On the PsbS-induced quenching in the plant major light-harvesting complex LHCII studied in proteoliposomes.

Non-photochemical quenching (NPQ) in photosynthetic organisms provides the necessary photoprotection that allows them to cope with largely and quickly varying light intensities. It involves deactivation of excited states mainly at the level of the antenna complexes of photosystem II using still largely unknown molecular mechanisms. In higher plants the main contribution to NPQ is the so-called qE-quenching, which can be switched on and off in a few seconds. This quenching mechanism is affected by the low pH-induced activation of the small membrane protein PsbS which interacts with the major light-harvesting complex of photosystem II (LHCII). We are reporting here on a mechanistic study of the PsbS-induced LHCII quenching using ultrafast time-resolved chlorophyll (Chl) fluorescence. It is shown that the PsbS/LHCII interaction in reconstituted proteoliposomes induces highly effective and specific quenching of the LHCII excitation by a factor ≥ 20 via Chl-Chl charge-transfer (CT) state intermediates which are weakly fluorescent. Their characteristics are very broad fluorescence bands pronouncedly red-shifted from the typical unquenched LHCII fluorescence maximum. The observation of PsbS-induced Chl-Chl CT-state emission from LHCII in the reconstituted proteoliposomes is highly reminiscent of the in vivo quenching situation and also of LHCII quenching in vitro in aggregated LHCII, indicating a similar quenching mechanism in all those situations. The PsbS mutant lacking the two proton sensing Glu residues induced significant, but much smaller, quenching than wild type. Added zeaxanthin had only minor effects on the yield of quenching in the proteoliposomes. Overall our study shows that PsbS co-reconstituted with LHCII in liposomes represents an excellent in vitro model system with characteristics that are reflecting closely the in vivo qE-quenching situation.

Spectroscopic and biochemical insight into an electron-bifurcating [FeFe] hydrogenase.

The heterotrimeric electron-bifurcating [FeFe] hydrogenase (HydABC) from Thermotoga maritima (Tm) couples the endergonic reduction of protons (H+) by dihydronicotinamide adenine dinucleotide (NADH) (∆G0 ≈ 18 kJ mol-1) to the exergonic reduction of H+ by reduced ferredoxin (Fdred) (∆G0 ≈ - 16 kJ mol-1). The specific mechanism by which HydABC functions is not understood. In the current study, we describe the biochemical and spectroscopic characterization of TmHydABC recombinantly produced in Escherichia coli and artificially maturated with a synthetic diiron cofactor. We found that TmHydABC catalyzed the hydrogen (H2)-dependent reduction of nicotinamide adenine dinucleotide (NAD+) in the presence of oxidized ferredoxin (Fdox) at a rate of ≈17 µmol NADH min-1 mg-1. Our data suggest that only one flavin is present in the enzyme and is not likely to be the site of electron bifurcation. FTIR and EPR spectroscopy, as well as FTIR spectroelectrochemistry, demonstrated that the active site for H2 conversion, the H-cluster, in TmHydABC behaves essentially the same as in prototypical [FeFe] hydrogenases, and is most likely also not the site of electron bifurcation. The implications of these results are discussed with respect to the current hypotheses on the electron bifurcation mechanism of [FeFe] hydrogenases. Overall, the results provide insight into the electron-bifurcating mechanism and present a well-defined system for further investigations of this fascinating class of [FeFe] hydrogenases.

Unique Spectroscopic Properties of the H-Cluster in a Putative Sensory [FeFe] Hydrogenase.

Sensory type [FeFe] hydrogenases are predicted to play a role in transcriptional regulation by detecting the H2 level of the cellular environment. These hydrogenases contain the hydrogenase domain with distinct modifications in the active site pocket, followed by a Per-Arnt-Sim (PAS) domain. As yet, neither the physiological function nor the biochemical or spectroscopic properties of these enzymes have been explored. Here, we present the characterization of an artificially maturated, putative sensory [FeFe] hydrogenase from Thermotoga maritima (HydS). This enzyme shows lower hydrogen conversion activity than prototypical [FeFe] hydrogenases and a reduced inhibition by CO. Using FTIR spectroelectrochemistry and EPR spectroscopy, three redox states of the active site were identified. The spectroscopic signatures of the most oxidized state closely resemble those of the Hox state from the prototypical [FeFe] hydrogenases, while the FTIR spectra of both singly and doubly reduced states show large differences. The FTIR bands of both the reduced states are strongly red-shifted relative to the Hox state, indicating reduction at the diiron site, but with retention of the bridging CO ligand. The unique functional and spectroscopic features of HydS are discussed with regard to the possible role of altered amino acid residues influencing the electronic properties of the H-cluster.

Proton Coupled Electronic Rearrangement within the H-Cluster as an Essential Step in the Catalytic Cycle of [FeFe] Hydrogenases.

The active site of [FeFe] hydrogenases, the H-cluster, consists of a [4Fe-4S] cluster connected via a bridging cysteine to a [2Fe] complex carrying CO and CN- ligands as well as a bridging aza-dithiolate ligand (ADT) of which the amine moiety serves as a proton shuttle between the protein and the H-cluster. During the catalytic cycle, the two subclusters change oxidation states: [4Fe-4S]H2+ ⇔ [4Fe-4S]H+ and [Fe(I)Fe(II)]H ⇔ [Fe(I)Fe(I)]H thereby enabling the storage of the two electrons needed for the catalyzed reaction 2H+ + 2e- ⇄ H2. Using FTIR spectro-electrochemistry on the [FeFe] hydrogenase from Chlamydomonas reinhardtii (CrHydA1) at different pH values, we resolve the redox and protonation events in the catalytic cycle and determine their intrinsic thermodynamic parameters. We show that the singly reduced state Hred of the H-cluster actually consists of two species: Hred = [4Fe-4S]H+ - [Fe(I)Fe(II)]H and HredH+ = [4Fe-4S]H2+ - [Fe(I)Fe(I)]H (H+) related by proton coupled electronic rearrangement. The two redox events in the catalytic cycle occur on the [4Fe-4S]H subcluster at similar midpoint-potentials (-375 vs -418 mV); the protonation event (Hred/HredH+) has a pKa ≈ 7.2.

Intercluster Redox Coupling Influences Protonation at the H-cluster in [FeFe] Hydrogenases.

[FeFe] hydrogenases catalyze proton reduction and hydrogen oxidation displaying high rates at low overpotential. Their active site is a complex cofactor consisting of a unique [2Fe] subcluster ([2Fe]H) covalently bound to a canonical [4Fe-4S] cluster ([4Fe-4S]H). The [FeFe] hydrogenase from Desulfovibrio desulfuricans is exceptionally active and bidirectional. This enzyme features two accessory [4Fe-4S]F clusters for exchanging electrons with the protein surface. A thorough understanding of the mechanism of this efficient enzyme will facilitate the development of synthetic molecular catalysts for hydrogen conversion. Here, it is demonstrated that the accessory clusters influence the catalytic properties of the enzyme through a strong redox interaction between the proximal [4Fe-4S]F cluster and the [4Fe-4S]H subcluster of the H-cluster. This interaction enhances proton-coupled electronic rearrangement within the H-cluster increasing the apparent pKa of its one electron reduced state. This may help to sustain H2 production at high pH values. These results may apply to all [FeFe] hydrogenases containing accessory clusters.

Pigment interactions in light-harvesting complex II in different molecular environments.

Extraction of plant light-harvesting complex II (LHCII) from the native thylakoid membrane or from aggregates by the use of surfactants brings about significant changes in the excitonic circular dichroism (CD) spectrum and fluorescence quantum yield. To elucidate the cause of these changes, e.g. trimer-trimer contacts or surfactant-induced structural perturbations, we compared the CD spectra and fluorescence kinetics of LHCII aggregates, artificial and native LHCII-lipid membranes, and LHCII solubilized in different detergents or trapped in polymer gel. By this means we were able to identify CD spectral changes specific to LHCII-LHCII interactions, at (-)-437 and (+)-484 nm, and changes specific to the interaction with the detergent n-dodecyl-ß-maltoside (ß-DM) or membrane lipids, at (+)-447 and (-)-494 nm. The latter change is attributed to the conformational change of the LHCII-bound carotenoid neoxanthin, by analyzing the CD spectra of neoxanthin-deficient plant thylakoid membranes. The neoxanthin-specific band at (-)-494 nm was not pronounced in LHCII in detergent-free gels or solubilized in the α isomer of DM but was present when LHCII was reconstituted in membranes composed of phosphatidylcholine or plant thylakoid lipids, indicating that the conformation of neoxanthin is sensitive to the molecular environment. Neither the aggregation-specific CD bands, nor the surfactant-specific bands were positively associated with the onset of fluorescence quenching, which could be triggered without invoking such spectral changes. Significant quenching was not active in reconstituted LHCII proteoliposomes, whereas a high degree of energetic connectivity, depending on the lipid:protein ratio, in these membranes allows for efficient light harvesting.

Hematological Alterations in Common Carp (Cyprinus carpio L.) Exposed to Herbicides: Pendimethalin and Ethofumesate Tested Separately and in Mixture.

Herbicides are used in large amounts in agriculture and the evaluation of their toxic effects is of major concern to environmental safety. The aim of the present study was to investigate common carp hematological alterations caused by herbicide exposure. Fish were treated with pendimethalin and ethofumesate tested separately and in mixture administered to aquarium water. Peripheral blood of treated fish was collected after 1, 3 and 7 days of exposure and compared to control. The total number of erythrocytes (RBC), total number of leukocytes (WBC), hematocrit value (Hct), total hemoglobin concentration (Hb), mean corpuscular hemoglobin (MCH), mean corpuscular volume (MCV), mean corpuscular hemoglobin concentration (MCHC) and leukograms were determined at once. The results indicate that herbicide exposure caused different changes in the hematological profile of the fish. In the case of exposure to individual herbicides, short-term fluctuations of various hematological indices were noted. Moreover, a significant increase in RBC and Hct after a short period of exposure (1-3 days) in fish exposed simultaneously to both tested herbicides was observed. Exposure to herbicides affected the leukocyte profile after 3 and 7 days of duration. Fluctuations of hematological parameters are a typical change in fish exposed to pesticides.

Effect of the LHCII pigment-protein complex aggregation on photovoltaic properties of sensitized TiO2 solar cells.

A modified dye-sensitized solar cell consisting of a thin TiO2 barrier layer sensitized with natural trimeric light-harvesting complex II (LHCII) from spinach was used as a biomimetic model to study the effects of LHCII aggregation on the photovoltaic properties. The aggregation of individual trimers induced molecular reorganization, which dramatically increased the photocurrent. The morphology of small- and large-size LHCII aggregates deposited on a surface was confirmed by atomic force microscopy. Enhanced LHCII immobilization was accomplished via electrostatic interaction with amine-functionalized photoanodes. The photocurrent responses of the assembled solar cells under illumination at three characteristic wavelength bands in the UV-Vis absorption spectra of LHCII solutions confirmed that a significant photocurrent was generated by LHCII photosensitizers. The enhanced photocurrent by large aggregated LHCII is shown to correlate with the quenching in the far-red fluorescence deriving from chlorophyll-chlorophyll charge transfer states that are effectively coupled with the TiO2 surface and thus inject electrons into the TiO2 conduction band. The large aggregated LHCII with more chlorophyll-chlorophyll charge transfer states is a much better sensitizer since it injects electrons more efficiently into the conduction band of TiO2 than the small aggregated LHCII mostly consisting of unquenched chlorophyll excited state. The assembled solar cells demonstrated remarkable stability in both aqueous buffer and acetonitrile electrolytes over 30 days.

The effect of methionine and folic acid administered in ovo on the blood biochemical parameters of chickens (Gallus gallus domesticus).

Methionine is one of the most frequently supplemented amino acids in raising of poultry. However, an overdose of methionine can cause hyperhomocysteinemia. Folic acid, taking part in the process of homocysteine remethylation, is a factor affecting the reduction of the concentration of this amino acid. The study was carried out in 2 stages. The experiment of step I was to investigate the effect of methionine and/or folic acid administration in ovo in the early stage of embryogenesis (E4), and the experiment of the second stage - in the late stage of embryogenesis (E17) on the following biochemical parameters of chicken blood: glucose concentration in whole blood and concentration of homocysteine and uric acid in plasma of domestic chickens (Gallus gallus domesticus). Our results confirm that methionine supplementation may increase the concentration of uric acid and homocysteine. Moreover, we demonstrated that folic acid administered during embryogenesis decreased homocysteine concentration, also in groups simultaneously supplemented with methionine, especially in the initial stage of postnatal life of the bird.

Effect of weak electromagnetic field on cardiac work, concentration of thyroid hormones and blood aminotransferase level in the chick embryo.

The aim of the study was to determine the effect of alternating electromagnetic field (EMF; 50 Hz frequency, 50 and 100 µT induction) on cardiac work of the chick embryo. Eggs from the experimental groups were exposed to EMF throughout incubation. During the experiment, heart rate (ballistocardiographic method), thyroxine (T4) and triiodothyronine (T3) concentrations, heart weight, ventricle wall thickness, and levels of alanine aminotransferase (ALT) and aspartate aminotransferase (AST) were determined. The results show, for the first time, that the exposure of chick embryos to EMF augments the heart rate, especially from 17 days of incubation. The increased heart rate in the embryos exposed to EMF was associated with considerable increases in plasma T4 and T3 concentrations, which were recorded during the final stage of embryogenesis. The significant effect of the 100-µT field on heart weight and blood AST levels in the embryos suggests that EMF has a direct effect on the physiological function of cardiac muscle.

A novel method for automatically analysing the activity of fast-moving animals: a case study of Callimico goeldii monkeys housed in a zoological garden.

Behavioural indices are recognised as important criteria for assessing animal welfare. One of the basic animal behaviours included in ethograms is their activity. The assessment of fast-moving animals, performed by humans using the visual observation method, is difficult and not very objective. Therefore, the aim of the research was to develop a method of automated analysis of animal activity, particularly useful in the observation of quick and lively individuals, and to prove its suitability for assessing the behaviour of fast-moving animals. A method of automatically assessing animal activity was developed using digital image analysis, with the Python programming language and the OpenCV library being the foundational tools. The research model was Callimico goeldii monkeys housed in a zoological garden. This method has been proved to correlate well (Rs = 0.76) with the visual method of animal behaviour analysis. The developed automatic evaluation of animal behaviour is many times faster than visual analysis, and it enables precise assessment of the daily activity of fast-moving groups of animals. The use of this system makes it possible to obtain an activity index with sub-second resolution, which allows it to be used in online mode as a detector of abnormal animal activity, e.g. early detection of illnesses or sudden events that are manifested by increased or decreased activity in relation to the standard activity pattern.

Health risks for children exercising in an air-polluted environment can be reduced by monitoring air quality with low-cost particle sensors.

A child's body is highly sensitive to air quality, especially regarding the concentration of particulate matter (PM). Nevertheless, due to the high cost of precision instruments, measurements of PM concentrations are rarely carried out in school areas where children spend most of their daily time. This paper presents the results of PM measurements made by a validated, low-cost university air pollution measurement system operating in a rural area near schools. An assessment of children's exposure to PM during school hours (8 a.m.-6 p.m.) at different times of the year was carried out. We show that PM10 concentrations in the air, particularly in winter, often exceeded the alert values of 50 µg m-3, posing a health risk to children, especially when children exercise outside the school building. We also calculated the rate and total PM10 deposition in the respiratory tract during various physical activities performed in clean and polluted air. Monitoring actual PM10 concentrations as presented in this paper, using a low cost sensors, offer school authorities and teachers an opportunity to reduce health risks for children. This can be achieved by adjusting the duration and exercise intensity of children's outdoor physical activities according to the measured air quality.

The concentration of particulate matter in the barn air and its influence on the content of heavy metals in milk.

Heavy metals are one of the components of smog, which is mainly the product of burning fossil fuels in residential buildings. These elements, introduced into the body of cattle by inhalation, may enter the milk. The goal of this study was to assess the impact of particulate pollution in the atmospheric air on the concentration of particulate matter in the air of a dairy cattle barn and on the content of selected heavy metals in milk from cows present in the building. Measurements were taken between November and April (148 measurement days). The calculations carried out showed a high correlation (RS = + 0.95) between the concentrations of particulates measured outside and inside the barn, which is indicative of a significant impact of the atmospheric air on the particulate pollution level of the livestock building. The number of days in excess of the daily standard for PM10 inside was 51. The conducted analysis of the chemical composition of the milk collected under high particulate pollution (February) showed that the permitted lead level had been exceeded-21.93 µg/kg (norm 20.00 µg/kg).

Chemical properties of the coffee grounds and poultry eggshells mixture in terms of soil improver.

Spent coffee grounds (SCG) as well as chicken (CES) or duck eggshells (DES) left over from the artificial hatching technology are proposed as potential soil improver and/or organic-mineral fertiliser components. Therefore, it seems interesting and necessary to evaluate the chemical composition of these wastes and their mixtures in terms of their possible use for that purpose. The study was conducted under the incubation experiment conditions using a mixture of SCG and eggshells (10:1 ratio). Macronutrients, i.e. C, N, S, were determined by the catalytic combustion method, while P, K, Mg, Ca, Na by atomic spectrometry. It was found that SCG were rich in C, N, P, and K, while eggshells in Ca, Mg, Na, and S. However, CES compared to DES were richer in deacidifying components (i.e. Ca, Mg, K). At the same time, the content of macronutrients in eggshells decreased gradually along with the embryo development. For this reason, the mixture of SCG and shells of unembryonated chicken eggs (CES I) had the best chemical and usable proprieties. To conclude, the chemical properties of the mixtures of spent coffee grounds and eggshells indicate their possible application in soil bioengineering.

Staphylococci and fecal bacteria as bioaerosol components in animal housing facilities in the Zoological Garden in Chorzów.

Zoos are places open for a large number of visitors, adults and children, who can admire exotic as well as indigenous animal species. The premises for animals may contain pathogenic microbes, including those exhibiting antibiotic resistance. It poses a threat to people remaining within the zoo premises, both for animal keepers who meet animals on a daily basis and visitors who infrequently have contact with animals. There are almost no studies concerning the presence on the concentration of airborne bacteria, especially staphylococci and fecal bacteria in animal shelters in the zoo. There is no data about antibiotic resistance of staphylococci in these places. The results will enable to determine the scale of the threat that indicator bacteria from the bioaerosol pose to human health within zoo premises. This study conducted in rooms for 5 animals group (giraffes, camels, elephants, kangaroos, and Colobinae (species of monkey)) in the Silesian Zoological Garden in Chorzów (Poland). The bioaerosol samples were collected using a six-stage Andersen cascade impactor to assess the concentrations and size distribution of airborne bacteria. Staphylococci were isolated from bioaerosol and tested for antibiotic resistance. In our study, the highest contamination of staphylococci and fecal bacteria was recorded in rooms for camels and elephants, and the lowest in rooms for Colobinae. At least 2/3 of bacteria in bioaerosol constituted respirable fraction that migrates into the lower respiratory tract of the people. In investigated animal rooms, the greatest bacteria contribution was recorded for bioaerosol fraction sized 1.1-3.3µm. Bacterial concentrations were particularly strong in spring and autumn, what is related to shedding fur by animals. Among the isolated staphylococci which most often occurred were Staphylococcus succinus, S. sciuri, and S. vitulinus. The highest antibiotic resistance was noted in the case of Staphylococcus epidermidis, while the lowest for S. xylosus. In addition to standard cleaning of animal rooms, periodic disinfection should be considered. Cleaning should be carried out wet, which should reduce dust, and thus the concentrations of bacteria in the air of animal enclosures.

Impact of bacterial aerosol, particulate matter, and microclimatic parameters on animal welfare in Chorzów (Poland) zoological garden.

Zoos are very popular facilities visited by entire families with children, who come there to watch live animals. Zoos also provide workplaces for a large number of people directly looking after the animals. For places designed to house animals, regardless of whether they are farm animals, pets, or zoo animals, a higher concentration of both dust and potentially harmful bioaerosols can be expected. Unfortunately, there are almost no studies concerning the concentration of bacterial bioaerosols and particulate matter in animal shelters that would answer the question whether the level of these pollutants is constant or variable and dependent on a particular zoo, group of animals, their number in enclosures, or season. This study aimed to assess the levels of bacterial aerosol in rooms intended for animals (giraffes, camels, elephants, kangaroos, and colobinae) in the Silesian Zoological Garden in Chorzów (Poland). The bioaerosol samples were collected using a six-stage Andersen cascade impactor to assess the concentrations and size distribution of airborne bacteria. Particulate matter (PM10) was assessed using an electronic dust meter. Measurements of microclimate parameters were carried out using the Airflow™ Instruments Velocity Meter TA440, while gas concentrations were determined applying GFG Microtector II G450. The results showed that the concentration of airborne bacteria varied significantly between facilities for the analyzed animal groups. The lowest concentration of the total bacterial aerosol was observed in enclosures for colobinae (approx. 850 CFU/m3), while the highest-in rooms for elephants (approx. 105,600 CFU/m3). The average share of respirable fraction of bacteria was quite high, with values ranging from 62.9 (colobinae) to 86.9% (elephants), indicating potential harmfulness to the health of exposed people. PM10 concentrations were relatively low (10-86 µg/m3) and did not exceed the limit values for occupational exposure. Moreover, the levels of bacterial bioaerosol in almost all cases did not exceed the limit values. As the animals constitute a significant source of bioaerosol, attention should be paid to thorough cleaning of animals and their shelters, as well as maintaining appropriate levels of microclimate parameters in the facilities.

The impact of smog on the concentration of particulate matter in the antelope house in the Silesian zoological garden.

Persistent negligence in the field of environmental protection in Poland as well as strong dependance of the energy sector on the fossil fuels have led to serious pollution of the air with particulate matter, which at high concentrations is capable of penetrating into the buildings. The aim of this study is to assess the impact of particulate airborne pollution on the concentration of particulate matter inside the antelope house in the Silesian Zoological Garden located within the Upper Silesia in Poland. The research was conducted from February to May in 2018. The records taken in the research period show that the concentration of PM10 exceeded the level of 50 µg/m3 outside the building during 26 days and 11 days when it comes to the concentration of particulate matter inside the antelope house. The quantity of particulate matter in the antelope house is strongly correlated with the concentration of the particles in the air. Despite fitting existing ventilation system with a filter that reduces the dust level by 60% during the highest level of smog, particulate matter concentration in the antelope house exceeded acceptable limit for PM10 more than twofold. Particle size-fraction analysis revealed that as much as 85% of the particles detected in the studied compartment constitute PM2.5.

Preliminary evaluation of application of a 3-dimensional network structure of siloxanes Dergall preparation on chick embryo development and microbiological status of eggshells.

The spatial network structure of Dergall is based on substances nontoxic to humans and the environment which, when applied on solid surfaces, creates a coating that reduces bacterial cell adhesion. The bacteriostatic properties of siloxanes are based on a purely physical action mechanism which excludes development of drug-resistant microorganisms. The aims of the present study were to 1) evaluate a Dergall layer formed on the eggshell surface regarding the potential harmful effects on the chick embryo; 2) evaluate antimicrobial activity and estimate the prolongation time of Dergall's potential antimicrobial activity. Dergall at a concentration of 0.6% formed a layer on the eggshell surface. In vitro testing of the potential harmful effects of Dergall by means of a hen embryo test of the chorioallantoic membrane showed no irritation reaction at a concentration of 3% and lower. The hatchability of the groups sprayed with a Dergall water solution with a concentration of 0 to 5% was 89.1 to 93.8% for fertilized eggs (P > 0.05) but decreased to 63.7% (P < 0.05) in the group sprayed with a 6% concentration of the solution. This phenomenon was caused by embryo mortality in the first week of incubation. At the concentration of 0.6%, Dergall exhibited strong antibacterial properties against bacteria such as Staphylococcus aureus, Escherichia coli, Shigella dysenteriae, Shigella flexneri, and Salmonella typhimurium. For Streptococcus pyogenes, the highest antibacterial activity of Dergall was reported in the concentrations of 100 and 50%. For Pseudomonas aeruginosa, no antibacterial activity of Dergall was generally observed, but in vivo testing showed a strong decrease of all gram-negative bacteria growth. Moreover, a prolonged antimicrobial effect lasting until 3 D after disinfection was observed, which makes Dergall a safe and efficient disinfectant.

Anisotropic Circular Dichroism of Light-Harvesting Complex II in Oriented Lipid Bilayers: Theory Meets Experiment.

Anisotropic circular dichroism (ACD) spectroscopy of macroscopically aligned molecules reveals additional information about their excited states that is lost in the CD of randomly oriented solutions. ACD spectra of light-harvesting complex II (LHCII)-the main peripheral antenna of photosystem II in plants-in oriented lipid bilayers were recorded from the far-UV to the visible wavelength region. ACD spectra show a drastically enhanced magnitude and level of detail compared to the isotropic CD spectra, resolving a greater number of bands and weak optical transitions. Exciton calculations show that the spectral features in the chlorophyll Q y region are well-reproduced by an existing Hamiltonian for LHCII, providing further evidence for the identity of energy sinks at chlorophylls a603 and a610 in the stromal layer and chlorophylls a604 and a613 in the luminal layer. We propose ACD spectroscopy to be a valuable tool linking the three-dimensional structure and the photophysical properties of pigment-protein complexes.