Chloroplast avoidance movement is not functional in plants grown under strong sunlight

Chloroplast movement in nine climbing plant species was investigated. It is thought that chloroplasts generally escape from strong light to avoid photodamage but accumulate towards weak light to perform photosynthesis effectively. Unexpectedly, however, the leaves of climbing plants grown under strong sunlight showed very low or no chloroplast photorelocation responses to either weak or strong blue light when detected by red light transmittance through leaves. Direct observations of Cayratia japonica leaves, for example, revealed that the average number of chloroplasts in upper periclinal walls of palisade tissue cells was only 1.2 after weak blue‐light irradiation and almost all of the chloroplasts remained at the anticlinal wall, the state of chloroplast avoidance response. The leaves grown under strong light have thin and columnar palisade tissue cells comparing with the leaves grown under low light. Depending on our analyses and our schematic model, the thinner cells in a unit leaf area have a wider total plasma membrane area, such that more chloroplasts can exist on the plasma membrane in the thinner cells than in the thicker cells in a unit leaf‐area basis. The same strategy might be used in other plant leaves grown under direct sunlight.     

Photochemical and photophosphorylation activities of chloroplasts and leaf mesostructure in Chinese cabbage plants grown under illumination with light-emitting diodes

Leaf mesostructure, photochemical activity, and chloroplast photophosphorylation (PP) in the fourth true leaf of 28-day-old Chinese cabbage (Brassica chinensis L.) plants were investigated. Plants were grown under a light source based on red (650 nm) and blue (470 nm) light-emitting diodes (LED) with red/blue photon flux ratio of 7: 1 and under illumination with high-pressure sodium lamp (HPSL) at photon flux densities of 391 ± 24 μmol/(m² s) (“normal irradiance”) and 107 ± 9 μmol/(m² s) (“low irradiance”) in photosynthetically active range. At normal irradiance, the leaf area in plants grown under HPSL was twofold higher than in LED-illuminated plants; other parameters of leaf mesostructure were little affected by spectral quality of incident light. The lowering of growth irradiance reduced the majority of leaf mesostructure parameters in plants grown under illumination with HPSL, whereas in LED-illuminated plants the lowered irradiance reduced only specific leaf weight but increased the leaf thickness and dimensions of mesophyll cells and chloroplasts. The photochemical activity of isolated chloroplasts was almost independent of growth irradiance and light spectral quality. Light quality and intensity used for plant growing had a considerable impact on PP in chloroplasts. At normal light intensity, the highest activity of noncyclic PP in chloroplasts was observed for plants grown under HPSL; at low light intensity the highest rates of PP were noted for plants grown under LED. The P/2e⁻ ratio, which characterizes the degree of PP coupling to electron transport in the chloroplast electron transport chain, showed a similar pattern. Thus, the narrow-band spectrum of the light source had little influence on leaf mesostructure and electron transport rates. However, this spectrum significantly affected the chloroplast PP activity. The PP patterns at low and normal light intensities were opposite for plants grown under LED and HPSL light sources. We suppose that growing plants under LED array at normal light intensity disturbed the chloroplast coupling system, thus preventing the effective use of light energy for ATP synthesis. At low light intensity, chloroplast PP activity was significantly higher under LED illumination, but plant growth was suppressed because of impaired adaptation to low light intensity.     

A New Model for Leaf Photosynthesis Incorporating the Gradients of Light Environment and of Photosynthetic Properties of Chloroplasts within a Leaf

A leaf photosynthesis model was constructed based upon the notionthat the leaf photosynthesis is a summation of photosynthesisof each chloroplast under in situ micro-environmental conditions.Intra-leaf light environment was calculated using the valuesof transmittance and reflectance of leaf tissues reported previously.Simulations of light response curves of whole leaf photosynthesiswere carried out for the model leaves with different patternsof gradients in light environment and/or in photosynthetic activities.The results indicate that the higher absorption coefficientof chlorophyll in spongy tissue than in palisade tissue andintra-leaf vertical gradient in photosynthetic activity of thechloroplasts as reported for real dorsiventral leaves are bothadvantageous to the productivity of the leaf because these propertiesraise the efficiency of the light utilization. Intra-leaf light environment, leaf photosynthesis, light utilization, palisade tissue, photosynthetic productivity, spongy tissue     

Structural Adaptation of the Leaf Mesophyll to Shading

Structural characteristics of the mesophyll were studied in five boreal grass species experiencing a wide range of light and water supply conditions. Quantitative indices of the palisade and spongy mesophyll tissues (cell and chloroplast sizes, the number of chloroplasts per cell, the total cell and chloroplast surface area per unit leaf surface area) were determined in leaves of each of the species. The cell surface area and the cell volume in spongy mesophyll were determined with a novel method based on stereological analysis of cell projections. An important role of spongy parenchyma in the photosynthetic apparatus was demonstrated. In leaves of the species studied, the spongy parenchyma constituted about 50% of the total volume and 40% of the total surface area of mesophyll cells. The proportion of the palisade to spongy mesophyll tissues varied with plant species and growth conditions. In a xerophyte Genista tinctoria, the total cell volume, cell abundance, and the total surface area of cells and chloroplasts were 30–40% larger in the palisade than in the spongy mesophyll. In contrast, in a shade-loving species Veronica chamaedris, the spongy mesophyll was 1.5–2 times more developed than the palisade mesophyll. In mesophyte species grown under high light conditions, the cell abundance and the total cell surface area were 10–20% greater in the palisade mesophyll than in the spongy parenchyma. In shaded habitats, these indices were similar in the palisade and spongy mesophyll or were 10–20% lower in the palisade mesophyll. In mesophytes, CO₂ conductance of the spongy mesophyll accounted for about 50% of the total mesophyll conductance, as calculated from the structural characteristics, with the mesophyll CO₂ conductance increasing with leaf shading.     

The spatial distribution of chloroplast water in Acer platanoides sun and shade leaves

We evaluated a new, two-dimensional (2-D) nuclear magnetic resonance (NMR) imaging technique as a method for measuring the distribution of chloroplasts in leaves. NMR images that showed the distribution of chloroplast water and of total water as a function of depth into Acer platanoides sun and shade leaves were compared with the distribution of chlorophyll in the same leaf types (as measured by fluorescence microscopy), with the cellular structure (by scanning electron microscopy), and with published information. Results showed that the volume fraction of chloroplast water was much larger in shade than in sun leaves, and that it averaged about one-third larger in the palisade than in the spongy parenchyma region of both leaf types. Chlorophyll fluorescence was more intense in shade than in sun leaves. In sun leaves, fluorescence was maximal in the palisade region near the junction with the spongy parenchyma, while in shade leaves, fluorescence was maximal in the upper part of the spongy layer. We concluded that 2-D NMR imaging reliably indicates the location of chloroplast water.     

CORRELATES OF LEAF OPTICAL PROPERTIES IN TROPICAL FOREST SUN AND EXTREME-SHADE PLANTS

Thirteen shade-adapted rain forest species were compared with twelve sun-adapted tropical forest species for correlates to leaf optical properties (described previously in Amer. J. Bot. 73: 1100–1108). The two samples were similar in absorptance of quanta for photosynthesis, but the shade-adapted taxa: 1) had significantly lower specific leaf weights, indicating a more metabolically efficient production of surface for quantum capture; 2) synthesized less chlorophyll per unit area; and 3) used less chlorophyll for capturing the same quanta for photosynthesis. The anatomical features that best correlate with this increased efficiency are palisade cell shape and chloroplast distribution. Palisade cells with more equal dimensions have more chloroplasts on their abaxial surfaces. This dense layer of chloroplasts maximizes the light capture efficiency limited by sieve effects. The more columnar palisade cells of sun-adapted taxa allow light to pass through the central vacuoles and spaces between cells, making chloroplasts less efficient in energy capture, but allowing light to reach chloroplasts in the spongy mesophyll. Pioneer species may be an exception to these two groups of species. Three pioneer taxa included in this study have columnar palisade cells that are extremely narrow and packed closely together. This layer allows little penetration of light, but exposure of the leaf undersurface may provide illumination of spongy mesophyll chloroplasts in these plants.     

Anatomical features of pepper plants (Capsicum annuum L.) grown under red light-emitting diodes supplemented with blue or far-red light

Pepper plants (Capsicum annuum L. cv., Hungarian Wax) were grown under metal halide (MH) lamps or light-emitting diode (LED) arrays with different spectra to determine the effects of light quality on plant anatomy of leaves and stems. One LED (660) array supplied 90% red light at 660 nm (25nm band-width at half-peak height) and 1% far-red light between 700-800nm. A second LED (660/735) array supplied 83% red light at 660nm and 17% far-red light at 735nm (25nm band-width at half-peak height). A third LED (660/blue) array supplied 98% red light at 660nm, 1% blue light between 350-550nm, and 1% far-red light between 700-800nm. Control plants were grown under broad spectrum metal halide lamps. Plants were gron at a mean photon flux (300-800nm) of 330 micromol m-2 s-1 under a 12 h day-night photoperiod. Significant anatomical changes in stem and leaf morphologies were observed in plants grown under the LED arrays compared to plants grown under the broad-spectrum MH lamp. Cross-sectional areas of pepper stems, thickness of secondary xylem, numbers of intraxylary phloem bundles in the periphery of stem pith tissues, leaf thickness, numbers of choloplasts per palisade mesophyll cell, and thickness of palisade and spongy mesophyll tissues were greatest in peppers grown under MH lamps, intermediate in plants grown under the 660/blue LED array, and lowest in peppers grown under the 660 or 660/735 LED arrays. Most anatomical features of pepper stems and leaves were similar among plants grown under 660 or 660/735 LED arrays. The effects of spectral quality on anatomical changes in stem and leaf tissues of peppers generally correlate to the amount of blue light present in the primary light source.     

Comparative ultrastructural properties of pallisade tissue and spongy tissue chloroplasts from normal and mutant leaves of Pisum sativum L.*

Abstract. The ultrastructure of chloroplasts from palisade and spongy tissue was studied in order to analyse the adaptation of chloroplasts to the light gradient within the bifacial leaves of pea. Chloroplasts of two nuclear gene mutants of Pisum sativum (chlorotica-29 and chlorophyll b-less 130A), grown under normal light conditions, were compared with the wild type (WT) garden-pea cv. ‘Dippes Gelbe Viktoria’. The differentiation of the thylakoid membrane system of plastids from normal pea leaves exhibited nearly the same degree of grana formation in palisade and in spongy tissue. Using morphometrical measurements, only a slight increase in grana stacking capacity was found in chloroplasts of spongy tissue. In contrast, chloroplasts of mutant leaves differed in grana development in palisade and spongy tissue, respectively. Their thylakoid systems appeared to be disorganized and not developed as much as in chloroplasts from normal pea leaves. Grana contained fewer lamellae per granum, the number of grana per chloroplast section was reduced and the length of appressed thylakoid regions was decreased. Nevertheless, chloroplasts of the mutants were always differentiated into grana and stroma thylakoids. The structural changes observed and the reduction of the total chlorophyll content correlated with alterations in the polypeptide composition of thylakoid membrane preparations from mutant chloroplasts. In sodium dodecyl sulphate polyacrylamide gel electrophoresis (SDS-PAGE), polypeptide bands with a relative molecular mass of 27 and 26 kilodalton (kD) were markedly reduced in mutant chloroplasts. These two polypeptides represented the major apoproteins of the light harvesting chlorophyll a/b complex from photosystem II (LHC-II) as inferred from a comparison with the electrophoretic mobility of polypeptides isolated from the LHC-II.     

Changes in chloroplast morphology of different parenchyma cells in leaves of Haberlea rhodopensis Friv. during desiccation and following rehydration

The size, shape, and number of chloroplasts in the palisade and spongy parenchyma layers of Haberlea rhodopensis leaves changed significantly during desiccation and following rehydration. The chloroplasts became smaller and more rounded during desiccation, and aggregated in the middle of the cell. The size and number of chloroplasts in the palisade parenchyma cells were higher than in spongy parenchyma. The good correlation observed between the size or number of chloroplasts and the cross-sectional area of mesophyll cells, the cross-sectional width of the leaf and its water content suggested that the palisade cells were more responsive to water availability than the spongy cells. Changes in chloroplast number during desiccation and rehydration process are characteristic features for desiccation-tolerant plants (especially in homoiochlorophyllous strategy).     

The effect of light quality on leaf production and development of in vitro-cultured plants of Alternanthera brasiliana Kuntze

We investigated the influence of light quality on the leaf development of Alternanthera brasiliana Kuntze (Amaranthaceae) grown in vitro. Growth parameters including specific leaf mass, thickness, and leaf density were lowest in plants grown under red light. Blue light induced the largest number of leaves/plant, and the largest thickness and area of the leafblade. Green and red lights induced the smallest leaf areas. The thickness of the abaxial-face epidermis and spongy parenchyma of the plants was significantly reduced in plants grown under red light. The thickness of the palisade parenchyma and upper epidermis were significantly increased in plants grown under blue light, compared to the other fluorescent-light treatments. The specific spectral band also influenced the differentiation of mesophyll cells. In the dark and under red light, the mesophyll was homogenous; and in the dark and under green light, the leaves were more compact. Under blue light, the cells displayed the characteristic palisade morphology. The results showed that the increase of a specific parenchyma type was related to a specific spectral band. All spectral-quality treatments reduced the numbers of stomata and trichomes. The results for green light were in some respects similar to those for red light, and in other respects similar to those for blue light, probably because phytochromes and cryptochromes are green-light receptors. This study indicated that Alternanthera plants have strong morphological plasticity induced by light. The results suggest that high-quality Alternanthera can be achieved by culturing the plants in vitro under a combination of blue and red light.     

光质对番茄和莴苣幼苗生长及叶绿体超微结构的影响

采用发光二极管(LED)精确调制不同光谱能量分布,以荧光灯光照为对照,研究光质对番茄和莴苣幼苗生长及叶绿体超微结构的影响.结果表明：红光下番茄、莴苣幼苗的可溶性糖、淀粉和碳水化合物含量均显著高于对照,叶片叶绿体中淀粉粒膨大显著;蓝光极显著抑制了番茄下胚轴伸长,显著提高了莴苣和番茄幼苗叶片叶绿素a和类胡萝卜素含量;红蓝光下莴苣幼苗叶片的可溶性糖、淀粉、碳水化合物、蔗糖含量和C/N均达到最大值且显著高于红光处理,番茄和莴苣幼苗的主根显著伸长,幼苗叶片中叶绿体形态正常,基粒增多,基质片层清晰,淀粉粒体积明显小于红光处理.光质对植物幼苗的光形态建成、生长、碳氮代谢及叶绿体发育有显著影响;红光下光合产物积累显著但运输受阻严重,在红光中添加适量蓝光更有利于莴苣幼苗的碳水化合物积累,并可促进幼苗根系生长,有利于同化产物输出.     

Measurement of chlorophyll fluorescence within leaves using a fibreoptic microprobe

Abstract. The distribution of chlorophyll fluorescence was measured within leaves of Medicago saliva with a fibre optic microprobe. Leaves were irradiated with broad band blue light (1000 μmol m⁻²s⁻¹) and chlorophyll fluorescence was measured at 688 nm. The amount of fluorescence measured within the leaf depended upon the direction in which the probe was inserted. When the probe was advanced directly through the leaf from the shaded towards the irradiated surface, the maximum amount of detected fluorescence occurred near the boundary between the palisade and spongy mesophyll. When the probe was advanced through the leaf from the opposite direction maximum detected fluorescence was at the boundary between the epidermis and palisade. These results appear to be a consequence of the blue light gradient, which declined exponentially within the palisade but was counterbalanced by increasing chlorophyll content within the leaf. Modelling indicates that the measured distribution of chlorophyll fluorescence can be explained by relatively uniform emission of fluorescence throughout the palisade layer, indicating that the chloroplasts may be photosynthetically specialized to their light environment within the leaf.     

Chloroplasts do not have a polarity for light-induced accumulation movement

Chloroplast photorelocation movement in green plants is generally mediated by blue light. However, in cryptogam plants, including ferns, mosses, and algae, both red light and blue light are effective. Although the photoreceptors required for this phenomenon have been identified, the mechanisms underlying this movement response are not yet known. In order to analyze this response in more detail, chloroplast movement was induced in dark-adapted Adiantum capillus-veneris gametophyte cells by partial cell irradiation with a microbeam of red and/or blue light. In each case, chloroplasts were found to move toward the microbeam-irradiated area. A second microbeam was also applied to the cell at a separate location before the chloroplasts had reached the destination of the first microbeam. Under these conditions, chloroplasts were found to change their direction of movement without turning and move toward the second microbeam-irradiated area after a lag time of a few minutes. These findings indicate that chloroplasts can move in any direction and do not exhibit a polarity for chloroplast accumulation movement. This phenomenon was analyzed in detail in Adiantum and subsequently confirmed in Arabidopsis thaliana palisade cells. Interestingly, the lag time for direction change toward the second microbeam in Adiantum was longer in the red light than in the blue light. However, the reason for this discrepancy is not yet understood. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The DCL gene of tomato is required for chloroplast development and palisade cell morphogenesis in leaves.

The defective chloroplasts and leaves-mutable (dcl-m) mutation of tomato was identified in a Ds mutagenesis screen. This unstable mutation affects both chloroplast development and palisade cell morphogenesis in leaves. Mutant plants are clonally variegated as a result of somatic excision of Ds and have albino leaves with green sectors. Leaf midribs and stems are light green with sectors of dark green tissue but fruit and petals are wild-type in appearance. Within dark green sectors of dcl-m leaves, palisade cells are normal, whereas in albino areas of dcl-m leaves, palisade cells do not expand to become their characteristic columnar shape. The development of chloroplasts from proplastids in albino areas is apparently blocked at an early stage. DCL was cloned using Ds as a tag and encodes a novel protein of approximately 25 kDa, containing a chloroplast transit peptide and an acidic alpha-helical region. DCL protein was imported into chloroplasts in vitro and processed to a mature form. Because of the ubiquitous expression of DCL and the proplastid-like appearance of dcl-affected plastids, the DCL protein may regulate a basic and universal function of the plastid. The novel dcl-m phenotype suggests that chloroplast development is required for correct palisade cell morphogenesis during leaf development.     

Light environment within a leaf. II. Progress in the past one-third century

Studies directly related to light environments within a leaf, conduced mainly in the past one-third century, are reviewed. In particular, studies that revealed the profiles of light absorption and photosynthetic capacity are highlighted. Progress in this research field has been accelerated by devising innovative techniques. Roles of the main photosynthetic tissues, the palisade and spongy tissues, as the light guide and diffuser, respectively, are discussed. When the leaf is illuminated with diffuse light, light is absorbed more by the chloroplasts located near the illuminated surface. The meanings of the occupation of the mesophyll surfaces facing the intercellular spaces by chloroplasts and chloroplast movement are also discussed. The discrepancy between the light absorption profile and that of photosynthetic capacity is examined most intensively.     

Epidermal focussing and effects upon photosynthetic light-harvesting in leaves of Oxalis

Abstract. In Oxalis , epidermal cells on both the adaxial and abaxial surface of the leaf concentrated light within the leaf by a lens mechanism. Focal lengths of epidermal cells were estimated using two methods: they were calculated from radius of curvature measurements taken from individual epidermal cells, and were measured directly in agarose replicas of the leaf surface. In the three species of Oxalis examined, light that was incident upon the adaxial leaf surface was concentrated within the palisade, whereas light that was incident upon the abaxial leaf surface was concentrated within the spongy mesophyll. Using sensiometric analysis, theoretically maximal focal intesifications were measured in leaf replicas at the focal maximum and at intermediate positions corresponding to the mid-region of the palisade and spongy mesophyll tissues. Focal intensifications ranged from 2.2 to 10.4 times incident light at the focal maximum, and 1.3 to 4.5 in the palisade or spongy mesophyll layers. Elimination of epidermal focussing, by covering the leaf surface with a thin layer of mineral oil, strongly affected chlorophyll fluorescence induction curves resulting in a decrease of 10–40% in the initial (F₀) and variable fluorescence (F_v). These results are consistent with the interpretation that the chloroplasts were adapted to their light microenvironment within the leaf and that focussing by the epidermis channelled light to a population of chloroplasts that were adapted to high light.     

Structural and functional properties of the coleoptile chloroplast: Photosynthesis and photosensory transduction

Recent studies have shown that guard cell and coleoptile chloroplasts appear to be involved in blue light photoreception during blue light-dependent stomatal opening and phototropic bending. The guard cell chloroplast has been studied in detail but the coleoptile chloroplast is poorly understood. The present study was aimed at the characterization of the corn coleoptile chloroplast, and its comparison with mesophyll and guard cell chloroplasts. Coleoptile chloroplasts operated the xanthophyll cycle, and their zeaxanthin content tracked incident rates of solar radiation throughout the day. Zeaxanthin formation was very sensitive to low incident fluence rates, and saturated at around 800–1000 mol m^–2 s^–1. Zeaxanthin formation in corn mesophyll chloroplasts was insensitive to low fluence rates and saturated at around 1800 mol m^–2 s^–1. Quenching rates of chlorophyll a fluorescence transients from coleoptile chloroplasts induced by saturating fluence rates of actinic red light increased as a function of zeaxanthin content. This implies that zeaxanthin plays a photoprotective role in the coleoptile chloroplast. Addition of low fluence rates of blue light to saturating red light also increased quenching rates in a zeaxanthin-dependent fashion. This blue light response of the coleoptile chloroplast is analogous to that of the guard cell chloroplast, and implicates these organelles in the sensory transduction of blue light. On a chlorophyll basis, coleoptile chloroplasts had high rates of photosynthetic oxygen evolution and low rates of photosynthetic carbon fixation, as compared with mesophyll chloroplasts. In contrast with the uniform chloroplast distribution in the leaf, coleoptile chloroplasts were predominately found in the outer cell layers of the coleoptile cortex, and had large starch grains and a moderate amount of stacked grana and stroma lamellae. Several key properties of the coleoptile chloroplast were different from those of mesophyll chloroplasts and resembled those of guard cell chloroplasts. We propose that the common properties of guard cell and coleoptile chloroplasts define a functional pattern characteristic of chloroplasts specialized in photosensory transduction.Abbreviations Ant or A antheraxanthin - dv/dt fluorescence quenching rate - Fm maximum yield of fluorescence with all PS II reaction centers closed - Fo yield of instantaneous fluorescence with all PS II reaction centers open - Vio or V violaxanthin - Zea or Z zeaxanthin     

葡萄属植物叶片显微和超微结构观察

利用光学显微镜和透射电镜,对葡萄属(Vitis L.)15种、1亚种和4栽培品种植物的叶片显微和超微结构进行了观察研究。结果显示:(1)东亚种群中桦叶葡萄的叶片总厚度和各组织厚度最大;北美种群的河岸葡萄和沙地葡葡叶片总厚度、上表皮厚度、栅栏组织厚度、海绵组织厚度和下表皮厚度较大,但叶片组织紧密度最小。(2)各种类间栅栏组织中叶绿体数目均多于海绵组织,且刺葡萄、桦叶葡萄、葛藟葡萄和秋葡萄叶肉细胞中叶绿体数目最多,北红、蘡薁、网脉葡萄和菱叶葡萄叶片中叶绿体数目最少,巨峰和桦叶葡萄叶绿体中含有大量的淀粉粒,桦叶葡萄、网脉葡萄、秋葡萄、葛藟葡萄、刺葡萄和华东葡萄含有较多的嗜锇颗粒。研究表明,中国野生葡萄的许多种类在叶片组织紧密度、叶绿体数量、淀粉粒数量、嗜锇颗粒数量方面均高于北美种群的河岸葡萄和沙地葡萄,具有更高的结构抗性基础。     

Measurement of gradients of absorbed light in spinach leaves from chlorophyll fluorescence profiles

Profiles of chlorophyll fluorescence were measured in spinach leaves irradiated with monochromatic light. The characteristics of the profiles within the mesophyll were determined by the optical properties of the leaf tissue and the spectral quality of the actinic light. When leaves were infiltrated with 10^?4M DCMU [3‐(3,4‐dichlorophenyl)‐1, 1‐dimethyl‐urea] or water, treatments that minimized light scattering, irradiation with 2000 μmol m^?2 s^?1 green light produced broad Gaussian‐shaped fluorescence profiles that spanned most of the mesophyll. Profiles for chlorophyll fluorescence in the red (680 ± 16 nm) and far red (λ > 710 nm) were similar except that there was elevated red fluorescence near the adaxial leaf surface relative to far red fluorescence. Fluorescence profiles were narrower in non‐infiltrated leaf samples where light scattering increased the light gradient. The fluorescence profile was broader when the leaf was irradiated on its adaxial versus abaxial surface due to the contrasting optical properties of the palisade and spongy mesophyll. Irradiation with blue, red and green monochromatic light produced profiles that peaked 50, 100 and 150 μm, respectively, beneath the irradiated surface. These results are consistent with previous measurements of the light gradient in spinach and they agree qualitatively with measurements of carbon fixation under monochromatic blue, red and green light. These results suggest that chlorophyll fluorescence profiles may be used to estimate the distribution of quanta that are absorbed within the leaf for photosynthesis.     

The functional significance of palisade tissue: penetration of directional versus diffuse light

Light gradients were measured in leaves that had different types of anatomical development of the mesophyll but similar pigment content. Leaves of the legume, Thermopsis montana, had columnar palisade and spongy mesophyll whereas leaves of the monocot, Smilacina stellata, had spongy mesophyll only. Light gradients were measured at 550 nm in both types of leaves when they were irradiated with collimated or diffuse light. When irradiated with collimated light, light gradients were steeper in leaves with spongy mesophyll in comparison to those that had palisade tissue. On the other hand, light gradients were similar between both leaf types when they were irradiated with diffuse light. Thus, columnar palisade cells facilitated the penetration of collimated light over diffuse light. These results suggest that palisade tissue may help distribute light more uniformly to chloroplasts within the leaf. Moreover, the functional significance of palisade tissue may be related to the amount of collimated light within the natural environment.