Comprehensive insights into the influence of supplemental green light on the photosynthesis of ginger (Zingiber officinale Roscoe)

Although green light is not considered to contribute to the photosynthesis of plants, the photosynthesis of ginger, a dual-purpose vegetable used as a medicine and food, is affected by the green wave band. In this study, the supplementary green band of sunlight (SG) increased the net photosynthetic rate (Pn), maximal photochemical efficiency of PSII (Fv/Fm), and actual photochemical efficiency of PSII (Y(II)) compared with the sunlight treatment (S). The Pn and Fv/Fm of the SG treatment were higher than those of the white light (W) treatment, while the Pn and Fv/Fm of the green light (G) treatment alone were lower than those of the W treatment. Further analysis found that the minimal fluorescence (Fo) of the S treatment increased, especially at noon, while the Fo of the SG treatment decreased. Similarly, the Fo of the W treatment increased significantly, while the Fo of the white–green mixed light (WG) treatment decreased. The relative fluorescence values of the K-J-I bands in the SG and WG treatments were lower than those in the S and W treatments, respectively. The photochemical quenching (qP) of the WG treatment was higher than that of the W treatment, while the primary thermal losses corresponded to the sum of nonregulated heat dissipation and fluorescence emission (Y(NO)) of the WG treatment was lower than that of the W treatment. The SG treatment reduced the accumulation of plastoglobules but increased the accumulation of starch granules and leaf thickness. Moreover, the green band supplemented with white light significantly increased the biomass of the aboveground plant parts and promoted the active growth of the aboveground parts. Supplementing green light plays a regulatory role in ginger based on the following four points. First, it effectively promotes the transfer of electrons between the acceptor side of photosystem II; second, it optimizes ginger photosynthesis; third, it alleviates strong light stress by reducing the accumulation of reactive oxygen species; and fourth, it promotes heat dissipation and reduces the rapid burst of active oxygen in the chloroplast caused by excess energy. In summary, green light can significantly optimize the photosynthetic characteristics of ginger.

     

Effects of light quality on the chloroplastic ultrastructure and photosynthetic characteristics of cucumber seedlings

To understand how light quality influences plant photosynthesis, we investigated chloroplastic ultrastructure, chlorophyll fluorescence and photosynthetic parameters, Rubisco and chlorophyll content and photosynthesis-related genes expression in cucumber seedlings exposed to different light qualities: white, red, blue, yellow and green lights with the same photosynthetic photon flux density of 100 μmol m^?2 s^?1. The results revealed that plant growth, CO₂ assimilation rate and chlorophyll content were significantly reduced in the seedlings grown under red, blue, yellow and green lights as compared with those grown under white light, but each monochromatic light played its special role in regulating plant morphogenesis and photosynthesis. Seedling leaves were thickened and slightly curled; Rubisco biosynthesis, expression of the rca, rbcS and rbcL, the maximal photochemical efficiency of PSII (Fv/Fm) and quantum yield of PSII electron transport (Ф_PSII) were all increased in seedlings grown under blue light as compared with those grown under white light. Furthermore, the photosynthetic rate of seedlings grown under blue light was significantly increased, and leaf number and chlorophyll content of seedlings grown under red light were increased as compared with those exposed to other monochromatic lights. On the contrary, the seedlings grown under yellow and green lights were dwarf with the new leaves etiolated. Moreover, photosynthesis, Rubisco biosynthesis and relative gene expression were greatly decreased in seedlings grown under yellow and green light, but chloroplast structural features were less influenced. Interestingly, the Fv/Fm, Ф_PSII value and chlorophyll content of the seedlings grown under green light were much higher than those grown under yellow light.     

Photosynthetic utilization of radiant energy by CAM Dendrobium flowers

14CO2 fixation was observed in orchid Dendrobium flowers; its rate decreased with the flower development. Chlorophyll (Chl) fluorescence in different developmental stages of flowers was compared to other green plant parts (leaf, inflorescence stalk, and fruit capsule). The photochemical efficiency of photosystem 2 (PS2) (Fv/Fm) of a leaf was 14-21 % higher than that of a mature flower perianth (sepal, petal, and labellum) which had a much lower total Chl content and Chl a/b ratio. A higher quantum yield of PS2 (ΦPS2) than in the mature flowers was observed in all green parts. Flower sepals had higher Chl content, Chl a/b ratio, and Fv/Fm values than the petal and labellum. During flower development the Chl content, Chl a/b ratio, Fv/Fm, and qN decreased while ΦPS2 and qP remained constant. An exposure of developing flowers to irradiances above 50 µmol m-2 s-1 resulted in a very drastic drop of ΦPS2 and qP, and a coherent increase of qN as compared to other green plant organs. A low saturation irradiance (PFD of 100 µmol m-2 s-1) and the increase in qN in the flower indicate that irradiation stress may occur since there is no further protection when the flower is exposed to irradiances above 100 µmol m-2 s-1. A low Chl/carotenoid ratio in mature flower perianth as a consequence of Chl content reduction in the course of flower development suggests a relief of irradiation stress via this mean.     

Effect of ambient ozone at the somma of Lake Mashu on growth and leaf gas exchange in Betula ermanii and Betula platyphylla var. japonica

We examined the effects of ambient ozone, at the somma of Lake Mashu in northern Japan, on the growth and photosynthetic traits of two common birch species in Japan (mountain birch and white birch). Seedlings of the two birch species were grown in open-top chambers and were exposed to charcoal-filtered ambient air (CF) or non-filtered ambient air (NF) at the somma of Lake Mashu during the growing season in 2009. For the mountain birch, ambient ozone significantly increased the ratio of aboveground dry mass to belowground dry mass (T/R ratio), although no difference in the whole-plant biomass was observed between the treatments. For the white birch, in contrast, ozone exposure at ambient level did not decrease in growth and photosynthesis. These results suggest that ambient O₃ at the somma of Lake Mashu may shift the allocation of biomass to above-ground rather than below-ground in the mountain birch.     

Nitric oxide plays a role as second messenger in the ultraviolet-B irradiated green alga <Emphasis Type="Italic">Chlorella pyrenoidosa</Emphasis>

Nitric oxide (NO) stimulated the activity of plasma membrane H⁺-ATPase, 5′-nucleotidase, peroxidase, ascorbate peroxidase and glutathione reductase in ultraviolet B (UV-B) irradiated Chlorella pyrenoidosa. It also boosted the activity of nitrogen-metabolism enzymes such as nitrate reductase, nitrite reductase, glutamine synthetase, which were inhibited by UV-B irradiation. The chlorophyll fluorescence ratio (Fv/Fm) of the UV-B irradiated algae and decreased continuously after the cells were transferred to UV-B irradiation. A continuing decrease of the Fv/Fm was observed even after the cells were transferred to photosynthetically active radiation (PAR). After adaptation for 8 h under PAR (after treatment with nitric oxide), Fv/Fm recovered to 55 % of normal levels — without NO the value approached zero. Exogenous NO stopped the decay of chlorophyll and thylakoid membrane in cells exposed to UV-B irradiation. NO plays probably a key role in damage induced by UV-B irradiation in green algae.     

红光下培养方式对雨生红球藻细胞增殖及其活力的影响

研究重点针对雨生红球藻绿色游动细胞的增殖培养阶段,分析了在利于细胞增殖的红光条件下,几种培养方式的调整对增殖过程和细胞活力的影响。结果显示:（1）在红光下,增殖平台期维持时间长,细胞活力稳定,细胞中性脂无累积,但进入平台期前,细胞中性脂有规律波动,进入平台期后相对稳定;通过更新率为20%的半连续培养,细胞数产出较批次培养提高57%;半连续培养中细胞呈现胁迫调节的时间较批次培养晚。随着培养时间增加,半连续培养下细胞营养盐吸收能力降低。（2）初始接种密度与细胞增殖速率及细胞光合活力呈负相关:初始密度低的细胞增殖速率较高,细胞光合作用活力高。（3）在培养过程中添加CO2时,最大密度均有提高,达6.0105 cells/mL,较无添加组提高54%;细胞分裂速率均有提高,但红光下较白光下增殖速率高（分别为0.223/d和0.198/d）;添加CO2降低培养液pH,利于维持适宜增殖的pH环境。叶绿素荧光参数以及细胞粒径在红光和白光下有显著差异:红光下,Fv/Fm显著高于白光下;红光下补充CO2显著减小细胞粒径,而白光下粒径无显著变化。研究结果显示,在红光下,采用间断式半连续培养补充CO2培养绿色游动细胞,有利于提升细胞活力与产出。     

The rapid yellowing of spruce at a mountain site in the Central Black Forest (Germany). Combined effects of Mg deficiency and ozone on biochemical, physiological and structural properties of the chloroplasts

Biochemical, physiological and ultrastructural changes of the chloroplasts were examined in the course of the rapid yellowing process of spruce (Picea abies (L.) Karst.) at a Mg-deficient and ozone polluted mountain site (Sch?llkopf mountain, Central Black Forest, Germany, 840 m a.s.l.). While at an early stage of yellowing the chlorophyll (Chl) content of the needles decreased slowly, significant changes occurred in the chloroplasts: The lability of the light-harvesting Chl a/b protein complex LHC II increased; the thylakoid cross-sectional area of chloroplasts in the outer mesophyll of the needles decreased, and their Chl fluorescence showed typical changes like the decrease of Fv/Fm and the increase of the photoinhibitory Fv quenching. Later on, the Chl content decreased rapidly, the changes in the chloroplasts continued and the needles turned yellow. Lutein and the pigments of the xanthophyll cycle were enhanced in relation to Chl a. Light and dark reactions of the xanthophyll cycle were highly active indicating efficient proton pumping and NADPH formation. The ratio of nonappressed to appressed thylakoid membranes increased with decreasing Fv/Fm suggesting that structural and fluorescence properties of the chloroplasts were related. The response of the needles to defined shading and improved Mg supply was also examined. The combined effects of strong sun light, low levels of non-Chl-bound Mg (Mg(free)) and ozone concentrations exceeding 80 microg m(-3) are shown to be necessary to induce the rapid yellowing process. For needles with Mg(free) < 0.12 mg g(-1) needle dry matter, the lability of the LHC II was correlated with the ozone concentration suggesting that the destabilization of the LHC II plays a central role in the rapid yellowing process.     

建兰叶艺品种光合色素含量及叶绿素荧光特性分析

以建兰栽培品种‘八宝奇珍’Cymbidium ensifolium ‘Ba Bao Qi Zhen’为材料,对其正常绿色叶片及黄化变异叶片的光合色素含量、叶绿素荧光参数进行比较。结果表明,‘八宝奇珍’黄化变异叶片的叶绿素总量、叶绿素a 和叶绿素b 含量均显著低于正常绿色叶片,且随着黄化面积的增大呈现递减趋势;黄化变异叶片的初始荧光量(Fo)、最大荧光产量(Fm)、Kautsky 诱导效应最大荧光(Fp)、稳态光适应光化学淬灭系数(qP)以及光适应稳态荧光产量(Ft_Lss)均显著低于正常绿色叶片;PSⅡ原初光能转化效率(Fv/Fm)与正常绿色叶片无明显差异;稳态非光化学荧光淬灭系数(NPQ)高于正常绿色叶片。     

玉米全生育期黄化材料的光合性能分析

通过对玉米(Zea mays L.)黄化材料和2份正常自交系的光合特性、光响应曲线及荧光动力学参数进行分析,结果表明,黄化材料yglm1的净光合速率(Pn)、气孔导度(Gs)和胞间CO2浓度(Ci)均显著低于正常自交系Mo17和RP128,而蒸腾速率(Tr)却显著高于正常自交系;黄化材料的光补偿点、光饱和点、暗呼吸速率及表观量子效率均高于正常自交系Mo17和RP128,但其净光合速率却低于正常自交系;黄化材料的荧光动力学参数初始荧光(Fo)、最大荧光(Fm)、PSⅡ原初光能转换效率(Fv/Fm)、PSⅡ原初光能捕获效率(Fv'/Fm')、PSⅡ光下实际光化学效率(ΦPSⅡ)、光化学猝灭系数(qP)均显著低于正常自交系Mo17和RP128,非光化学猝灭系数(qN)显著高于正常自交系。相对于正常自交系,黄化材料具有较低的电子传递潜力、原初光能捕获效率和转换效率、光能分配及利用率,这可能是导致其净光合能力降低的根本原因。     

ACCLIMATION OF THE PHOTOSYNTHETIC APPARATUS OF PALMARIA PALMATA (RHODOPHYTA) TO LIGHT QUALITIES THAT PREFERENTIALLY EXCITE PHOTOSYSTEM I OR PHOTOSYSTEM II1

Acclimation of the photosynthetic apparatus to light absorbed primarily by phycobilisomes (which transfer energy predominantly to photosystem II) or absorbed by chlorophyll a (mainly present in the antenna of photosystem I) was studied in the macroalga Palmaria palmata L. In addition, the influence of blue and yellow light, exciting chlorophyll a and phycobilisomes, respectively, ivas investigated. All results were compared to a white light control. Complementary chromatic adaptation in terms of an enhanced ratio of phycoerythrin to phycocyanin under green light conditions was observed. Red light (mainly absorbed by chlorophyll a) and green light (mainly absorbed by phycobilisomes) caused an increase of the antenna system, which was not preferentially excited. Yellow and blue light led to intermediate states comparable to each other and white light. Growth was reduced under all light qualities in comparison to white light, especially under conditions preferably exciting phycobilisomes (green light-adapted algae had a 58% lower growth rate compared to white light-adapted algae). Red and blue light-adapted algae showed maximal photosynthetic capacity with white light excitation and significantly lower values with green light excitation. In contrast, green and yellow light-adapted algae exhibited comparable photosynthetic capacities at all excitation wavelengths. Low-temperature fluorescence emission analysis showed an increase of photosystem II emission in red light-adapted algae and a decrease in green light-adapted algae. A small increase of photosystem I emission teas also found in green light-adapted algae, but this was much less than the photosystem II emission increase observed in red light-adapted algae (both compared to phycobilisome emission). Efficiency of energy transfer from phycobilisomes to photosystem II was higher in red than in green light-adapted algae. The opposite was found for the energy transfer efficiency from phycobilisomes to photosystem I. Zeaxanthin content increased in green and blue light-adapted algae compared to red, white, and yellow light-adapted algae. Results are discussed in comparison to published data on unicellular red algae and cyanobacteria.     

Effects of ozone exposure or fungal pathogen on white lupin leaves as determined by imaging of chlorophyll a fluorescence.

Chlorophyll fluorescence has been used routinely to investigate photosynthetic activity in plants subjected to both biotic and abiotic stresses. The aim of this work was to compare the perturbations in photosynthesis induced by ozone and by a pathogen. By using a conventional fluorometer a similar response pattern was observed in inoculated and O(3)-fumigated leaves. The application of chlorophyll fluorescence imaging provided further detailed information on the spatial-temporal heterogeneity of the response of white lupin leaves to fungal pathogen or to ozone fumigation. In particular, 48 h after artificial inoculation with the necrotrophic fungal pathogen Pleiochaeta setosa, the leaves showed a remarkable alteration in PSII operating efficiency (Phi(PSII)), which affected the whole surface. Afterwards, the infection site was surrounded by a ring of increased photosynthetic activity. The response of ozonated leaves was quite different. The reduction in Phi(PSII) was already evident 24h after fumigation; moreover, a distinct heterogeneity of the fluorescence yield was observed and the major veins displayed a lowered Phi(PSII).     

Photosynthetic utilization of radiant energy by CAM Dendrobium flowers

14CO2 fixation was observed in orchid Dendrobium flowers; its rate decreased with the flower development. Chlorophyll (Chl) fluorescence in different developmental stages of flowers was compared to other green plant parts (leaf, inflorescence stalk, and fruit capsule). The photochemical efficiency of photosystem 2 (PS2) (Fv/Fm) of a leaf was 14-21 % higher than that of a mature flower perianth (sepal, petal, and labellum) which had a much lower total Chl content and Chl a/b ratio. A higher quantum yield of PS2 (PS2) than in the mature flowers was observed in all green parts. Flower sepals had higher Chl content, Chl a/b ratio, and Fv/Fm values than the petal and labellum. During flower development the Chl content, Chl a/b ratio, Fv/Fm, and qN decreased while PS2 and qP remained constant. An exposure of developing flowers to irradiances above 50 µmol m-2 s-1 resulted in a very drastic drop of PS2 and qP, and a coherent increase of qN as compared to other green plant organs. A low saturation irradiance (PFD of 100 µmol m-2 s-1) and the increase in qN in the flower indicate that irradiation stress may occur since there is no further protection when the flower is exposed to irradiances above 100 µmol m-2 s-1. A low Chl/carotenoid ratio in mature flower perianth as a consequence of Chl content reduction in the course of flower development suggests a relief of irradiation stress via this mean.     

Structural changes in the vascular bundles of light-exposed and shaded spruce needles suffering from Mg deficiency and ozone pollution

The correlation between structural changes of the vascular bundles and needle yellowing was examined for needles of damaged spruce (Picea abies (L.) Karst.) growing at a Mg-deficient and ozone polluted mountain site in the Central Black Forest (840m a.s.l.). In the previous year's sun-exposed needles, the following sequence of events was observed: (1) rapid needle yellowing, (2) hypertrophy and anomalous divisions of cambium cells, (3) phloem collapse, and, (4) production of atypical xylem tracheids. Under defined shade (reduction of the photosynthetically active photon flux density of the ambient light by 85-90%), the needles remained green, while the phloem collapsed completely within the first 6 weeks of shading; subsequently, a reversal of the collapse was observed. Under both light conditions, the content of Mg not bound to chlorophyll (Mg(free)) was in the range of 0.1 mg g(-1) needle dry matter, and hardly changed throughout the investigation period. After Mg fertilization, the Mg(free) level of the previous year's needles increased to 0.2 mg g(-1) dry matter, the light-exposed needles remained green, and the vascular bundles developed no anomalies. The data show that the rapid needle yellowing of ozone-exposed Mg-deficient needles did not depend on the collapse of the phloem. Mg deficiency played a key role in the development of anomalous vascular bundles under light, and also appears to explain the transient changes in sieve cell structure under shade. The role of Mg deficiency, rather than ozone pollution, in the damage of the sieve cells was confirmed in a long-term ozone exposure experiment with young clonal spruce growing under defined conditions.     

不同光质对烟草叶片生长及光合作用的影响

通过对烟草植株覆盖白、红、黄、蓝、紫色滤膜获得不同光质,研究了光质对烟叶生长及光合作用的影响。结果表明,与白膜处理相比,红膜与蓝膜处理下的烟草叶片较厚,比叶面积较小,叶绿素a/b比值、净光合速率、可变荧光强度（Fv）和最大荧光强度（Fm）的比值Fv/Fm（PSⅡ最大光化学量子效率）、PSⅡ实际光化学量子效率（ΦPSⅡ）、光饱和点和CO2饱和点均较高。黄膜处理下的叶片较白膜处理的更薄,净光合速率、Fv/Fm、ΦPSⅡ、光饱和点、CO2饱和点均较低。紫膜处理的叶片比叶面积比白膜处理的小,净光合速率和Fv/Fm比白膜的大。实验结果表明红光、蓝光和紫光促进了烟叶的生长,这种促进作用是与其高光合效率紧密相连的;而黄光对烟叶的生长有一定程度的抑制作用。     

Life-cycles of lotic populations of Spongilla lacustris and Eunapius fragilis (Porifera: Spongillidae)

SUMMARY. The life-cycles of green and white morphs of the freshwater sponge Spongilla lacustris were examined in the light of past evidence that zoochlorellae may augment their sponge host's nutrition. Field collections from a lotic population of S. lacustris were supplemented by laboratory experiments on gemmule hatching and gemmule size. Both white and green S. lacustris produced sperm for a 6-week period in 1976 starting in the middle of May. Out of thirty white and thirty green sponges examined during this period, twenty white and ten green sponges contained sperm. Sperm production in both morphs was limited primarily to the basal 3.18mm of sponge tissue, and the density of sperm packets in the two morphs was the same. Out of 180 white and green sponges examined in 1976, only four eggs, no embryos, and no larvae were observed. White sponges gemmulated a week or two earlier, and produced smaller gemmules which were more uniform in size than those of green sponges. White and green gemmules hatched synchronously in the spring. In 1977 one female and numerous male specimens of S. lacustris , and numerous females but no males of another sponge, Eunapius fragilis , were found. The life-cycles are discussed in the light of other recent studies on freshwater sponges.     

The effect of ozone on the yellowing process of magnesium-deficient clonal Norway spruce grown under defined conditions

During two vegetation periods, young clonal spruce trees (Picea abies (L.) Karst.) with sufficient and poor magnesium (Mg) supply were exposed in the environmental chambers of the GSF phytotron to three levels of ozone (daily means: 18-22, 88-130, and 135-190 microg m(-3); 10% reduction at night). Previous year's needles were examined at 4-week intervals with respect to their contents of Mg, Ca, K, Mn, N, P, and chlorophyll (Chl), various parameters of Chl fluorescence, and the stability of the isolated light-harvesting Chl-a/b-protein complex LHC II. The needles of the two nutrition variants contained more than 0.53 or less than 0.27mg Mg g(-1) needle dry matter, respectively. The ratio of variable to maximal Chl-a fluorescence of the dark-adapted needles, Fv/Fm, and the photoinhibitory quenching of Fv after light treatment, SVi.v, were affected by the Mg content of the needles rather than the ozone levels. Changes of the Chl content and the behavior of the LHC II allowed differentiating between a slow process of needle yellowing occurring under Mg deficiency only, and a rapid process of needle yellowing occurring under the combined action of Mg deficiency and ozone pollution. Only the rapid yellowing process was accompanied by destabilization of the LHC II, and the degree of destabilization was correlated with the ozone concentration present in the days before sampling. The results are consistent with observations obtained at a research site in the Central Black Forest (J Plant Physiol 161 (2004) 423).     

Examination of chlorophyll fluorescence decay kinetics in sulfur deprived algae Chlamydomonas reinhardtii

Chlorophyll fluorescence decay kinetics was measured in sulfur deprived cells of green alga Chlamydomonas reinhardtii with a home made picosecond fluorescence laser spectrometer. The measurements were carried out on samples either shortly adapted to the dark (‘Fo conditions’) or treated to reduce Qa (‘Fm conditions’). Bi-exponential fitting of decay kinetics was applied to distinguish two components one of them related to energy trapping (fast component) and the other to charge stabilization and recombination in PS 2 reaction centers (slow component). It was found that the slow component yield increased by 2.0 and 1.2 times when measured under ‘Fo’ and ‘Fm conditions’, respectively, in sulfur deprived cells as compared to control ones. An additional rapid rise of the slow component yield was observed when incubation was carried out in a sealed bioreactor and cell culture turned to anaerobic conditions. The obtained results strongly indicate the existence of the redox control of PS 2 activity during multiphase adaptation of C. reinhardtii to sulfur deficiency stress. Probable mechanisms responsible for the observed increased recombinant fluorescence yield in starved cells are discussed.     

Response to ozone in two lettuce varieties on chlorophyll a fluorescence, photosynthetic pigments and lipid peroxidation.

The effect of different O3 concentrations on two lettuce (Lactuca sativa L.) varieties (Valladolid and Morella) was investigated through chlorophyll (Chl) a fluorescence parameters, photosynthetic pigments (Chl a, b and total carotenoid), lipid peroxidation and crop yield. Ozone fumigation caused: a decrease in maximum quantum yield of photosystem II (PSII) photochemistry (Fv/Fm) in mature leaves, a reduction in the non-cyclic electron flow (phiPSII) and a lower capacity to reoxidize the QA pool (qP). These reductions were significant in the Valladolid var. but not in the Morella var. A significant decrease in Chl a, b and in the total carotenoids was observed in the Valladolid var. but not in the Morella var. mainly under O3 fumigation conditions. We observed that the NPQ parameter did not increase in parallel to the qP reduction seen in the Valladolid var. O3 fumigation with respect to air charcoal filtered air conditions. This fact could be associated with a lower capacity for dissipation of non-radiative excess energy and it may be closely correlated with significant decreases in photosynthetic pigment concentration. A decrease in NPQ from air ozone-free to ozone fumigation in the Morella var. can be explained by the need to maintain the photochemical quenching under O3 stress. It may also be associated with a slight increase in photosynthetic pigments. The differences between the two varieties may indicate that the Valladolid var. is more susceptible to O3 damage.     

Effect of Green and White Flashes on the Activity of the Enzyme Ribulose-diphonphate Carboxylase in Darkgrown Barley Plants

The effect of short impulses of white and green light on the activity of ribulosediphosphate carboxylase in dark-grown barley leaves has been studied Intermittent white light induces an increase in the activity of the enzyme. After 90 flashes a 30% increase in the activity is observed. Intermittent green light has no effect on the activity of the enzyme. A correlation between the increase in the activity of the enzyme and the accumulation of chlorophyll under the effect of intermittent white light has been observed.