extensor, spherical coated vesicles are located near the microtubules. The possible function of these structures in regulating intracellular shuttling processes is discussed.     

Dynamic aspects of the response of the pulvinus in the leaf of bean plants(Phaseolus vulgaris L.) to photoexcitation

The pulvinus of bean (Phaseolus vulgaris L) responds to unilateral photo-excitation by phototropic curvature. Osmotically active solutes and water are transported from its exposed to the opposite sector of its motor tissue, resulting in differential changes in turgor pressure in these sectors and generation of a trans-pulvinar torque. A null-point approach was used to non-invasively study these dynamic changes in the terminal leaflet of bean. A variable torque was applied perpendicular to the midrib, to restrain laminar movement by precisely and continuously counteracting the generated torque. This equilibrium prevented curvature of the pulvinus and the associated opposite axial changes in volume in the opposite sectors of its motor tissue. The laminar torques measured were used to estimate stresses (changes in turgor pressure) generated within the motor tissue. These stress values were used to derive the corresponding changes in osmotic pressures and in solute concentration. Skotonastically downfolded leaflets were excited with white light to study their combined dynamic response to photonastic and phototropic excitation. Photonastically unfolded (horizontal) leaflets were excited with blue and red light, alone and in combination, to determine the spectral dependence of the dynamic pulvinar responses tophototropic excitation by itself.     

Acropetal leaflet initiation of Eschscholzia californica is achieved by constant spacing of leaflets and differential growth of leaf

In compound leaves, leaflet primordia are initiated directionally along the lateral sides. Our understanding of the molecular basis of leaflet initiation has improved, but the regulatory mechanisms underlying spatio-temporal patterns remain unclear. In this study, we investigated the mechanisms of acropetal (from the base to the tip) progression of leaflet initiation in Eschscholzia californica. We established an ultraviolet-laser ablation system to manipulate compound-leaf development. Local ablation at the leaflet incipient site generated leaves with asymmetric morphology. In the majority of cases, leaflets that were initiated on the ablated sides shifted apically. Finite time-course observation revealed that the timing of leaflet initiation was delayed, but the distance from the leaf tip did not decrease. These results were suggestive of the local spacing mechanism in leaflet initiation, whereby the distance from the leaf tip and adjacent pre-existing leaflet determines the position of leaflet initiation. To understand how such a local patterning mechanism generates a global pattern of successive leaflet initiation, we assessed the growth rate gradient along the apical–basal axis. Our time-course analysis revealed differential growth rates along the apical–basal axis of the leaf, which can explain the acropetal progression of leaflet initiation. We propose that a leaflet is initiated at a site where the distances from pre-existing leaflets and the leaf tip are sufficient. Furthermore, the differential growth rate may be a developmental factor underlying the directionality of leaflet initiation.     

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.     

Use of an oriented transmembrane protein to probe the assembly of a supported phospholipid bilayer.

Planar-supported phospholipid bilayers formed by the adsorption of vesicles are increasingly used in the investigation of lipid-dependent reactions. We have studied the way in which these bilayers are formed with phospholipid vesicles containing the transmembrane protein Tissue Factor (TF). TF complexed with the serine protease, factor VIIa, is the primary initiator of blood coagulation by way of activation of the zymogen factor X. TF has been shown to orient randomly on the inner and outer leaflets of vesicles. We used proteolytic digestion to produce vesicles in which the extracellular domain of TF is located on the inner leaflet. These vesicles show no cofactor activity for factor VIIa as a result of the inability of the extracellular domain of TF to bind VIIa. After freeze/thawing, 50% of the cofactor activity was regained, indicating reorientation of the sequestered, inner leaflet TF. Adsorption of these vesicles to the inner surface of glass microcapillaries results in a continuous phospholipid bilayer. The microcapillaries were perfused with a solution of factors VIIa and X, and the effluent was monitored for factor Xa production, a sensitive measure of the activity of the TF-VIIa complex. For coatings produced with the digested vesicles, minimal TF-VIIa activity was observed, showing that the supported bilayer preserves the orientation of the leaflets in the vesicles, i.e., the outer leaflet of the vesicles forms the outer leaflet of the supported bilayer.     

Elemental analysis of freeze-dried thin sections of Samanea motor organs: barriers to ion diffusion through the apoplast

Leaflet movements in the legume Samanea saman are dependent upon massive redistribution of potassium (K), chloride (Cl), and other solutes between opposing (extensor and flexor) halves of the motor organ (pulvinus). Solutes are known to diffuse through the apoplast during redistribution. To test the possibility that solute diffusion might be restricted by apoplastic barriers, we analyzed elements in the apoplast in freeze-dried cryosections of pulvini using scanning electron microscopy/x-ray microanalysis. Large discontinuities in apoplastic K and Cl at the extensor-flexor interface provide evidence for a barrier to solute diffusion. The barrier extends from the epidermis on upper and lower sides of the pulvinus to cambial cells in the central vascular core. It is completed by hydrophobic regions between phloem and cambium, and between xylem rays and surrounding vascular tissue, as deduced by discontinuities in apoplastic solutes and by staining of fresh sections with lipid-soluble Sudan dyes. Thus, symplastic pathways are necessary for ion redistribution in the Samanea pulvinus during leaflet movement. In pulvini from leaflets in the closed state, all cells on the flexor side of the barrier have high internal as well as external K and Cl, whereas cells on the extensor side have barely detectable internal or external K or Cl. Approximately 60% of these ions are known to migrate to the extensor during opening; all return to the flexor during subsequent closure. We propose that solutes lost from shrinking cells in the outer cortex diffuse through the apoplast to plasmodesmata-rich cells of the inner cortex, collenchyma, and phloem; and that solutes cross the barrier by moving through plasmodesmata.     

Genetic transfer of a twospotted spider mite (Acari: Tetranychidae) repellent in tomato hybrids

Lycopersicon hirsutum Dunal is very resistant to arthropod herbivory, and research on causes of resistance has often implicated trichomes and their secretions. To better understand relationships among resistance, repellency, and 2,3-dihydrofarnesoic acid, a trichome-borne sesquiterpenoid spider mite repellent, two tomato, Lycopersicon esculentum Miller, varieties were interbred with a highly resistant, spider mite repellent accession (LA1363) of L. hirsutum. Backcross and F2 generations were produced with each tomato variety. Whole leaves of 99 hybrids were bioassayed with twospotted spider mites, Tetranychus urticae Koch, allowing selection of six hybrids (two susceptible and four resistant) for each generation of each family. When these 24 hybrids were characterized for spider mite repellency with thumbtack bioassays, two hybrids had repellent leaflets, demonstrating that repellency was genetically transferred to interspecific tomato hybrids. Leaflet washes containing trichome secretions from each of three hybrids, including the two having repellent leaflets, were repellent in bridge bioassays. For the two hybrids having repellent leaflets and leaflet washes, removal of trichome secretions by dipping leaflets in methanol eliminated leaflet repellency. 2,3-Dihydrofarnesoic acid was present in trichome secretions of the hybrids having leaflet repellency, and it also was present in secretions of other hybrids, indicating that its presence is essential, but not sufficient for leaflet repellency. With regard to resistance, 16 of the hybrids tested had been identified as resistant in a whole leaf bioassay, but only two had repellent leaflets, indicating that other mechanisms of resistance are present in the resistant L. hirsutum parent.     

Light-driven leaf movements*

Abstract. Mature leaves of many plants re-orientate their laminae photonastically in response to non-directional light signals, and/or phototropically in response to directional light signals, by flexing of pulvini, most commonly subtending their bases. Physiological and structural specializations of the pulvinus enable it to flex, by rapidly undergoing differential and repeatedly reversible axial volume changes (expansion/contraction) in opposite sectors of its motor tissue. Light-driven leaf movements are adaptations that contribute to the efficiency of the photosynthetic apparatus in the leaf. The phototropic response maximizes the harvesting of photosynthetically radiant energy. The photonastic response to dark-to-light transitions increases the interception of light by unfolding the lamina. Another photonastic response modulates the interception of radiant energy by the lamina, allowing it to evade damage by light in excess of its photosynthetic capacity when the leaf is under stress. The same unidentified blue-absorbing pigment system appears to be involved in all these responses. Non-directional light signals are perceived in the pulvinus. Perception of directional light signals may be localized in other parts of the leaf in different plants: for example, the pulvinus in most leguminous species, and the lamina in malvaceous and at least one leguminous species. The perception of non-directional and directional light signals, their transduction to differential volume changes in the target cells, and their transmission between the two where the sites are separate, are discussed.     

A method to record circadian plant movements, with application to Oxalis leaf rhythms

A transducer was developed to record the circadian movement of the individual leaflets in Oxalis regnellii Mig. The method can easily be adapted to measure other kinds of plant movements as well. It is based on the detection of the shadow each leaflet casts on the small side of a specially formed Perspex plate. The light is guided through the Perspex and collected by a phototransistor, which provides an electrical signal that is proportional to the light intensity falling onto it. The output signal can be made a linear function of the leaf angle. This equipment was used in experiments to study the coupling between the 3 leaflets in Oxalis . Pulses of 4 h of red light were given to one of the leaflets, the two others were shielded from the light. A phase response curve was determined for each leaflet, but there was no significant difference in the phase response between the 3 leaflets. Experiments were also made in which the 3 leaflets were separated physically by cuts along the petiole between the pulvini. In this case ultradian oscillations were observed.     

Reduced leaf complexity in tomato wiry mutants suggests a role for PHAN and KNOX genes in generating compound leaves

Recent work on species with simple leaves suggests that the juxtaposition of abaxial (lower) and adaxial (upper) cell fates (dorsiventrality) in leaf primordia is necessary for lamina outgrowth. However, how leaf dorsiventral symmetry affects leaflet formation in species with compound leaves is largely unknown. In four non-allelic dorsiventrality-defective mutants in tomato, wiry, wiry3, wiry4 and wiry6, partial or complete loss of ab-adaxiality was observed in leaves as well as in lateral organs in the flower, and the number of leaflets in leaves was reduced significantly. Morphological analyses and expression patterns of molecular markers for ab-adaxiality [LePHANTASTICA (LePHAN) and LeYABBY B (LeYAB B)] indicated that ab-adaxial cell fates were altered in mutant leaves. Reduction in expression of both LeT6 (a tomato KNOX gene) and LePHAN during post-primordial leaf development was correlated with a reduction in leaflet formation in the wiry mutants. LePHAN expression in LeT6 overexpression mutants suggests that LeT6 is a negative regulator of LePHAN. KNOX expression is known to be correlated with leaflet formation and we show that LeT6 requires LePHAN activity to form leaflets. These phenotypes and gene expression patterns suggest that the abaxial and adaxial domains of leaf primordia are important for leaflet primordia formation, and thus also important for compound leaf development. Furthermore, the regulatory relationship between LePHAN and KNOX genes is different from that proposed for simple-leafed species. We propose that this change in the regulatory relationship between KNOX genes and LePHAN plays a role in compound leaf development and is an important feature that distinguishes simple leaves from compound leaves.     

MORPHOLOGY AND VASCULATURE OF THE LEAF OF POTATO (SOLANUM TUBEROSUM)

The leaf and stem of the potato plant (Solanum tuberosum L. cv. Russet Burbank) were studied by light microscopy to determine their morphology and vasculature; scanning electron microscopy provided supplemental information on the leaf's morphology. The morphology of the basal leaves of the potato shoot is quite variable, ranging from simple to pinnately compound. The upper leaves of the shoot are more uniform, being odd pinnate with three major pairs of lateral leaflets and a number of folioles. The primary vascular system of the stem is comprised of six bundles, three large and three small ones. The three large bundles form a highly interconnected system through a repeated series of branchings and arch-producing mergers. Two of the three large bundles give rise to short, lateral leaf traces at each node. Each of the small bundles in the stem is actually a median leaf trace which extends three internodes before diverging into a leaf. The three leaf traces enter the petiole through a single gap; thus the nodel anatomy is three-trace unilacunar. Upon entering the petiole, each of the laterals splits into an upper and a lower lateral. Whereas the upper laterals diverge entirely into the first pair of leaflets, the lower laterals feed all of the lateral leaflets through a series of bifurcations. Prior to their entering the terminal leaflet, the lower laterals converge on the median bundle to form a single vascular crescent which progresses acropetally into the terminal leaflet as the midvein, or primary vein. In the midrib, portions of the midvein diverge outward and continue as secondaries to the margin on either side of the lamina. Near the tip of the terminal leaflet, the midvein consists of a single vascular bundle which is a continuation of the median bundle. Six to seven orders of veins occur in the terminal leaflet.     

AXIAL ROTATION OF ERYTHRINA HERBACEA LEAFLETS

The leaflets of Erythrina herbacea, a leguminous plant native to subtropical North America, undergo daily movements. During the night, leaflets are oriented with their tips pointing downwards and show no rotation around the midrib of each leaflet. In the morning, leaflets rise, passing through a horizontal orientation and, somewhat before noon, reach a maximum average midrib elevation of approximately sixty degrees above the horizontal plane. Simultaneously, each leaflet rotates about the midrib by an average angle of forty to fifty degrees. These leaflet movements result in an initial increase in the relative leaflet surface area presented to the sun during the morning, a decrease in interception of direct solar radiation around noon, followed by a second maximum in light interception in early afternoon.