Chlorophyll and light gradients in sun and shade leaves of Spinacia oleracea

Abstract. Light gradients were measured and correlated with chlorophyll concentration and anatomy of leaves in spinach (Spinacia oleracea L.). Light gradients were measured at 450, 550 and 680 nm within thin (455 μm) and thick (630 μm) leaves of spinach grown under sun and shade conditions. The light gradients were relatively steep in both types of leaves and 90% of the light at 450 and 680 nm was absorbed by the initial 140 μm of the palisade. In general, blue light was depleted faster than red light which, in turn was depleted faster than green light. Light penetrated further into the thicker palisade of sun leaves in comparison to the shade leaves. The distance that blue light at 450 nm travelled before it became 90% depleted was 120 μm in sun leaves versus 76 μm in shade leaves. Red light at 680 nm and green light at 550 nm travelled further but the trends were similar to that measured at 450nm. The steeper light gradients within the palisade-of shade leaves were caused by increased scattering of light within the intercellular air spaces and/or cells which were less compact than those in sun leaves. The decline in the amount of light within the leaf appeared to be balanced by a gradient in chlorophyll concentration measured in paradermal sections. Progressing from the adaxial epidermis, chlorophyll content increased through the palisade and then declined through the spongy mesophyll. Chlorophyll content was similar in the palisade of both sun and shade leaves. Chloroplast distribution within both sun and shade leaves was relatively uniform so that the chlorophyll gradient appeared to be caused by greater amounts of chlorophyll within chloroplasts located deeper within the leaf. These results indicate that the anatomy of the palisade may be of special importance for controlling the penetration of photo-synthetically active radiation into the leaf. Changing the structural characteristics of individual palisade cells or their arrangement may be an adaptation that maximizes the absorption of light in leaves with varying mesophyll thickness due to different ambient light regimes.     

Evidence for the role of the oxygen-evolving manganese complex in photoinhibition of Photosystem II

Photoinhibition of PSII occurs at the same quantum efficiency from very low to very high light, which raises a question about how important is the rate of photosynthetic electron transfer in photoinhibition. We modulated electron transfer rate and light intensity independently of each other in lincomycin-treated pea leaves and in isolated thylakoids, in order to elucidate the specific effects of light and PSII electron transport on photoinhibition. Major changes in the rate of electron transport caused only small changes in the rate of photoinhibition, suggesting the existence of a significant photoinhibitory pathway that contains an electron-transfer-independent phase. We compared the action spectrum of photoinhibition with absorption spectra of PSII components that could function as photoreceptors of the electron-transfer-independent phase of photoinhibition and found that the absorption spectra of Mn(III) and Mn(IV) compounds resemble the action spectrum of photoinhibition, showing a steep decrease from UV-C to blue light and a low visible-light tail. Our results show that the release of a Mn ion to the thylakoid lumen is the earliest detectable step of both UV- and visible-light-induced photoinhibition. After Mn release from the oxygen-evolving complex, oxidative damage to the PSII reaction center occurs because the Mn-depleted oxygen-evolving complex cannot reduce P680+ normally.     

Blue,Green and Red Fluorescence Signatures and Images of Tobacco Leaves*

Laser-induced fluorescence images of the leaf of an aurea mutant of Nicotiana tabacum were recorded for the blue and green fluorescence at 440 and 520 nm and the red chlorophyll fluorescence at 690 and 735 nm. The results obtained were compared with direct measurements of the fluorescence emission spectra of leaves using a conventional spectrofluorometer. The highest emission of blue (F440) and green fluorescence (F520) within the leaf was found in the leaf veins, particularly the main leaf vein. In contrast, the intercostal fields of leaves, which exhibited the highest chlorophyll content, showed only a very low blue and green fluorescence emission, which was much lower than the red and far-red chlorophyll fluorescence emission bands (F690 and F735). Correspondingly, the ratio of blue to red leaf fluorescence F440/F690 of upper and lower leaf side was much higher in the leaf veins (values 1.2 to 1.5) than in intercostal fields (values of 0.6 to 0.7). The results also demonstrated that in the intercostal fields the major part of the blue-green fluorescence was reabsorbed by chlorophylls and carotenoids. A partial reabsorption of the red fluorescence band near 690 nm by leaf chlorophyll took place, but did not affect the far-red fluorescence band near F735. As a consequence the chlorophyll fluorescence ratio F690/F735 exhibited significantly higher values in the chlorophyll-poor leaf vein regions (1.7 to 1.8) than in the chlorophyll-rich intercostal fields (0.8 to 1.3). Imaging spectroscopy of leaves was shown to be much more precise than the screening of fluorescence signatures by conventional fluorometers. It clearly demonstrated that the blue-green fluorescence and the red chlorophyll fluorescence of leaves exhibit an inverse contrast to each other. The advantage of the fluorescence imaging spectroscopy, which allows the simultaneous screening of the whole leaf surface and distinct parts of it, and its possible application in the detection of stress effects or local damage by insects and pathogens, is discussed.     

Characterization of the reversible state of photoinhibition occurring in vitro under anaerobic conditions

Thylakoid membranes were subjected to photoinhibitory illumination. The use of oxygen-consuming enzymes to obtain strictly anaerobic conditions showed that while the absence of oxygen is a prerequisite for the formation of a reversibly photoinhibited state, the presence of oxygen is required for the recovery in the dark. The formation of the reversibly photoinhibited state did not protect the thylakoid membranes against irreversible damage. The effects of both bicarbonate and formate were found to be qualitatively different for photoinhibition under strictly anaerobic conditions compared to the effects observed under normal aerobic photoinhibition. It is suggested that there are two different mechanisms of photoinhibition, occurring to different extent under aerobic and anaerobic conditions, respectively, involving Q_A in both cases, but the Q_B-site in the former only.Abbreviations chl chlorophyll - PpBQ phenyl-p-benzoquinone - PS 2 Photosystem 2 - Q_A and Q_B primary and secondary quinone acceptors of Photosystem 2     

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.     

环境强光诱导玉簪叶片光抑制的机制

为进一步阐述光抑制的强光诱导和发生机制, 该文以喜阴植物玉簪(Hosta spp.)为材料研究其光抑制发生规律及其与环境光强的关系。结果表明, 全日照和遮阴条件下玉簪叶片发育分别形成适应强光和弱光的形态特征; 与遮阴处理相比, 强光下生长的玉簪光合速率和叶绿素含量较低, 但两种处理叶片最大光化学效率差异很小, 证明强光下植株可以正常生长且光合机构未发生严重的光抑制。将遮阴处生长的植株转移到全日照下, 光合速率和最大光化学效率急剧下降; 荧光诱导动力学曲线发生明显改变, 而且光系统II供体侧和受体侧荧光产量的变化幅度分别达到24.3%和34.2%, 表明玉簪由弱光转入强光后光系统II发生不可逆失活, 且受体侧受到的伤害较供体侧更严重。因此, 作者认为环境光强骤然提高并超过玉簪生长光强时很容易诱导其光合机构发生严重的光抑制。该研究对于理解植物适应光环境的策略以及喜阴植物的优质栽培有重要意义。     

Photoinhibition of manganese enzymes: insights into the mechanism of photosystem II photoinhibition

Evidence has recently been presented that photoinhibition of photosystem II (PSII) is triggered by absorption of light by the oxygen-evolving manganese cluster. To get insight into the effects of light on enzymes containing manganese or other transition metal cofactors, the photosensitivities of Mn catalase, Mn superoxide dismutase, the haem (Fe)-containing bovine liver catalase, and CuZn superoxide dismutase were investigated. Glucose oxidase was studied as an example of an enzyme that does not have a metal cofactor. Sensitivities of these five enzymes to UVC, UVA, and visible light were compared in anaerobic conditions. The Mn(III)-oxo-Mn(III)-containing Mn catalase was found to be more sensitive to both visible and UV light than bovine liver catalase. Furthermore, the action spectrum of photoinhibition of Mn catalase was found to be fairly similar to that of photoinhibition of PSII. The Mn(II)-containing Mn superoxide dismutase was sensitive to UVC light and somewhat sensitive to UVA light, while only UVC light caused some inhibition of CuZn superoxide dismutase. Glucose oxidase was the least photosensitive of the enzymes studied. The photosensitivity of Mn enzymes supports the hypothesis that the oxygen-evolving manganese complex of PSII can be damaged by UV and visible light absorbed by its Mn(III) or Mn(IV) ions.     

Mechanisms of photoinhibition induced by high light in Hosta grown outdoors

Aims It has long been recognized that photoinhibition of photosynthesis is induced by high light. However, our recent studies are not consistent with this traditional view. Therefore, the objective of this study is to explore the induction of photoinhibition and its mechanisms under full sunlight outdoors.
Methods Changes of leaf morphology, gas exchange, and chlorophyll a fluorescence were measured to investigate the induction and mechanisms of photoinhibition under high light in Hosta, which is a typical shade-tolerant plant.
Important findings Hosta plants grown under full sunlight (HT) and low light (LT) developed sun- and shade-type leaf morphological characteristics, respectively. Under a full sunlight, Hosta plants had lower photosynthetic rate and chlorophyll content than under the LT; whereas, there were only slight difference in the maximum quantum yield of photosystem II (F_v/F_m) between the two treatments, suggesting that Hosta plants could grow normally under full sunlight without severe photoinhibition. After transition from the low to a high light (LHT), the photosynthetic rate and maximum quantum yield of photosystem II decreased sharply, reflecting that the LHT treatment led to irreversibly inactivation of photosystem II. Additionally, the shape of chlorophyll a fluorescence transients also changed significantly; the relative fluorescence yield of the K and J steps were reduced by 24.3% and 34.2%, respectively, indicating that the acceptor side of photosystem II was damaged more severely than the donor side. Consequently, we postulate that photoinhibition in Hosta leaves is mainly induced by the sudden enhancement of light intensity outdoors. Hosta can acclimate to high irradiance through leaf development outdoors. Our finding is of great significance in understanding the acclimation of plants to high light and cultivation of shade-tolerant plants in field.     

苗期遮光光质对生姜光合及生长的影响

以不同颜色塑料薄膜为遮光材料,研究了苗期遮光光质对生姜生长及光合作用的影响.结果表明:幼苗覆膜期,生姜叶片叶绿素含量以蓝膜及绿膜处理较高,白膜次之,红膜较低;叶片P_n则以绿膜处理较高,为14.9 μmol·m^-2·s^-1（第4叶）,分别较白膜、红膜及蓝膜提高5.7%、10.4%和18.3%.旺盛生长期撤膜后,P_n较幼苗期升高,但处理间的变化趋势与幼苗期相似;新生叶片叶绿素含量除红膜处理较低外,其它处理无显著差异,但下位叶片叶绿素含量则以蓝膜和红膜处理显著低于绿膜和白膜处理.蓝膜处理生姜植株茎秆增高、变细,分枝数较少;绿膜处理植株根、茎、叶及根茎鲜质量较高,白膜、红膜及蓝膜处理依次降低,收获时,其产量分别达57 000、53 709、51 487和48 712 kg·hm^-2.说明生姜苗期采用绿膜遮光,可增强叶片光合作用,促进植株生长,提高生姜产量.     

Mechanistic differences in photoinhibition of sun and shade plants

Leaf discs of the shade plant Tradescantia albiflora Kunth grown at 50 μmol · m^?2 · s^?1, and the facultative sun/shade plant Pisum sativum L. grown at 50 or 300 μmol · m^?2, s^?1, were photoinhibited for 4 h in 1700 μmol photons m^?2 · s^?1 at 22° C. The effects of photoinhibition on the following parameters were studied: i) photosystem II (PSII) function; ii) amount of D1 protein in the PSII reaction centre; iii) dependence of photoinhibition and its recovery on chloroplast-encoded protein synthesis; and, iv) the sensitivity of photosynthesis to photoinhibition in the presence or absence of the carotenoid zeaxanthin. We show that: i) despite different sensitivities to photoinhibition, photoinhibition in all three plants occurred at the reaction centre of PSII; ii) there was no correlation between the extent of photoinhibition and the degradation of the D1 protein; iii) the susceptibility to photoinhibition by blockage of chloroplas-tencoded protein synthesis was much less in shade plants than in plants acclimated to higher light; and iv) inhibition of zeaxanthin formation increased the sensitivity to photoinhibition in pea, but not in the shade plant Tradescantia. We suggest that there are mechanistic differences in photoinhibition of sun and shade plants. In sun plants, an active repair cycle of PSII replaces photoinhibited reaction centres with photochemically active ones, thereby conferring partial protection against photoinhibition. However, in shade plants, this repair cycle is less important for protection against photoinhibition; instead, photoinhibited PSII reaction centres may confer, as they accumulate, increased protection of the remaining connected, functional PSII centres by controlled, nonphotochemical dissipation of excess excitation energy.     

Resistance to low temperature photoinhibition is not associated with isolated thylakoid membranes of winter rye

In vivo measurements of chlorophyll a fluorescence indicate that cold-hardened winter rye (Secale cereale L. cv Musketeer) develops a resistance to low temperature-induced photoinhibition compared with nonhardened rye. After 7.2 hours at 5°C and 1550 micromoles per square meter per second, the ratio of variable fluorescence/maximum fluorescence was depressed by only 23% in cold-hardened rye compared with 46% in nonhardened rye. We have tested the hypothesis that the principal site of this resistance to photoinhibition resides at the level of rye thylakoid membranes. Thylakoids were isolated from cold-hardened and nonhardened rye and exposed to high irradiance (1000-2600 micromoles per square meter per second) at either 5 or 20°C. The photoinhibitory response measured by room temperature fluorescence induction, photosystem II electron transport, photoacoustic spectroscopy, or [¹⁴C]atrazine binding indicates that the differential resistance to low temperature-induced photoinhibition in vivo is not observed in isolated thylakoids. Similar results were obtained whether isolated rye thylakoids were photoinhibited or thylakoids were isolated from rye leaves preexposed to a photoinhibitory treatment. Thus, we conclude that increased resistance to low temperature-induced photoinhibition is not a property of thylakoid membranes but is associated with a higher level of cellular organization.     

Operation of dual mechanisms that both lead to photoinactivation of Photosystem II in leaves by visible light

Photosystem II (PS II) is photoinactivated during photosynthesis, requiring repair to maintain full function during the day. What is the mechanism(s) of the initial events that lead to photoinactivation of PS II? Two hypotheses have been put forward. The 'excess-energy hypothesis' states that excess energy absorbed by chlorophyll (Chl), neither utilized in photosynthesis nor dissipated harmlessly in non-photochemical quenching, leads to PS II photoinactivation; the 'Mn hypothesis' (also termed the two-step hypothesis) states that light absorption by the Mn cluster in PS II is the primary effect that leads to dissociation of Mn, followed by damage to the reaction centre by light absorption by Chl. Observations from various studies support one or the other hypothesis, but each hypothesis alone cannot explain all the observations. We propose that both mechanisms operate in the leaf, with the relative contribution from each mechanism depending on growth conditions or plant species. Indeed, in a single system, namely, the interior of a leaf, we could observe one or the other mechanism at work, depending on the location within the tissue. There is no reason to expect the two mechanisms to be mutually exclusive.     

Reversible photoinhibition of unhardened and cold-acclimated spinach leaves at chilling temperatures

The photoinhibition of photosynthesis at chilling temperatures was investigated in cold-acclimated and unhardened (acclimated to +18° C) spinach (Spinacia oleracea L.) leaves. In unhardened leaves, reversible photoinhibition caused by exposure to moderate light at +4° C was based on reduced activity of photosystem (PS) II. This is shown by determination of quantum yield and capacity of electron transport in thylakoids isolated subsequent to photoinhibition and recovery treatments. The activity of PSII declined to approximately the same extent as the quantum yield of photosynthesis of photoinhibited leaves whereas PSI activity was only marginally affected. Leaves from plants acclimated to cold either in the field or in a growth chamber (+1° C), were considerably less susceptible to the light treatment. Only relatively high light levels led to photoinhibition, characterized by quenching of variable chlorophyll a fluorescence (F_V) and slight inhibition of PSII-driven electron transport. Fluorescence data obtained at 77 K indicated that the photoinhibition of cold-acclimated leaves (like that of the unhardened ones) was related to increased thermal energy dissipation. But in contrast to the unhardened leaves, 77 K fluorescence of cold-acclimated leaves did not reveal a relative increase of PSI excitation. High-light-treated, cold-acclimated leaves showed increased rates of dark respiration and a higher light compensation point. The photoinhibitory fluorescence quenching was fully reversible in low light levels both at +18° C and +4° C; the recovery was much faster than in unhardened leaves. Reversible photoinhibition is discussed as a protective mechanism against excess light based on transformation of PSII reaction centers to fluorescence quenchers.Abbreviations F_O initial fluorescence - F_M maximal fluorescence - F_V devariable fluorescence (f_m-f_o) - PFD photon flux density - PS photosystem - SD standard deviation The authors thank the Deutsche Forschungsgemeinschaft and the Academy of Finland for financial support.     

Light adaptation and the role of autotrophic epiphytes in primary production of the temperate seagrass, Zostera marina L.

Photosynthetic features of Zostera marina L. and its autotrophic epiphyte community were investigated in a population inhabiting a shallow (1.3 m depth) water meadow in Great Harbor, Woods Hole, MA (U.S.A.). Photosynthesis versus irradiance (P-I) relationships were measured with respect to leaf age determined by the leaf position in the shoot bundle and by location of the tissue along the leaf axis. Therefore both age and light intensity gradients along the leaf axis were considered. The maximum photosynthesis (P_max) per dm² typically increased nearly two-fold along the leaf axis from leaf bases to apices. Photosynthetic rate on a chlorophyll (Chl) basis did not increase as dramatically along the leaf axis, and rates were usually lowest in tissues with the highest Chl content. The P-I relationships of leaves of different ages did not reveal photoinhibition even at light intensities > 1400 μE • m⁻² • s ⁻¹. Furthermore, no photoinhibition was observed in tissues from leaf blade bases, which never experienced high light levels (> 500 μE • m ⁻² • s⁻¹) in situ in Great Harbor. The initial slopes of the P-I curves and light compensation and saturation values varied along the leaf axis in relation to in situ light intensity gradients and in relation to leaf or tissue age. It appeared that leaf and/or tissue age was more important than light environment in determining P-I responses. The contribution of the autotrophic epiphyte community on Z. marina leaves to total photosynthesis per dm² was between 27 and 50%, and between 10 and 44% per mg chlorophyll. These levels of epiphyte photosynthesis can double the primary production of Z. marina leaves. No detrimental effects of epiphyte cover were realized in leaf maximal photosynthesis or P-I relationships. Non-epiphytized leaves and leaves from which epiphytes were removed showed essentially identical photosynthetic features. Light intensity and age gradients along the leaf axis control both the photosynthetic performance of the leaves and epiphyte biomass and photosynthesis.     

Reduced photoinhibition with stem curling in the resurrection plant Selaginella lepidophylla

Summary Selaginella lepidophylla, the resurrection plant, curls dramatically during desiccation and the hypothesis that curling may help limit bright light-induced damage during desiccation and rehydration was tested under laboratory conditions. Restraint of curling during desiccation at 25° C and a constant irradiance of 2000 mol m^–2 s^]t-1 significantly decreased PSII and whole-chain electron transport and the F_v/F_m fluorescence yield ratio following rehydration relative to unrestrained plants. Normal curling during desiccation at 37.5°C and 200 mol m^–2 s^–1 irradiance did not fully protect against photoinhibition or chlorophyll photooxidation indicating that some light-induced damage occurred early in the desiccation process before substantial curling. Photosystem I electron transport was less inhibited by high-temperature, high-irradiance desiccation than either PSII or whole-chain electron transport and PSI was not significantly affected by restraint of curling during desiccation at 25°C and high irradiance. Previous curling also helped prevent photoinhibition of PSII electron transport and loss of whole-plant photosynthetic capacity as the plants uncurled during rehydration at high light. These results demonstrate that high-temperature desiccation exacerbated photoinhibition, PSI was less photoinhibited than PSII or whole-chain electron transport, and stem curling ameliorated bright light-induced damage helping to make rapid recovery of photosynthetic competence possible when the plants are next wetted.     

Use of chlorophyll fluorescence to estimate the effect of photoinhibition in outdoor cultures ofSpirulina platensis

Chlorophyll fluorescence measurements were used to evaluate the effect of temperature on photoinhibition inSpirulina platensis cultures grown in tubular reactors outdoors. Cultures grown at 35 °C during the day time showed a lower reduction in the Fv/Fm ratio as compared to cultures grown at 25 °C. It is demonstrated that the lower temperature photoinhibited cells can undergo a complete recovery once transferred to low light and higher temperature. This recovery does not take place when 100 µg ml^-1 chloramphenicol is added to cells. The recovery is light dependent and cells incubated in the dark at low temperature do not show a recovery in the Fv/Fm ratio. The data presented strongly support the hypothesis that photoinhibition takes place in outdoorSpirulina cultures. At the same time it is demonstrated that fluorescence measurements can be used as a fast reliable indication for photoinhibition in outdoor algal cultures.Author for correspondencePublication No. 69 of the Microalgal Biotechnology Laboratory.     

Strength of winter leaf redness as an indicator of stress vulnerable individuals in Pistacia lentiscus

Leaf anthocyanins are believed to afford protection against photoinhibition, yet the dependence of protection on actual anthocyanin concentrations has not been investigated. To that aim, non-invasive optical methods (spectral reflectance and chlorophyll fluorescence) were used to assess the levels of anthocyanins and chlorophylls as well as photosystem II photochemical efficiency in hundreds of leaves from the mastic tree (Pistacia lentiscus), which displays in the field a continuum of leaf tints during winter from fully green to fully red. Contrary to expectations based on the photoprotective hypothesis, the strength of leaf redness was positively correlated to the extent of mid-winter chronic photoinhibition and negatively correlated to leaf chlorophyll contents. Hence, a photoprotective role for anthocyanins is not substantiated. Instead, we argue that winter leaf redness may be used reliably, quickly and non-invasively to locate vulnerable individuals in the field.     

Light scattering,chlorophyll fluorescence and state of the adenylate system in illuminated spinach leaves

Scattering of green light and chlorophyll fluorescence by spinach leaves kept in a stream of air or nitrogen were compared with leaf adenylate levels during illumination with blue, red or far-red light. Energy charge and ATP-ADP ratios exhibited considerable variability in different leaves both in the dark and in the light. Variability is explained by different possible states of the reaction oxidizing triose phosphate or reducing 3-phosphoglycerate. Except when oxygen levels were low, there was an inverse relationship between light scattering and chlorophyll fluorescence during illumination with blue or red light. When CO₂ was added to a stream of CO₂-free air, chlorophyll fluorescence increased, sometimes after a transient decrease, and both light scattering and leaf $\text{ATP}\text{ADP}$ ratios decreased. Similar observations were made when air was replaced by nitrogen under blue or high-intensity red light. Under these conditions, over-reduction caused inhibition of electron transport and phosphorylation in chloroplasts. However, when air was replaced by nitrogen during illumination with low-intensity red light or far-red light, light scattering increased instead of decreasing. Under these light conditions, $\text{ATP}\text{ADP}$ ratios were maintained in the light. They decreased drastically only after darkening. Although $\text{ATP}\text{ADP}$ ratios responded faster than light scattering or the slow secondary decline of chlorophyll fluorescence due to illumination, it appeared that in the steady state, light scattering and chlorophyll fluorescence are useful indicators of the phosphorylation state of the leaf adenylate system at least under aerobic conditions, when chloroplast and extrachloroplast adenylate systems can effectively communicate.     

Photosynthetic Efficiency, and Photodamage by UV and Visible Radiation, in Red versus Green Leaf Coleus Varieties

The maximum quantum yield for photosynthetic O₂ evolution inred leaf coleus varieties having anthocyanin in their upperepidermis is much lower in green light and slightly lower inwhite light than in a green leaf variety lacking anthocyanin.A similar degree of photoinhibition occurred under excess visiblelight in the red versus green varieties; whereas, the red leafvarieties were less damaged by UV-B and UV-C radiation suggestingprotection by anthocyanin in their epidermal tissue. (Received October 3, 1995; Accepted February 7, 1996)