Dissipation Of Excess Excitation Energy Research Articles

Genotype-specific nonphotochemical quenching responses to nitrogen deficit are linked to chlorophyll a to b ratios

Non-photochemical quenching (NPQ) protects plants from photodamage caused by excess light energy. Substantial variation in NPQ has been reported among different genotypes of the same species. However, comparatively little is known about how environmental perturbations, including nutrient deficits, impact natural variation in NPQ kinetics. Here, we analyzed a natural variation in NPQ kinetics of a diversity panel of 225 maize (Zea mays L.) genotypes under nitrogen replete and nitrogen deficient field conditions. Individual maize genotypes from a diversity panel exhibited a range of changes in NPQ in response to low nitrogen. Replicated genotypes exhibited consistent responses across two field experiments conducted in different years. At the seedling and pre-flowering stages, a similar portion of the genotypes (∼33%) showed decrease, no-change or increase in NPQ under low nitrogen relative to control. Genotypes with increased NPQ under low nitrogen also showed greater reductions in dry biomass and photosynthesis than genotypes with stable NPQ when exposed to low nitrogen conditions. Maize genotypes where an increase in NPQ was observed under low nitrogen also exhibited a reduction in the ratio of chlorophyll a to chlorophyll b. Our results underline that since thermal dissipation of excess excitation energy measured via NPQ helps to balance the energy absorbed with energy utilized, the NPQ changes are the reflection of broader molecular and biochemical changes which occur under the stresses such as low soil fertility. Here, we have demonstrated that variation in NPQ kinetics resulted from genetic and environmental factors, are not independent of each other. Natural genetic variation controlling plastic responses of NPQ kinetics to environmental perturbation increases the likelihood it will be possible to optimize NPQ kinetics in crop plants for different environments.

Photosystem II monomeric antenna CP26 plays a key role in nonphotochemical quenching in Chlamydomonas.

Thermal dissipation of excess excitation energy, called nonphotochemical quenching (NPQ), is 1 of the main photoprotective mechanisms in oxygenic photosynthetic organisms. Here, we investigated the function of the monomeric photosystem II (PSII) antenna protein CP26 in photoprotection and light harvesting in Chlamydomonas reinhardtii, a model organism for green algae. We used CRISPR/Cas9 genome editing and complementation to generate cp26 knockout mutants (named k6#) that did not negatively affect CP29 accumulation, which differed from previous cp26 mutants, allowing us to compare mutants specifically deprived of CP26, CP29, or both. The absence of CP26 partially affected PSII activity, causing reduced growth at low or medium light but not at high irradiances. However, the main phenotype observed in k6# mutants was a more than 70% reduction of NPQ compared to the wild type (Wt). This phenotype was fully rescued by genetic complementation and complemented strains accumulating different levels of CP26, demonstrating that ∼50% of CP26 content, compared to the Wt, was sufficient to restore the NPQ capacity. Our findings demonstrate a pivotal role for CP26 in NPQ induction, while CP29 is crucial for PSII activity. The genetic engineering of these 2 proteins could be a promising strategy to regulate the photosynthetic efficiency of microalgae under different light regimes.

Exogenous melatonin improved photosynthetic efficiency of photosystem II by reversible phosphorylation of thylakoid proteins in wheat under osmotic stress.

It has been well demonstrated that melatonin plays an important protective role in photosynthesis of plants under various environmental stresses, while the detailed mechanisms by which melatonin protects photosystem II (PSII) under environmental stress are still unclear. In the study, the effects of melatonin on photosynthetic efficiency, energy dissipation, PSII protein composition, and reversible phosphorylation of thylakoid proteins were investigated in wheat plants under osmotic stress. The results showed that osmotic stress significantly reduced pigment content, photochemical efficiency of PSII, oxygen-evolving activity, and dissipation of excess excitation energy, while 25 μM melatonin applications greatly alleviated their decline under osmotic stress. Western blot data of PSII proteins revealed that melatonin upregulated the levels of D1, Lhcb5, Lhcb6, PsbQ, and PsbS proteins in wheat exposed to osmotic stress. In addition, thylakoid membrane proteins were strongly phosphorylated in wheat under osmotic stress with or without melatonin. Furthermore, the results from PSII protein dephosphorylation showed that exogenous melatonin promoted the dephosphorylation of LCHII, CP43, and D1 under osmotic stress. Therefore, our findings suggest that melatonin can provide an effective protection for the photosynthetic apparatus by the regulation of PSII proteins and the reversible phosphorylation of thylakoid proteins under drought stress.

Exogenous paclobutrazol can relieve the low irradiance stress in Capsicum annuum seedlings

Irradiance is one of the main limiting factors affecting the production of pepper (Capsicum annuum L.) in facility production. It is therefore important to measure the growth of pepper seedlings under low irradiance and to understand how to relieve low radiation stress. In this study, pepper seedlings were cultivated under low irradiance and were treated with gibberellic acid (GA3) and paclobutrazol (PP333). Agronomic and physiological characteristics of the pepper seedlings were analyzed. Under low irradiance, the plant height, leaf area, and the chlorophyll (Chl) and malondialdehyde (MDA) content of pepper seedlings were higher than under normal irradiance, while the content of proline, and soluble protein and Chl a/b ratio of pepper seedlings were lower than those under normal irradiance; exogenous GA3 had a similar effect. When PP333 was applied to the seedlings, abnormal growth was mitigated. The effects of exogenous GA3 and PP333 on Chl fluorescence parameters were also analyzed. Under low irradiance, the maximum quantum yield reduction of photosystem (PS II) (Fv/Fm) increased significantly, and the reaction center initiated the corresponding defense mechanism for timely dissipation of excess excitation energy to reduce the damage to the plant. These results showed that spraying gibberellin under normal irradiance conditions had similar effects on pepper seedlings as under low irradiance conditions. Exogenous PP333 relieved the growth reduction by increasing the content of Chl and soluble protein and enhancing some photosynthetic parameters. These results suggest that exogenous PP333 can alleviate the abnormal growth of pepper under low radiation stress.

Melatonin-mediated photosynthetic performance of tomato seedlings under high-temperature stress

Photosynthesis is a fundamental biosynthetic process in plants that can enhance carbon absorption and increase crop productivity. Heat stress severely inhibits photosynthetic efficiency. Melatonin is a bio-stimulator capable of regulating diverse abiotic stress tolerances. However, the underlying mechanisms of melatonin-mediated photosynthesis in plants exposed to heat stress largely remain elucidated. Our results revealed that melatonin treatment (100 μM) in tomato seedlings increased the endogenous melatonin levels and photosynthetic pigment content along with upregulated of their biosynthesis gene expression under high-temperature stress (42 °C for 24 h), whereas heat stress significantly decreased the values of gas exchange parameters. Under heat stress, melatonin boosted CO2 assimilation, i.e., Vc,max (maximum rate of ribulose-1,5-bisphosphate carboxylase, Rubisco), and Jmax (electron transport of Rubisco generation) and also enhanced the Rubisco and FBPase activities, which resulted in upregulated photosynthetic related gene expression. In addition, heat stress greatly reduced the photochemical chemistry of photosystem II (PSII) and photosystem I (PSI), particularly the maximum quantum efficiency of PSII (Fv/Fm) and PSI (Pm). Conversely, melatonin supplementation increased the chlorophyll a fluorescence parameters led to amplifying the electron transport efficiency. Moreover, heat stress decreased the actual PSII efficiency (ΦPSII), electron transport rate (ETR) and photochemical quenching coefficient (qP), while increasing nonphotochemical quenching (NPQ); however, melatonin reversed these values, which helps to fostering the dissipation of excess excitation energy. Taken together, our results provide a concrete insight into the efficacy of melatonin-mediated photosynthesis performance in a high-temperature regime.

Mechanisms shaping the synergism of zeaxanthin and PsbS in photoprotective energy dissipation in the photosynthetic apparatus of plants.

Safe operation of photosynthesis is vital to plants and is ensured by the activity of processes protecting chloroplasts against photo-damage. The harmless dissipation of excess excitation energy is considered to be the primary photoprotective mechanism and is most effective in the combined presence of PsbS protein and zeaxanthin, a xanthophyll accumulated in strong light as a result of the xanthophyll cycle. Here we address the problem of specific molecular mechanisms underlying the synergistic effect of zeaxanthin and PsbS. The experiments were conducted with Arabidopsis thaliana, using wild-type plants, mutants lacking PsbS (npq4), and mutants affected in the xanthophyll cycle (npq1), with the application of molecular spectroscopy and imaging techniques. The results lead to the conclusion that PsbS interferes with the formation of densely packed aggregates of thylakoid membrane proteins, thus allowing easy exchange and incorporation of xanthophyll cycle pigments into such structures. It was found that xanthophylls trapped within supramolecular structures, most likely in the interfacial protein region, determine their photophysical properties. The structures formed in the presence of violaxanthin are characterized by minimized dissipation of excitation energy. In contrast, the structures formed in the presence of zeaxanthin show enhanced excitation quenching, thus protecting the system against photo-damage.

Zeaxanthin, a Molecule for Photoprotection in Many Different Environments.

Conversion of sunlight into photochemistry depends on photoprotective processes that allow safe use of sunlight over a broad range of environmental conditions. This review focuses on the ubiquity of photoprotection associated with a group of interconvertible leaf carotenoids, the xanthophyll cycle. We survey the striking plasticity of this process observed in nature with respect to (1) xanthophyll cycle pool size, (2) degree and speed of interconversion of its components, and (3) flexibility in the association between xanthophyll cycle conversion state and photoprotective dissipation of excess excitation energy. It is concluded that the components of this system can be independently tuned with a high degree of flexibility to produce a fit for different environments with various combinations of light, temperature, and other factors. In addition, the role of genetic variation is apparent from variation in the response of different species growing side-by-side in the same environment. These findings illustrate how field studies can generate insight into the adjustable levers that allow xanthophyll cycle-associated photoprotection to support plant photosynthetic productivity and survival in environments with unique combinations of environmental factors.

A Protein Environment-Modulated Energy Dissipation Channel in LHCII Antenna Complex.

SummaryThe major light-harvesting complex of photosystem II (LHCII) is the main contributor to sunlight energy harvesting in plants. The flexible design of LHCII underlies a photoprotective mechanism whereby this complex switches to a dissipative state in response to high light stress, allowing the rapid dissipation of excess excitation energy (non-photochemical quenching, NPQ). In this work, we locked single LHCII trimers in a quenched conformation after immobilization of the complexes in polyacrylamide gels to impede protein interactions. A comparison of their pigment excited-state dynamics with quenched LHCII aggregates in buffer revealed the presence of a new spectral band at 515 nm arising after chlorophyll excitation. This is suggested to be the signature of a carotenoid excited state, linked to the quenching of chlorophyll singlet excited states. Our data highlight the marked sensitivity of pigment excited-state dynamics in LHCII to structural changes induced by the environment.

Changes in Photo-Protective Energy Dissipation of Photosystem II in Response to Beneficial Bacteria Consortium in Durum Wheat under Drought and Salinity Stresses

The present research aimed at evaluating the harmless dissipation of excess excitation energy by durum wheat (Triticum durum Desf.) leaves in response to the application of a bacterial consortium consisting of four plant growth-promoting bacteria (PGPB). Three pot experiments were carried out under non-stress, drought (at 40% field capacity), and salinity (150 mM NaCl) conditions. The results showed that drought and salinity affected photo-protective energy dissipation of photosystem II (PSII) increasing the rate of non-photochemical chlorophyll fluorescence quenching (NPQ (non-photochemical quenching) and qCN (complete non-photochemical quenching)), as well as decreasing the total quenching of chlorophyll fluorescence (qTQ), total quenching of variable chlorophyll fluorescence (qTV) and the ratio of the quantum yield of actual PSII photochemistry, in light-adapted state to the quantum yield of the constitutive non-regulatory NPQ (PQ rate). Our results also indicated that the PGPB inoculants can mitigate the adverse impacts of stresses on leaves, especially the saline one, in comparison with the non-fertilized (control) treatment, by increasing the fraction of light absorbed by the PSII antenna, PQ ratio, qTQ, and qTV. In the light of findings, our beneficial bacterial strains showed the potential in reducing reliance on traditional chemical fertilizers, in particular in saline soil, by improving the grain yield and regulating the amount of excitation energy.

Salicylic Acid Protects Photosystem II by Alleviating Photoinhibition in Arabidopsis thaliana under High Light.

Salicylic acid (SA) is considered to play an important role in plant responses to environmental stresses. However, the detailed protective mechanisms in photosynthesis are still unclear. We therefore explored the protective roles of SA in photosystem II (PSII) in Arabidopsis thaliana under high light. The results demonstrated that 3 h of high light exposure resulted in a decline in photochemical efficiency and the dissipation of excess excitation energy. However, SA application significantly improved the photosynthetic capacity and the dissipation of excitation energy under high light. Western blot analysis revealed that SA application alleviated the decrease in the levels of D1 and D2 protein and increased the amount of Lhcb5 and PsbS protein under high light. Results from photoinhibition highlighted that SA application could accelerate the repair of D1 protein. Furthermore, the phosphorylated levels of D1 and D2 proteins were significantly increased under high light in the presence of SA. In addition, we found that SA application significantly alleviated the disassembly of PSII-LHCII super complexes and LHCII under high light for 3 h. Overall, our findings demonstrated that SA may efficiently alleviate photoinhibition and improve photoprotection by dissipating excess excitation energy, enhancing the phosphorylation of PSII reaction center proteins, and preventing the disassembly of PSII super complexes.

A Flexible LHCII Structure Allows for Fine-Tuning of Excitation Energy Dissipation

The major light-harvesting complex of photosystem II (LHCII) is the main contributor to sunlight energy harvesting in plants. The flexible design of LHCII underlies a photoprotective mechanism whereby this complex switches to a dissipative state in response to high light stress, allowing the rapid dissipation of excess excitation energy (NPQ). In this work, we stabilized the quenched conformation of single LHCII trimers after immobilization of the complexes in polyacrylamide gels to impede protein interactions. A comparison of their pigment excited-state dynamics with quenched LHCII aggregates in buffer revealed the presence of a new spectral band at 515 nm arising after chlorophyll excitation. This is suggested to be the signature of a carotenoid excited state, linked to the quenching of chlorophyll singlet excited states. Our data highlight the marked sensitivity of pigment excited-state dynamics in LHCII to structural changes induced by the environment.

Photorespiration is the major alternative electron sink under high light in alpine evergreen sclerophyllous Rhododendron species

Owing to the high leaf mass per area, alpine evergreen sclerophyllous Rhododendron have low values of mesophyll conductance (gm). The resulting low chloroplast CO2 concentration aggravates photorespiration, which requires a higher ATP/NADPH ratio. However, the significance of photorespiration and underlying mechanisms of energy balance in these species are little known. In this study, eight alpine evergreen sclerophyllous Rhododendron species grown in a common garden were tested for their gm, electron flow to photorespiration, and energy balancing. Under saturating light, gm was the most limiting factor for net photosynthesis (AN) in all species, and the species differences in AN were primarily driven by gm rather than stomatal conductance. The total electron flow through photosystem II (ETRII) nearly equaled the electron transport required for Rubisco carboxylation and oxygenation. Furthermore, blocking electron flow to photosystem I with appropriate inhibitors showed that electron flow to plastic terminal oxidase was not observed. As a result, these studied species showed little alternative electron flow mediated by water-water cycle or plastic terminal oxidase. By comparison, the ratio of electron transport consumed by photorespiration to ETRII (JPR/ETRII), ranging from 43%∼55%, was negatively correlated to AN and gm. Furthermore, the increased ATP production required by enhanced photorespiration was regulated by cyclic electron flow around photosystem I. These results indicate that photorespiration is the major electron sink for dissipation of excess excitation energy in the alpine evergreen sclerophyllous Rhododendron species. The coordination of gm, photorespiration and cyclic electron flow is important for sustaining leaf photosynthesis.

The positive regulation of putrescine on light-harvesting complex II and excitation energy dissipation in salt-stressed cucumber seedlings

Non-photochemical quenching (NPQ) is an important mechanism employed by higher plants to protect themselves against the adverse effects of excess excitation energy. The regulatory mechanism of exogenous putrescine (Put) on the status of light-harvesting complex II (LHCII) and its dissipation of excess excitation energy in cucumber seedlings (Cucumis sativus L.) were investigated under salt stress in this study. Put significantly increased the maximum quantum photochemical efficiency in photosystem II (Fv/Fm), NPQ, and the actual yield of PSII (ΦPSII) under salt stress. Quantum yield of regulated energy dissipation in PSII (ΦNPQ) significantly increased and non-regulated energy dissipation (ΦNO) decreased when was sprayed on stressed plants. Put increased the expression levels of D1, D2, Lhcb4 and PsbS, thereby minimizing degradation to the monomeric and trimeric states of LHCII in the thylakoid membranes of salt-stressed cucumber leaves. In addition, Put up-regulated the activity of transglutaminase (TGase) and significantly increased the levels of total endogenous polyamines in monomeric and trimeric LHCII under salt stress. These results suggest that Put could play a crucial role in dissipating excess excitation energy by regulating endogenously increased polyamines to alter the adaptation of LHCII, which are most probably involved in an improvement of photochemical efficiency in salt stressed-cucumber seedlings.

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Non-photochemical quenching (NPQ) is an important mechanism employed by higher plants to protect themselves against the adverse effects of excess excitation energy. The regulatory mechanism of exogenous putrescine (Put) on the status of light-harvesting complex II (LHCII) and its dissipation of excess excitation energy in cucumber seedlings (Cucumis sativus L.) were investigated under salt stress in this study. Put significantly increased the maximum quantum photochemical efficiency in photosystem II (Fv/Fm), NPQ, and the actual yield of PSII (ΦPSII) under salt stress. Quantum yield of regulated energy dissipation in PSII (ΦNPQ) significantly increased and non-regulated energy dissipation (ΦNO) decreased when was sprayed on stressed plants. Put increased the expression levels of D1, D2, Lhcb4 and PsbS, thereby minimizing degradation to the monomeric and trimeric states of LHCII in the thylakoid membranes of salt-stressed cucumber leaves. In addition, Put up-regulated the activity of transglutaminase (TGase) and significantly increased the levels of total endogenous polyamines in monomeric and trimeric LHCII under salt stress. These results suggest that Put could play a crucial role in dissipating excess excitation energy by regulating endogenously increased polyamines to alter the adaptation of LHCII, which are most probably involved in an improvement of photochemical efficiency in salt stressed-cucumber seedlings.

Light Energy Partitioning and Photoprotection in an Exotic Species (Salix Psammophila) Grown in a Semi-Arid Area of Northwestern China

Thermal dissipation of excess excitation energy is an important photoprotective mechanism that plants have evolved to cope with surplus illumination. However, light-energy-partitioning dynamics in an exotic sand-dune willow (Salix psammophila) commonly used in restoring and/or stabilizing sand lands in northwestern China is largely unknown. In this study, chlorophyll fluorescence (ChlF) of photosystem II (PSII) was continuously monitored in situ in Salix psammophila to investigate plant acclimation processes driven by excessive solar radiation and extreme air temperatures (Ta). As part of a heat-regulation mechanism, energy partitioning is shown to vary with prevailing environmental conditions. In this investigation, energy absorbed during periods of moderate photosynthetically active radiation (PAR < 1200 μmoL·m−2·s−1) was largely allocated towards photochemistry (ΦPSII) with nominal amounts to thermal dissipation through reversible thermal dissipation (ΦNPQr). In extremely high solar radiation (PAR > 1500 μmoL·m−2·s−1) or in a cold temperature (Ta < 0 °C), more energy was dissipated by way of non-regulated thermal energy (Φf,D) and sustained thermal dissipation (ΦNPQs), leading to non-reversible photoinhibition or photodamage. This was mainly as a result of the low utilization and high absorption of light energy by PSII under cold conditions and physiologically-induced vulnerability. It was concluded that Salix psammophila had a clear tolerance to high temperatures and moderate solar radiation, but tended to be more vulnerable to high solar radiation and cold temperature. Based on species sensitivity to extreme environmental conditions, practical application and extension of Salix psammophila for land-restoration purposes should be approached cautiously, especially in high-latitude or high-altitude desert ecosystems commonly affected by events of high solar radiation and cold temperature.

Dynamic feedback of the photosystem II reaction centre on photoprotection in plants.

Photosystem II of higher plants is protected against light damage by thermal dissipation of excess excitation energy, a process that can be monitored through non-photochemical quenching of chlorophyll fluorescence. When the light intensity is lowered, non-photochemical quenching largely disappears on a time scale ranging from tens of seconds to many minutes. With the use of picosecond fluorescence spectroscopy, we demonstrate that one of the underlying mechanisms is only functional when the reaction centre of photosystem II is closed, that is when electron transfer is blocked and the risk of photodamage is high. This is accompanied by the appearance of a long-wavelength fluorescence band. As soon as the reaction centre reopens, this quenching, together with the long-wavelength fluorescence, disappears instantaneously. This allows plants to maintain a high level of photosynthetic efficiency even in dangerous high-light conditions.

Variation in short-term and long-term responses of photosynthesis and isoprenoid-mediated photoprotection to soil water availability in four Douglas-fir provenances

For long-lived forest tree species, the understanding of intraspecific variation among populations and their response to water availability can reveal their ability to cope with and adapt to climate change. Dissipation of excess excitation energy, mediated by photoprotective isoprenoids, is an important defense mechanism against drought and high light when photosynthesis is hampered. We used 50-year-old Douglas-fir trees of four provenances at two common garden experiments to characterize provenance-specific variation in photosynthesis and photoprotective mechanisms mediated by essential and non-essential isoprenoids in response to soil water availability and solar radiation. All provenances revealed uniform photoprotective responses to high solar radiation, including increased de-epoxidation of photoprotective xanthophyll cycle pigments and enhanced emission of volatile monoterpenes. In contrast, we observed differences between provenances in response to drought, where provenances sustaining higher CO2 assimilation rates also revealed increased water-use efficiency, carotenoid-chlorophyll ratios, pools of xanthophyll cycle pigments, β-carotene and stored monoterpenes. Our results demonstrate that local adaptation to contrasting habitats affected chlorophyll-carotenoid ratios, pool sizes of photoprotective xanthophylls, β-carotene, and stored volatile isoprenoids. We conclude that intraspecific variation in isoprenoid-mediated photoprotective mechanisms contributes to the adaptive potential of Douglas-fir provenances to climate change.

Artificial Photosynthetic Reaction Center Exhibiting Acid-Responsive Regulation of Photoinduced Charge Separation.

Nonphotochemical quenching (NPQ) is a photoprotective regulatory mechanism employed by many photosynthetic organisms to dynamically modulate energy flow within the photosynthetic apparatus in response to fluctuating light conditions. Activated by decreases in lumen pH produced during periods of high photon flux, NPQ induces rapid thermal dissipation of excess excitation energy. As a result, the rate of charge separation (CS) decreases, thereby limiting the accumulation of potentially deleterious reactive intermediates and byproducts. Herein, a molecular triad that functionally mimics the effects of NPQ associated with an artificial photosynthetic reaction center is described. Steady-state absorption and emission, time-resolved fluorescence, and transient absorption spectroscopies have been used to demonstrate a 1 order of magnitude reduction in the CS quantum yield via reversible protonation of an excited-state-quenching molecular switch moiety. As in the natural system, the populations of unquenched and quenched states and therefore the overall yields of CS were found to be dependent on acid concentration.

Dissipation of excess excitation energy of the needle leaves in Pinus trees during cold winters.

Photooxidative damage to the needle leaves of evergreen trees results from the absorption of excess excitation energy. Efficient dissipation of this energy is essential to prevent photodamage. In this study, we determined the fluorescence transients, absorption spectra, chlorophyll contents, chlorophyll a/b ratios, and relative membrane permeabilities of needle leaves of Pinus koraiensis, Pinus tabulaeformis, and Pinus armandi in both cold winter and summer. We observed a dramatic decrease in the maximum fluorescence (F m) and substantial absorption of light energy in winter leaves of all three species. The F m decline was not correlated with a decrease in light absorption or with changes in chlorophyll content and chlorophyll a/b ratio. The results suggested that the winter leaves dissipated a large amount of excess energy as heat. Because the cold winter leaves had lost normal physiological function, the heat dissipation depended solely on changes in the photosystem II supercomplex rather than the xanthophyll cycle. These findings imply that more attention should be paid to heat dissipation via changes in the photosystem complex structure during the growing season.

The Type II NADPH Dehydrogenase Facilitates Cyclic Electron Flow, Energy-Dependent Quenching, and Chlororespiratory Metabolism during Acclimation of Chlamydomonas reinhardtii to Nitrogen Deprivation.

When photosynthetic organisms are deprived of nitrogen (N), the capacity to grow and assimilate carbon becomes limited, causing a decrease in the productive use of absorbed light energy and likely a rise in the cellular reduction state. Although there is a scarcity of N in many terrestrial and aquatic environments, a mechanistic understanding of how photosynthesis adjusts to low-N conditions and the enzymes/activities integral to these adjustments have not been described. In this work, we use biochemical and biophysical analyses of photoautotrophically grown wild-type and mutant strains of Chlamydomonas reinhardtii to determine the integration of electron transport pathways critical for maintaining active photosynthetic complexes even after exposure of cells to N deprivation for 3 d. Key to acclimation is the type II NADPH dehydrogenase, NDA2, which drives cyclic electron flow (CEF), chlororespiration, and the generation of an H(+) gradient across the thylakoid membranes. N deprivation elicited a doubling of the rate of NDA2-dependent CEF, with little contribution from PGR5/PGRL1-dependent CEF The H(+) gradient generated by CEF is essential to sustain nonphotochemical quenching, while an increase in the level of reduced plastoquinone would promote a state transition; both are necessary to down-regulate photosystem II activity. Moreover, stimulation of NDA2-dependent chlororespiration affords additional relief from the elevated reduction state associated with N deprivation through plastid terminal oxidase-dependent water synthesis. Overall, rerouting electrons through the NDA2 catalytic hub in response to photoautotrophic N deprivation sustains cell viability while promoting the dissipation of excess excitation energy through quenching and chlororespiratory processes.