Model parameterization to simulate and compare the PAR absorption potential of two competing plant species

Mountain pastures dominated by the pasture grass Setaria sphacelata in the Andes of southern Ecuador are heavily infested by southern bracken (Pteridium arachnoideum), a major problem for pasture management. Field observations suggest that bracken might outcompete the grass due to its competitive strength with regard to the absorption of photosynthetically active radiation (PAR). To understand the PAR absorption potential of both species, the aims of the current paper are to (1) parameterize a radiation scheme of a two-big-leaf model by deriving structural (LAI, leaf angle parameter) and optical (leaf albedo, transmittance) plant traits for average individuals from field surveys, (2) to initialize the properly parameterized radiation scheme with realistic global irradiation conditions of the Rio San Francisco Valley in the Andes of southern Ecuador, and (3) to compare the PAR absorption capabilities of both species under typical local weather conditions. Field data show that bracken reveals a slightly higher average leaf area index (LAI) and more horizontally oriented leaves in comparison to Setaria. Spectrometer measurements reveal that bracken and Setaria are characterized by a similar average leaf absorptance. Simulations with the average diurnal course of incoming solar radiation (1998–2005) and the mean leaf–sun geometry reveal that PAR absorption is fairly equal for both species. However, the comparison of typical clear and overcast days show that two parameters, (1) the relation of incoming diffuse and direct irradiance, and (2) the leaf–sun geometry play a major role for PAR absorption in the two-big-leaf approach: Under cloudy sky conditions (mainly diffuse irradiance), PAR absorption is slightly higher for Setaria while under clear sky conditions (mainly direct irradiance), the average bracken individual is characterized by a higher PAR absorption potential. (∼74 MJ m⁻² year⁻¹). The latter situation which occurs if the maximum daily irradiance exceeds 615 W m⁻² is mainly due to the nearly orthogonal incidence of the direct solar beam onto the horizontally oriented frond area which implies a high amount of direct PAR absorption during the noon maximum of direct irradiance. Such situations of solar irradiance favoring a higher PAR absorptance of bracken occur in ∼36% of the observation period (1998–2005). By considering the annual course of PAR irradiance in the San Francisco Valley, the clear advantage of bracken on clear days (36% of all days) is completely compensated by the slight but more frequent advantage of Setaria under overcast conditions (64% of all days). This means that neither bracken nor Setaria show a distinct advantage in PAR absorption capability under the current climatic conditions of the study area.     

Why is chlorophyll <Emphasis Type="Italic">b</Emphasis> only used in light-harvesting systems?

Chlorophylls (Chl) are important pigments in plants that are used to absorb photons and release electrons. There are several types of Chls but terrestrial plants only possess two of these: Chls a and b. The two pigments form light-harvesting Chl a/b-binding protein complexes (LHC), which absorb most of the light. The peak wavelengths of the absorption spectra of Chls a and b differ by c. 20 nm, and the ratio between them (the a/b ratio) is an important determinant of the light absorption efficiency of photosynthesis (i.e., the antenna size). Here, we investigated why Chl b is used in LHCs rather than other light-absorbing pigments that can be used for photosynthesis by considering the solar radiation spectrum under field conditions. We found that direct and diffuse solar radiation (PAR_dir and PAR_diff, respectively) have different spectral distributions, showing maximum spectral photon flux densities (SPFD) at c. 680 and 460 nm, respectively, during the daytime. The spectral absorbance spectra of Chls a and b functioned complementary to each other, and the absorbance peaks of Chl b were nested within those of Chl a. The absorption peak in the short wavelength region of Chl b in the proteinaceous environment occurred at c. 460 nm, making it suitable for absorbing the PAR_diff, but not suitable for avoiding the high spectral irradiance (SIR) waveband of PAR_dir. In contrast, Chl a effectively avoided the high SPFD and/or high SIR waveband. The absorption spectra of photosynthetic complexes were negatively correlated with SPFD spectra, but LHCs with low a/b ratios were more positively correlated with SIR spectra. These findings indicate that the spectra of the photosynthetic pigments and constructed photosystems and antenna proteins significantly align with the terrestrial solar spectra to allow the safe and efficient use of solar radiation.     

The effect of solar UV-B radiation on terpenes and biomass production in Grindelia chiloensis (Asteraceae), a woody perennial of Patagonia,Argentina

UV-B radiation is absorbed effectively by nucleic acids and other sensitive targets, potentially causing harmful photochemical effects. Protection against UV-B radiation may be afforded by flavonoids and other phenolics, which absorb strongly in the UV region, but little is known about the role played by other compounds, such as terpenes. Grindelia chiloensis, native of Patagonia (Argentina), can accumulate as much as 25% resin (terpenes) in its leaves. The present investigation was carried out to test the effect of solar UV-B radiation on the allocation of photoassimilates to biomass and terpenes. Exposure to UV-B radiation reduced whole plant biomass, plant height and leaf area, and increased leaf thickness and resin accumulation in Grindelia chiloensis. Higher absorbance was found for refined resin in the UV-B waveband from plants grown under solar UV-B radiation than plants without UV-B radiation. These chemical and structural changes could protect the plant from UV radiation, and may help elucidate the importance of epicuticular resins for a species as G. chiloensis native to an environment with maximum daily integrated values of solar UV-B irradiance.     

光诱导下杂种杨无性系叶角和叶绿体的运动

自然光结合人工光源对几个杂种杨无性系叶片进行处理后,对其在两种光环境下３种叶角（方位角、方向角、悬挂角）和叶绿体的运动方式进行了研究。结果发现：各无性系的叶角运动在晴天比比阴天强烈,特别是三倍体无性系最为明显,晴天上午９：００－１１：００,三倍体无性系ＺＨ６和Ｂ３４６通过方位角运动来避免强光胁迫,各无性系主要通过方向角和悬挂角的变化来调节获得最佳太阳辐射,在中午受到强光胁迫时存在明显的“避光运动”。三倍体无性系和某些二倍体无性系在避光运动方式上是不同的,三倍体无性系ＺＨ６和Ｂ３４６采用叶片下垂形式,而二倍体无性系Ｂ１１则采用叶片竖立方式,无论是晴天还是阴天,植物的节律性运动可能参与了叶角运动。受光胁迫时,栅栏组织的叶绿通过不同的运动排列方式来实现对光辐射的最佳吸收。强光胁迫下叶绿体沿径向细胞壁排列,以尽量养活接受过量的太阳辐射,处于弱光条件时,叶绿体则充满足整个细胞,以扩大上太阳辐射的表面积,三倍体无性系对光诱导的敏感程度程度要高于二倍体无性系。     

Changes in leaf optical properties associated with light-dependent chloroplast movements

We surveyed 24 plant species to examine how leaf anatomy influenced chloroplast movement and how the optical properties of leaves change with chloroplast position. All species examined exhibited light-dependent chloroplast movements but the associated changes in leaf absorptance varied considerably in magnitude. Chloroplast movement-dependent changes in leaf absorptance were greatest in shade species, in which absorptance changes of >10% were observed between high- and low-light treatments. Using the Kubelka-Munk theory, we found that changes in the absorption (k) and chlorophyll a absorption efficiency (k*) associated with chloroplast movement correlated with cell diameter, such that the narrower, more columnar cells found in sun leaves restricted the ability of chloroplasts to move. The broader, more spherical cells of shade leaves allowed greater chloroplast rearrangements and in low-light conditions allowed efficient light capture. Across the species tested, light-dependent chloroplast movements modulated leaf optical properties and light absorption efficiency by manipulating the package (sieve or flattening) effect but not the detour (path lengthening) effect.     

Modification of leaf cytology and anatomy in Brassica napus grown under above ambient levels of supplemental UV-B radiation.

Plants exposed to natural solar radiation usually show acclimation responses on a daily and seasonal basis. Many of these responses are complex and modified by interactions with acclimation responses to other climatic factors. While changes in photosynthetically active radiation (PAR, 400-700 nm) are the driving force for many acclimation responses in plants, radiation outside the PAR range is also important. Recently, interest has increased in the potential role of UV-A (320-400 nm) and UV-B (280-320 nm) components of sunlight in plant developmental, physiological and daily acclimation processes. In order to explore the role of UV-B further, Brassica napus L. cv Paroll plants were grown to maturity under 13 kJ d(-1) of biologically effective ultraviolet-B radiation (UV-B(BE), 280-320 nm) plus 800 micromol photons m(-2) s(-1) photosynthetically active radiation (PAR, 400-700 nm) or PAR alone. Leaf anatomy and palisade cell structure were quantified using stereological techniques. The leaves of plants grown under UV-B radiation exhibited an increase in overall leaf width, although no change in leaf anatomy was discerned. Palisade cells in UV-B exposed leaves showed a significant decrease in chloroplast, mitochondrial, starch, and microbody volume density (Vv), while the vacuolar Vv increased compared to cells exposed to PAR only. In UV-B exposed leaves, there was an increase in the appressed and non-appressed thylakoid surface area density (Sv) within the chloroplasts. Since the relative proportion of appressed to non-appressed thylakoid surface area did not change, both thylakoid systems changed in concert with each other. Thylakoid stacks were broader and shorter in leaves subjected to UV-B. In general these responses were similar to those which occurred in plants moved from a high to low PAR environment and similar to mature plants exposed to 13 kJ d(-1) UV-B(BE) for only a short period of time. Although UV absorbing pigments increased by 21% in UV-B exposed leaves, there was no significant difference in chlorophyll a,b or carotenoid content compared to plants exposed to only PAR.     

Light absorption by phytoplankton and chromophoric dissolved organic matter in the drainage basin and estuary of the Neuse River, North Carolina (U.S.A.)

1. We examined the absorption of solar radiation by phytoplankton and chromophoric dissolved organic matter (CDOM) taking into account riparian shading in the rivers, reservoirs, swamps of the Neuse River Estuary and its drainage basin. 2. In the streams, CDOM typically absorbed 55 and 64% of photons in the spectral range of 400–700 nm (photosynthetically active radiation, PAR) and 500–600 nm, respectively. The large proportion of photons absorbed by CDOM indicates high potential for abiotic photochemial reactions in the 500–600 nm region. 3. Despite the high concentration of nutrients, phytoplankton contributed little (2%) to the total absorption of PAR in the streams. Small (<30 m wide) streams typically received only 7% of incident PAR that impinged onto the more exposed reservoirs and estuary. Riparian shading and the low contribution of phytoplankton to the total absorption resulted in conditions where phytoplankton absorbed nearly two orders of magnitude less PAR in the streams than in the estuary and reservoirs. 4. The results indicated that riparian shading and non‐algal absorbing components can significantly restrict phytoplankton production in nutrient‐rich streams with a high concentration of CDOM flowing throughout forested catchments.     

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.     

The combined effects of CO2 concentration and enhanced UV-B radiation on faba bean. 3. Leaf optical properties,pigments, stomatal index and epidermal cell density

Seedlings of Vicia faba L. (cv. Minica) were grown in a factorial experiment in a greenhouse. The purpose of the study was to determine whether CO₂ enrichment and supplemental UV-B radiation affect leaf optical properties and whether the combined effects differ from single factor effects. Seedlings were grown at either 380 mol mol^-1 or 750 mol mol^-1 CO₂ and at four levels of UV-B radiation. After 20 and 40 days of treatment, absorptance, transmittance and reflectance of photosynthetically active radiation (PAR) were measured on the youngest fully developed leaf. On the same leaf, the specific leaf area on a fresh weight basis (SLA_fw), chlorophyll content, UV-B absorbance, transmittance of UV light and stomatal index were measured. UV-B radiation significantly increased PAR absorptance and decreased PAR transmittance. The increased PAR absorptance can be explained by an increased chlorophyll content in response to UV-B radiation. Leaf transmittance of UV radiation decreased with increasing UV-B levels mainly caused by increased absorbance of UV absorbing compounds. UV-B radiation decreased both the stomatal density and epidermal cell density of the abaxial leaf surface, leaving the stomatal index unchanged. Effects of CO₂ enrichment were less pronounced than those of UV-B radiation. The most important CO₂ effect was an increase in stomatal density and epidermal cell density of the adaxial leaf surface. The stomatal index was not affected. No interaction between CO₂ and UV-B radiation was found. The results are discussed in relation to the internal light environment of the leaf.     

Physiological role of anthocyanin accumulation in common hazel juvenile leaves

Common hazel (Corylus avellana L., Fusca rubra Dipp.) juvenile leaves from the periphery of the canopy and thus subjected to high fluxes of solar radiation are characterized by red coloration due to anthocyanin accumulation disappearing in mature leaves. To elucidate the physiological role of anthocyanin accumulation, the interrelations between anthocyanin content, a degree of attenuation by the pigments of the light reaching the photosynthetic apparatus (PSA), and PSA tolerance to photoinhibition in C. avellana juvenile leaves were studied. Absorption spectra were calculated taking into account the light losses due to reflection by the leaf. The analysis of the spectra showed that, in red common hazel leaves accumulating high amounts of anthocyanins in the vacuoles of the upper and lower epidermal cells, up to 95% of visible radiation entering the leaf blade was absorbed by these pigments. The rate of the linear electron transport (ETR) in the chloroplast electron transport chain (ETC) was closely correlated with the anthocyanin content (r ² = 0.87). In red leaves, the saturation of ETR dependence on irradiance was observed at the higher values of PAR than in green leaves. In red juvenile leaves, this value was close to that in mature green leaves tolerant to high light. There were no differences between red and green leaves in the level of non-photochemical quenching, the content of violaxanthin cycle pigments, a degree of their de-epoxidation under natural illumination and at irradiation with high PAR fluxes. Basing on the data obtained, one may conclude that anthocyanins in C. avellana juvenile leaves serve PSA photoprotection, preventing injury of immature PSA with excessive fluxes of PAR.     

Pubescence and leaf spectral characteristics in a desert shrub,Encelia farinosa

Summary The effects of leaf hairs (pubescence) on leaf spectral characteristics were measured for the drought-deciduous desert shrub Encelia farinosa. Leaf absorptance to solar radiation is diminished by the presence of pubescence. The pubescence appears to be reflective only after the hairs have dried out. There are seasonal changes in leaf absorptance; leaves produced at the beginning of a growing season have high absorptances, whereas leaves produced during the growing season are more pubescent and have lower absorptances. The decrease in leaf absorptance is the result of an increase in pubescence density and thickness. Between 400 and 700 nm (visible wavelengths), pubescence serves as a blanket reflector. However, over the entire solar spectrum (400–3000 nm), the pubescence preferentially reflects near infrared radiation (700–3000 nm) over photosynthetically useful solar radiation (400–700 nm). Leaf absorptance to solar radiation (400–3000 nm) varies between 46 and 16%, depending on pubescence; whereas leaf absorptance to photosynthetically useful radiation (400–700 nm) may vary from 81 to 29%.C.I.W.-D.P.B. Publication No. 612     

MICROALGAL LIGHT-HARVESTING IN EXTREME LOW-LIGHT ENVIRONMENTS IN MCMURDO SOUND, ANTARCTICA1

Microalgal pigment composition, photosynthetic characteristics, single-cell absorption efficiency (Qa_(λ)) spectra, and fluorescence-excitation (FE) spectra were determined for platelet ice and benthic communities underlying fast ice in Mc Murdo Sound, Antarctica, during austral spring 1988. Measurements of spectral irradiance (E_(λ)) and photosynthetically active radiation (PAR) as well as samples for particulate absorption measurements were taken directly under the congelation ice, within the platelet layer, as profiles vertically through the water column, and at the benihic surface. Light attenuation by.sea ice, algal pigments, and particulates reduced PAR reaching the platelet ice layer to 3%(9–33 fimol photons m-²-^?s-¹) of surface values and narrowed its spectral distribution to a band between 400 and 580 nm. Attenuation by the water column further reduced PAR reaching the sea floor (28–m depth) to 0.05% of surface levels (< 1 μmol photons m-² s-¹), with a spectral distribution dominated by 470–580–nm wavelengths. The photoadaptive index (I) for platelet ice algae (5.9–12.6 μmol photons m-^2.s-¹) was similar to ambient PAR, indicating that algae had acclimated to their light environment (i.e. the algae were light-replete). Maximum Qa_(λ) at the blue absorption peak (440 nm) was 0.63, and enhanced absorption was observed from 460–500 nm and was consistent with observed high cellular chlorophyll (chi) c:chl a and fucoxanthin: chl a molar ratios (0.4 and 1.2, respectively). Benthic algae were light-limited despite the maintenance of very low I_k values (4–11 μmol photons.m-^2.s-¹). Extremely high fucoxanthin: chi a ratios (1.6) in benthic algae produced enhanced green light absorption, resulting in a high degree of complementation between algal absorption and ambient spectral irradiance. Qa_(λ) values for benthic algae were maximal (0.9) between 400 and 510 nm but remained >0.35 even at absorption minima. Strong spectral flattening, a characteristic of intense pigment packaging, was also apparent in the Qa_(λ) spectra for benthic algae. FE and Qa_(λ) spectra were similar in shape for platelet ice algae, indicating that the efficiency at which absorbed energy was transferred to photosystem II (PSII) was independent of wavelength. Fluorescence emission by benthic algae was greatest for the 500–560–nm excitation wavelengths, suggesting that most energy absorbed by accessory pigments was transferred to PSII. These results suggest that under ice algae employ complementary pigmentation and maximize absorption efficiency as adaptive strategies to low-light stress. Regulating the distribution of absorbed energy between PSI and PSII may be an adaptive response to the restricted spectral distribution of irradiance.     

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.     

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.     

Moderate heat stress of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> leaves causes chloroplast swelling and plastoglobule formation

Photosynthesis is inhibited by heat stress. This inhibition is rapidly reversible when heat stress is moderate but irreversible at higher temperature. Absorbance changes can be used to detect a variety of biophysical parameters in intact leaves. We found that moderate heat stress caused a large reduction of the apparent absorbance of green light in light-adapted, intact Arabidopsis thaliana leaves. Three mechanisms that can affect green light absorbance of leaves, namely, zeaxanthin accumulation (absorbance peak at 505 nm), the electrochromic shift (ECS) of carotenoid absorption spectra (peak at 518 nm), and light scattering (peak at 535 nm) were investigated. The change of green light absorbance caused by heat treatment was not caused by changes of zeaxanthin content nor by the ECS. The formation of non-photochemical quenching (NPQ), chloroplast movements, and chloroplast swelling and shrinkage can all affect light scattering inside leaves. The formation of NPQ under high temperature was not well correlated with the heat-induced absorbance change, and light microscopy revealed no appreciable changes of chloroplast location because of heat treatment. Transmission electron microscopy results showed swollen chloroplasts and increased number of plastoglobules in heat-treated leaves, indicating that the structural changes of chloroplasts and thylakoids are significant results of moderate heat stress and may explain the reduced apparent absorbance of green light under moderately high temperature.     

Leaf anatomy during leaf development of photoautotrophically <Emphasis Type="Italic">in vitro</Emphasis>-grown tobacco plants as affected by growth irradiance

Tobacco (Nicotiana tabacum L.) plants were cultured in vitro photoautotrophically at three levels of irradiance (PAR 400–700 nm): low (LI, 60 μmol m⁻² s⁻¹), middle (MI, 180 μmol m⁻² s⁻¹) and high (HI, 270 μmol m⁻² s⁻¹). Anatomy of the fourth leaf from bottom was followed during leaf development. In HI and MI plants, leaf area expansion started earlier as compared to LI plants, and both HI and MI plants developed some adaptations of sun species: leaves were thicker with higher proportion of palisade parenchyma to spongy parenchyma tissue. Furthermore, in HI and MI plants palisade and spongy parenchyma cells were larger and relative abundance of chloroplasts in parenchyma cells measured as chloroplasts cross-sectional area in the cell was lower than in LI plants. During leaf growth, chloroplasts crosssectional area in both palisade and spongy parenchyma cells in all treatments considerably decreased and finally it occupied only about 5 to 8 % of the cell cross-sectional area. Thus, leaf anatomy of photoautotrophically in vitro cultured plants showed a similar response to growth irradiance as in vivo grown plants, however, the formation of chloroplasts and therefore of photosynthetic apparatus was strongly impaired.     

Epidermal focussing and effects on light utilization in Oxalis acetosella

lmage analysis was used to quantify the lateral heterogeneity of the radiation field within the palisade of Oxialis acetosella L. leaves. Oxalis acetosella epidermal cells focus light up to four times incident irradiance, resulting in regions of high and low internal fluenec rate within the palisade. Chlorophyll fluorescence from leaves irradiated with directional light was found to originate primarily from palisade cell chloroplasts located within focal zones. When the internal radiation field was made more homogeneous by using diffuse light or by coating the leaf with a layer of mineral oil to eliminate epidermal focussing, the characteristics of the chlorophyll fluorescence signal were altered: non-photochemical quenching (qN) increased, while relaxation of qN was slowed. This indicates that upper palisade chloroplasts may fine-tune their light utilization to the intra leaf light microenvironment.     

Light absorption by isolated chloroplasts and leaves: effects of scattering and ‘packing’

Light absorption was quantified in the following systems: isolated chloroplasts and leaves of spinach (Spinacea oleracea L.), a mutant of geranium (Pelargonium zonale L.) widely differing in pigment content, and coleus (Coleus blumei Benth.) at different stages of leaf ontogenesis. For these species and pea (Pisum sativum L.), scattering-compensated absorption spectra of chloroplast suspensions are presented. Comparison of leaf and chloroplast spectra showed considerable changes in the extent of the ‘package’ effect and the lengthening of the effective optical path in a leaf. The difference between leaf and isolated chloroplast absorption could be quantitatively described by adapting Duysens’s treatment of flattening. It was found that the accumulation of chlorophyll in leaves is accompanied by a monotonous enhancement of the package effect. The results are discussed with special reference to the role of light scattering in leaf optics, light utilization in photosynthesis and wavelength-dependent light gradients in a leaf.     

Spectral composition of photosynthetically active radiation penetrating into a Norway spruce canopy: the opposite dynamics of the blue/red spectral ratio during clear and overcast days

The spectral composition of photosynthetically active radiation (PAR) during clear and overcast days was studied above the canopy (U) and at two layers of a dense Norway spruce stand [Picea abies (L.) Karst.] characterized with an average LAI = 7.3 ± 0.8 (middle layer: M) and 12.3 ± 0.7 (lower layer: L). Whereas the spectral composition of PAR incoming on the canopy surface during cloudy days (characterized by diffuse index DI > 0.7) was almost independent of the solar elevation angle, the proportion of the blue-green spectral region of PAR was significantly reduced at low elevation angles during days with prevailing direct radiation (DI < 0.3). The PAR spectrum at both M and L levels was only slightly enriched in the green spectral region (more pronounced for DI < 0.3). The penetration of diffuse radiation into the canopy resulted in a slight (approx. 5%) reduction of the blue region proportion that remained stable during the day. On the contrary, under clear sky conditions the penetration of blue and red radiation was dependent on the solar elevation in an opposite manner in comparison with the spectral composition of PAR incident on canopy, giving almost twofold proportion of the blue part of the spectrum at a low elevation angle at M layer. We suggest that the blue enhancement of the spectrum within the Norway spruce canopy during clear days is due to a specific spatial arrangement of the assimilatory apparatus of a coniferous stand. Further, the possible consequences of the observed dynamics of the PAR spectrum inside the canopy during clear days on the efficiency of PAR absorption of the needles located within the canopy are discussed.     

Living off the Sun: chlorophylls,bacteriochlorophylls and rhodopsins

Pigments absorbing 350–1,050 nm radiation have had an important role on the Earth for at least 3.5 billion years. The ion pumping rhodopsins absorb blue and green photons using retinal and pump ions across cell membranes. Bacteriochlorophylls (BChl), absorbing in the violet/blue and near infra red (NIR), power anoxygenic photosynthesis, with one photoreaction centre; and chlorophylls (Chl), absorbing in the violet/blue and red (occasionally NIR) power oxygenic photosynthesis, with two photoreaction centres. The accessory (bacterio)chlorophylls add to the spectral range (bandwidth) of photon absorption, e.g., in algae living at depth in clear oceanic water and in algae and photosynthetic (PS) bacteria in microbial mats. Organism size, via the package effect, determines the photon absorption benefit of the costs of synthesis of the pigment–protein complexes. There are unresolved issues as to the evolution of Chls vs. BChls and the role of violet/blue and NIR radiation in PS bacteria.