Manganese concentration affects chloroplast structure and the photosynthetic apparatus in Marchantia polymorpha.

Manganese (Mn) is an essential metal for plant growth. The most important Mn-containing enzyme is the Mn4CaO5 cluster that catalyzes water oxidation in photosystem II (PSII). Mn deficiency primarily affects photosynthesis, whereas Mn excess is generally toxic. Here, we studied Mn excess and deficiency in the liverwort Marchantia polymorpha, an emerging model ideally suited for analysis of metal stress since it accumulates rapidly toxic substances due to the absence of well-developed vascular and radicular systems and a reduced cuticle. We established growth conditions for Mn excess and deficiency and analyzed the metal content in thalli and isolated chloroplasts. In vivo super-resolution fluorescence microscopy and transmission electron microscopy revealed changes in the organization of the thylakoid membrane under Mn excess and deficiency. Both Mn excess and Mn deficiency increased the stacking of the thylakoid membrane. We investigated photosynthetic performance by measuring chlorophyll fluorescence at room temperature and 77 K, measuring P700 absorbance, and studying the susceptibility of thalli to photoinhibition. Nonoptimal Mn concentrations changed the ratio of PSI to PSII. Upon Mn deficiency, higher non-photochemical quenching was observed, electron donation to PSI was favored, and PSII was less susceptible to photoinhibition. Mn deficiency seemed to favor cyclic electron flow around PSI, thereby protecting PSII in high light. The results presented here suggest an important role of Mn in the organization of the thylakoid membrane and photosynthetic electron transport.

Ocular diagnostics, ophthalmic findings, and conjunctival microbiome in the Chilean flamingo (Phoenicopterus chilensis).

OBJECTIVE: To establish normative data for selected ocular diagnostic tests and commensal conjunctival microflora and describe the incidence of ocular pathology in Chilean flamingos. ANIMALS STUDIED: A total of 41 Chilean flamingos were examined at the Blank Park Zoo in Des Moines, Iowa. PROCEDURES: In 20 flamingos, blink rate was assessed undisturbed in their exhibit, then gentle manual restraint was used to assess palpebral fissure length (PFL), aqueous tear production (phenol red thread test [PRTT] in one eye, endodontic absorbent paper point tear test [EAPPTT] in the other), intraocular pressure (IOP; rebound tonometry), and fluorescein staining. Twenty-one other flamingos were brought to a darkened area for neuro-ophthalmic examination, slit lamp biomicroscopy, and indirect ophthalmoscopy. Swabs from seven flamingos were used for ocular microbiome evaluation. RESULTS: Results are presented as mean ± standard deviation (range). Flamingos comprised 23 females/18 males, aged 11 ± 9.1 (0.7-40) years. Test results: blink rate, 3.7 ± 2 (1-9) blinks/min; PFL, 11.2 ± 1.2 (9-14) mm; IOP, 14 ± 3.2 (10-22) mmHg; EAPPT, 10.2 ± 2.8 (9-14) mm/min; PRTT, 6.8 ± 2.5 (3-13) mm/15 s. Dazzle reflex was positive in four birds examined. Pathologies included cataracts (n = 7 birds), corneal fibrosis (n = 3), endothelial pigment (n = 2), uveal cysts (n = 1), lens luxation (n = 1), and uveitis (n = 1). Ocular microbiome showed high diversity of taxa. CONCLUSIONS: Baseline ocular parameters and incidence of ophthalmic pathology assist veterinarians with disease screening for Chilean flamingos, while the ocular microbiome showed high diversity.

DEXMEDETOMIDINE, BUTORPHANOL, AND MIDAZOLAM AS A REVERSIBLE INDUCTION PROTOCOL IN NORTH AMERICAN RIVER OTTERS (LONTRA CANADENSIS).

Captured free-ranging North American river otters (Lontra canadensis) were immobilized for the placement of intra-abdominal radio transmitters in cooperation with the Iowa Department of Natural Resources. Twenty-four otters were induced with dexmedetomidine (0.03 mg/kg, IM), butorphanol (0.2 mg/kg, IM), and midazolam (0.15 mg/kg, IM) combined in one syringe. The otters were maintained on isoflurane during the surgical procedure. Heart rate and rhythm, respiratory rate, rectal temperature, and peripheral capillary oxygen saturation were recorded every 5 min for the duration of the procedures. The otters were reversed with atipamezole (0.3-2 mg/kg, IM), naltrexone (0.6 mg/kg, IM), and flumazenil (0.05 mg/kg, IM). Rapid and smooth induction was seen, with lateral recumbency reached within 6.2 ± 5.6 min. Episodes of resedation were seen in four otters that received 0.3 mg/kg atipamezole so the dose was increased to 1 mg/kg, and no further resedation events were noted. Two fatal complications occurred secondary to suspected respiratory arrest during recovery. This drug protocol provided a smooth and rapid induction in free-ranging river otters, but further research is required to determine the safety of this protocol for river otters in both zoo and free-ranging situations.

Developing outreach events that impact underrepresented students: Are we doing it right?

Many outreach programs share the common goals of serving underrepresented groups in STEM and improving public attitudes toward science. To meet these goals, scientists must find ways to both reach the appropriate audience and communicate the importance of science in meaningful and accessible ways. This requires careful consideration of the outreach method being used. Two common outreach methods include in-school visits (scientist in the classroom) and science fairs or open houses. Here, we compare the effectiveness of these two outreach methods in meeting the goals of reaching underrepresented students and/or students with less initial interest in science. We have found that in-school visits reached more underrepresented students and that initial attitudes toward science scores were lower for in-school visit participants than for open house event participants. Importantly, positive attitudes toward science increased significantly after in-school outreach events. Taken together, these data suggest that outreach events that are taken out into the community will reach a less enthusiastic but more diverse audience and can have a positive impact on attitudes toward science within these populations. These studies highlight the importance of knowing the goals of your outreach program and choosing the method that is best suited to meeting those goals.

Immediate and Long-Term Implications of the COVID-19 Pandemic for People With Disabilities.

Some people with disabilities may have greater risk of contracting COVID-19 or experiencing worse outcomes if infected. Although COVID-19 is a genuine threat for people with disabilities, they also fear decisions that might limit lifesaving treatment should they contract the virus.During a pandemic, health systems must manage excess demand for treatment, and governments must enact heavy restrictions on their citizens to prevent transmission. Both actions can have a negative impact on people with disabilities.Ironically, the sociotechnical advances prompted by this pandemic could also revolutionize quality of life and participation for people with disabilities. Preparation for future disasters requires careful consideration.

IDENTIFICATION AND CORRELATION OF A NOVEL SIADENOVIRUS IN A FLOCK OF BUDGERIGARS (MELOPSITTACUS UNDULATES) INFECTED WITH SALMONELLA TYPHIMURIUM IN THE UNITED STATES.

A flock of budgerigars (Melopsittacus undulates) was purchased from a licensed breeder and quarantined at a zoologic facility within the United States in 2016. Following 82 deaths within the flock, the remaining 66 birds were depopulated because of ongoing clinical salmonellosis despite treatment. Gross necropsy was performed on all 66 birds. Histopathologic examination was performed on 10 birds identified with gross lesions and 10 birds without. Pathologic findings were most often observed in the liver, kidney, and spleen. Lesions noted in the livers and spleens were consistent with published reports of salmonellosis in psittacine species. Multisystemic changes associated with septicemia were not noted, most likely because of antibiotic intervention before euthanasia. Of the 20 budgerigars evaluated by histopathology, six had large basophilic intranuclear inclusion bodies within tubular epithelia in a portion of the kidneys. Electronic microscopy, next-generation sequencing, Sanger sequencing, and phylogenetic analyses were used to identify and categorize the identified virus as a novel siadenovirus strain BuAdV-1 USA-IA43444-2016. The strain was 99% similar to budgerigar adenovirus 1 (BuAdV-1), previously reported in Japan, and to a psittacine adenovirus 5 recently identified in a U.S. cockatiel. Salmonella typhimurium carriers were identified via polymerase chain reaction (PCR) and bacterial culture and compared with viral carriers identified via PCR. Inclusion bodies and Salmonella detection were significant in birds with gross lesions versus those without; however, there was no correlation between budgerigars positive with siadenovirus by PCR and concurrent Salmonella infection. Identifying subclinical siadenovirus strain BuAdV-1 USA-IA43444-2016 infection in this flock significantly differs from a previous report of clinical illness in five budgerigars resulting in death caused by BuAdV-1 in Japan. S. typhimurium remains a significant pathogen in budgerigars, and zoonotic concerns prompted depopulation to mitigate the public health risks of this flock.

An Effective Model for Engaging Faculty and Undergraduate Students in Neuroscience Outreach with Middle Schoolers.

Engaging undergraduate students in science outreach events is critical for improving future communication between scientists and community members. Outreach events are opportunities for faculty and undergraduates to utilize active learning strategies to engage non-scientists in scientific questions and principles. Through careful design of outreach events, undergraduate students can practice science communication skills while reaching populations of the public that remain underserved and underrepresented in scientific fields. Here we describe a classroom outreach event designed to give a broad overview of the field of neuroscience to middle school students of all backgrounds by delivering the content in school, during school hours. Through a variety of active learning strategies, middle school students learned about basic structures of the brain and their corresponding functions. Additionally, these students participated in demonstrations during which they generated and tested their own hypotheses and learned about sensory transmission and responses. We designed the lesson to meet the educational goals for middle school students, fulfilling the criteria for the Next Generation Science Standard MS-LS1-8 (NGSS Lead States, 2013). We evaluated the impact of the event on both undergraduate student instructors and middle school participants. Our results demonstrate that these outreach events effectively deliver new content to middle school students while also reinforcing the importance and value of outreach to undergraduate instructors.

Apoprotein heterogeneity increases spectral disorder and a step-wise modification of the B850 fluorescence peak position.

It has already been established that the quaternary structure of the main light-harvesting complex (LH2) from the photosynthetic bacterium Rhodopseudomonas palustris is a nonameric 'ring' of PucAB heterodimers and under low-light culturing conditions an increased diversity of PucB synthesis occurs. In this work, single molecule fluorescence emission studies show that different classes of LH2 'rings' are present in "low-light" adapted cells and that an unknown chaperon process creates multiple sub-types of 'rings' with more conformational sub-states and configurations. This increase in spectral disorder significantly augments the cross-section for photon absorption and subsequent energy flow to the reaction centre trap when photon availability is a limiting factor. This work highlights yet another variant used by phototrophs to gather energy for cellular development.

LESIONS OF THE FEMALE REPRODUCTIVE TRACT IN JAPANESE MACAQUE ( MACACA FUSCATA) FROM TWO CAPTIVE COLONIES.

Reproductive lesions have been described in various nonhuman primate species, including rhesus macaques ( Macaca mulatta), cynomolgus macaques ( Macaca fascicularis), baboons ( Papio spp.), squirrel monkeys ( Saimiri sciureus), and chimpanzees ( Pan spp.); however, there are few publications describing reproductive disease and pathology in Japanese macaques ( Macaca fuscata). A retrospective evaluation of postmortem reports for two captive M. fuscata populations housed within zoos from 1982 through 2015 was completed, comparing reproductive diseases diagnosed by gross pathology and histopathology. Disease prevalence, organs affected, and median age at death between the two institutions was also compared. Fifteen female captive M. fuscata, ranging in age from 15 to 29 yr were identified with reproductive tract lesions, including endometriosis, endometritis, leiomyoma, leiomyosarcoma, and adenomyosis. No significant differences were identified in disease prevalence, organs affected, and median age of death between the two institutions. Endometriosis was the most common disease process identified and was found in 10 of the 15 cases (66.7%), followed by leiomyoma (4 of 15; 26.7%). In four cases (26.7%), severe endometriosis and secondary hemorrhage was indicated as the cause of death or the primary reason for humane euthanasia. These findings were compared with a separate population of Japanese macaques managed within a research facility in the United States, with a prevalence of endometriosis of 7.6%. This study discusses possible risk factors and potential treatment options for the management of endometriosis in captive M. fuscata.

Vibrational techniques applied to photosynthesis: Resonance Raman and fluorescence line-narrowing.

Resonance Raman spectroscopy may yield precise information on the conformation of, and the interactions assumed by, the chromophores involved in the first steps of the photosynthetic process. Selectivity is achieved via resonance with the absorption transition of the chromophore of interest. Fluorescence line-narrowing spectroscopy is a complementary technique, in that it provides the same level of information (structure, conformation, interactions), but in this case for the emitting pigment(s) only (whether isolated or in an ensemble of interacting chromophores). The selectivity provided by these vibrational techniques allows for the analysis of pigment molecules not only when they are isolated in solvents, but also when embedded in soluble or membrane proteins and even, as shown recently, in vivo. They can be used, for instance, to relate the electronic properties of these pigment molecules to their structure and/or the physical properties of their environment. These techniques are even able to follow subtle changes in chromophore conformation associated with regulatory processes. After a short introduction to the physical principles that govern resonance Raman and fluorescence line-narrowing spectroscopies, the information content of the vibrational spectra of chlorophyll and carotenoid molecules is described in this article, together with the experiments which helped in determining which structural parameter(s) each vibrational band is sensitive to. A selection of applications is then presented, in order to illustrate how these techniques have been used in the field of photosynthesis, and what type of information has been obtained. This article is part of a Special Issue entitled: Vibrational spectroscopies and bioenergetic systems.

Mapping energy transfer channels in fucoxanthin-chlorophyll protein complex.

Fucoxanthin-chlorophyll protein (FCP) is the key molecular complex performing the light-harvesting function in diatoms, which, being a major group of algae, are responsible for up to one quarter of the total primary production on Earth. These photosynthetic organisms contain an unusually large amount of the carotenoid fucoxanthin, which absorbs the light in the blue-green spectral region and transfers the captured excitation energy to the FCP-bound chlorophylls. Due to the large number of fucoxanthins, the excitation energy transfer cascades in these complexes are particularly tangled. In this work we present the two-color two-dimensional electronic spectroscopy experiments on FCP. Analysis of the data using the modified decay associated spectra permits a detailed mapping of the excitation frequency dependent energy transfer flow with a femtosecond time resolution.

Conformational switching in a light-harvesting protein as followed by single-molecule spectroscopy.

Among the ultimate goals of protein physics, the complete, experimental description of the energy paths leading to protein conformational changes remains a challenge. Single protein fluorescence spectroscopy constitutes an approach of choice for addressing protein dynamics, and, among naturally fluorescing proteins, light-harvesting (LH) proteins from purple bacteria constitute an ideal object for such a study. LHs bind bacteriochlorophyll a molecules, which confer on them a high intrinsic fluorescence yield. Moreover, the electronic properties of these pigment-proteins result from the strong excitonic coupling between their bound bacteriochlorophyll a molecules in combination with the large energetic disorder due to slow fluctuations in their structure. As a result, the position and probability of their fluorescence transition delicately depends on the precise realization of the disorder of the set of bound pigments, which is governed by the LH protein dynamics. Analysis of these parameters using time-resolved single-molecule fluorescence spectroscopy thus yields direct access to the protein dynamics. Applying this technique to the LH2 protein from Rhodovulum (Rdv.) sulfidophilum, the structure-and consequently the fluorescence properties-of which depends on pH, allowed us to follow a single protein, pH-induced, reversible, conformational transition. Hence, for the first time, to our knowledge, a protein transition can be visualized through changes in the electronic structure of the intrinsic cofactors, at a level of a single LH protein, which opens a new, to our knowledge, route for understanding the changes in energy landscape that underlie protein function and adaptation to the needs of living organisms.

Coherence and population dynamics of chlorophyll excitations in FCP complex: Two-dimensional spectroscopy study.

Energy transfer processes and coherent phenomena in the fucoxanthin-chlorophyll protein complex, which is responsible for the light harvesting function in marine algae diatoms, were investigated at 77 K by using two-dimensional electronic spectroscopy. Experiments performed on femtosecond and picosecond timescales led to separation of spectral dynamics, witnessing evolutions of coherence and population states of the system in the spectral region of Qy transitions of chlorophylls a and c. Analysis of the coherence dynamics allowed us to identify chlorophyll (Chl) a and fucoxanthin intramolecular vibrations dominating over the first few picoseconds. Closer inspection of the spectral region of the Qy transition of Chl c revealed previously not identified, mutually non-interacting chlorophyll c states participating in femtosecond or picosecond energy transfer to the Chl a molecules. Consideration of separated coherent and incoherent dynamics allowed us to hypothesize the vibrations-assisted coherent energy transfer between Chl c and Chl a and the overall spatial arrangement of chlorophyll molecules.

PHARMACOKINETIC PROPERTIES OF A SINGLE ADMINISTRATION OF ORAL GABAPENTIN IN THE GREAT HORNED OWL (BUBO VIRGINIANUS).

Gabapentin (1-[aminomethyl] cyclohexane acetic acid) is a γ-aminobutyric acid analogue that has been shown to be efficacious for neuropathic pain control in humans. Plasma gabapentin concentrations >2 µg/ml are considered effective in treating epilepsy in humans and are suggested to provide analgesia for neuropathic pain. This study investigated the pharmacokinetics of a single oral dose of gabapentin suspension (11 mg/kg) in great horned owls ( Bubo virginianus ). Plasma gabapentin concentrations were determined in six healthy birds for 48 hr using high-performance liquid chromatography with mass spectrometric detection. Plasma gabapentin concentrations were estimated by noncompartmental pharmacokinetic analysis. The harmonic mean (±SD) maximum concentration (Cmax), time to maximum concentration (Tmax), and elimination half-life (tv2λZ) for gabapentin (11 mg/kg) were 6.17±0.83 µg/ml, 51.43±5.66 min, and 264.60±69.35 min, respectively. In this study, plasma gabapentin concentrations were maintained above 2 µg/ml for 528 min (8.8 hr), suggesting that gabapentin administered orally every 8 hr may be appropriate in great horned owls.

Light-dependent conformational change of neoxanthin in a siphonous green alga, Codium intricatum, revealed by Raman spectroscopy.

Siphonous green algae, a type of deep-sea green algae, appear olive drab and utilize blue-green light for photosynthesis. A siphonous green alga, Codium (C.) intricatum, was isolated from Okinawa prefecture in Japan, and a clonal algal culture in filamentous form was established. The major light-harvesting antenna was analogous to the trimeric LHCII found in higher plants, but the C. intricatum complex contained an unusual carbonyl carotenoid siphonaxanthin. Culture conditions were optimized to achieve high siphonaxanthin content in intact lyophilized filamentous bodies. Interestingly, the carotenoid composition was different when cultured under high irradiance: all-trans neoxanthin was accumulated in addition to the normal 9'-cis form in whole cell extract. Resonance Raman spectra of intact filamentous bodies, cultured under high- and low-light conditions, confirmed the accumulation of all-trans neoxanthin under high irradiance conditions. A plausible function of the presence of all-trans neoxanthin will be discussed in relation to the regulation against high light stress.

Functional organization of 3D plant thylakoid membranes as seen by high resolution microscopy.

In the field of photosynthesis, only a limited number of approaches of super-resolution fluorescence microscopy can be used, as the functional architecture of the thylakoid membrane in chloroplasts is probed through the natural fluorescence of chlorophyll molecules. In this work, we have used a custom-built fluorescence microscopy method called Single Pixel Reconstruction Imaging (SPiRI) that yields a 1.4 gain in lateral and axial resolution relative to confocal fluorescence microscopy, to obtain 2D images and 3D-reconstucted volumes of isolated chloroplasts, obtained from pea (Pisum sativum), spinach (Spinacia oleracea) and Arabidopsis thaliana. In agreement with previous studies, SPiRI images exhibit larger thylakoid grana diameters when extracted from plants under low-light regimes. The three-dimensional thylakoid architecture, revealing the complete network of the thylakoid membrane in intact, non-chemically-fixed chloroplasts can be visualized from the volume reconstructions obtained at high resolution. From such reconstructions, the stromal connections between each granum can be determined and the fluorescence intensity in the stromal lamellae compared to those of neighboring grana.

The light intensity under which cells are grown controls the type of peripheral light-harvesting complexes that are assembled in a purple photosynthetic bacterium.

The differing composition of LH2 (peripheral light-harvesting) complexes present in Rhodopseudomonas palustris 2.1.6 have been investigated when cells are grown under progressively decreasing light intensity. Detailed analysis of their absorption spectra reveals that there must be more than two types of LH2 complexes present. Purified HL (high-light) and LL (low-light) LH2 complexes have mixed apoprotein compositions. The HL complexes contain PucAB(a) and PucAB(b) apoproteins. The LL complexes contain PucAB(a), PucAB(d) and PucB(b)-only apoproteins. This mixed apoprotein composition can explain their resonance Raman spectra. Crystallographic studies and molecular sieve chromatography suggest that both the HL and the LL complexes are nonameric. Furthermore, the electron-density maps do not support the existence of an additional Bchl (bacteriochlorophyll) molecule; rather the density is attributed to the N-termini of the α-polypeptide.

Keep Your Mask On: The Benefits of Masking for Behavior and the Contributions of Aging and Disease on Dysfunctional Masking Pathways.

Environmental cues (e.g., light-dark cycle) have an immediate and direct effect on behavior, but these cues are also capable of "masking" the expression of the circadian pacemaker, depending on the type of cue presented, the time-of-day when they are presented, and the temporal niche of the organism. Masking is capable of complementing entrainment, the process by which an organism is synchronized to environmental cues, if the cues are presented at an expected or predictable time-of-day, but masking can also disrupt entrainment if the cues are presented at an inappropriate time-of-day. Therefore, masking is independent of but complementary to the biological circadian pacemaker that resides within the brain (i.e., suprachiasmatic nucleus) when exogenous stimuli are presented at predictable times of day. Importantly, environmental cues are capable of either inducing sleep or wakefulness depending on the organism's temporal niche; therefore, the same presentation of a stimulus can affect behavior quite differently in diurnal vs. nocturnal organisms. There is a growing literature examining the neural mechanisms underlying masking behavior based on the temporal niche of the organism. However, the importance of these mechanisms in governing the daily behaviors of mammals and the possible implications on human health have been gravely overlooked even as modern society enables the manipulation of these environmental cues. Recent publications have demonstrated that the effects of masking weakens significantly with old age resulting in deleterious effects on many behaviors, including sleep and wakefulness. This review will clearly outline the history, definition, and importance of masking, the environmental cues that induce the behavior, the neural mechanisms that drive them, and the possible implications for human health and medicine. New insights about how masking is affected by intrinsically photosensitive retinal ganglion cells, temporal niche, and age will be discussed as each relates to human health. The overarching goals of this review include highlighting the importance of masking in the expression of daily rhythms, elucidating the impact of aging, discussing the relationship between dysfunctional masking behavior and the development of sleep-related disorders, and considering the use of masking as a non-invasive treatment to help treat humans suffering from sleep-related disorders.

Compassionate reappraisal and rumination impact forgiveness, emotion, sleep, and prosocial accountability.

Sufficient sleep quality and quantity are important for biopsychosocial well-being. Correlational research has linked trait forgiveness to better sleep. Prior experimental evidence also demonstrated contrasting effects of offense rumination versus compassionate reappraisal on forgiveness and psychophysiological responses, suggesting the value of testing effects on sleep. The present study assessed 180 participants (90 M, 90 F). First, we replicated an individual difference model of forgiveness, rumination, depressed and anxious affect, and sleep. Second, we conducted a quasi-experiment inducing offense rumination and compassionate reappraisal on two consecutive nights. Compassionate reappraisal (vs. rumination) replicated past research by prompting more empathic, forgiving, positive, and social responses, with less negative emotion including anger. New findings revealed that compassionate reappraisal (vs. rumination) was also associated with faster sleep onset, fewer sleep disturbances, and fewer sleep impairing offense intrusions. The morning after compassionate reappraisal, participants reported less rumination and intrusive impact of the offense, with more hedonic well-being and accountability to others. Compared to rumination, compassionate reappraisal was associated with more empathy and forgiveness, better sleep, well-being, and prosociality.

Molecular adaptation of photoprotection: triplet states in light-harvesting proteins.

The photosynthetic light-harvesting systems of purple bacteria and plants both utilize specific carotenoids as quenchers of the harmful (bacterio)chlorophyll triplet states via triplet-triplet energy transfer. Here, we explore how the binding of carotenoids to the different types of light-harvesting proteins found in plants and purple bacteria provides adaptation in this vital photoprotective function. We show that the creation of the carotenoid triplet states in the light-harvesting complexes may occur without detectable conformational changes, in contrast to that found for carotenoids in solution. However, in plant light-harvesting complexes, the triplet wavefunction is shared between the carotenoids and their adjacent chlorophylls. This is not observed for the antenna proteins of purple bacteria, where the triplet is virtually fully located on the carotenoid molecule. These results explain the faster triplet-triplet transfer times in plant light-harvesting complexes. We show that this molecular mechanism, which spreads the location of the triplet wavefunction through the pigments of plant light-harvesting complexes, results in the absence of any detectable chlorophyll triplet in these complexes upon excitation, and we propose that it emerged as a photoprotective adaptation during the evolution of oxygenic photosynthesis.