RESUMEN
Neurulation is a crucial process in the formation of the central nervous system (CNS), which begins with the folding and fusion of the neural plate, leading to the generation of the neural tube and subsequent development of the brain and spinal cord. Environmental and genetic factors that interfere with the neurulation process promote neural tube defects (NTDs). Connexins (Cxs) are transmembrane proteins that form gap junctions (GJs) and hemichannels (HCs) in vertebrates, allowing cell-cell (GJ) or paracrine (HCs) communication through the release of ATP, glutamate, and NAD+; regulating processes such as cell migration and synaptic transmission. Changes in the state of phosphorylation and/or the intracellular redox potential activate the opening of HCs in different cell types. Cxs such as Cx43 and Cx32 have been associated with proliferation and migration at different stages of CNS development. Here, using molecular and cellular biology techniques (permeability), we demonstrate the expression and functionality of HCs-Cxs, including Cx46 and Cx32, which are associated with the release of ATP during the neurulation process in Xenopus laevis. Furthermore, applications of FGF2 and/or changes in intracellular redox potentials (DTT), well known HCs-Cxs modulators, transiently regulated the ATP release in our model. Importantly, the blockade of HCs-Cxs by carbenoxolone (CBX) and enoxolone (ENX) reduced ATP release with a concomitant formation of NTDs. We propose two possible and highly conserved binding sites (N and E) in Cx46 that may mediate the pharmacological effect of CBX and ENX on the formation of NTDs. In summary, our results highlight the importance of ATP release mediated by HCs-Cxs during neurulation.
Asunto(s)
Conexinas , Defectos del Tubo Neural , Animales , Conexinas/metabolismo , Neurulación , Uniones Comunicantes/metabolismo , Tubo Neural/metabolismo , Defectos del Tubo Neural/metabolismo , Adenosina Trifosfato/metabolismoRESUMEN
BACKGROUND: Early seed germination and a functional root system development during establishment are crucial attributes contributing to nutrient competence under marginal nutrient soil conditions. Chenopodium quinoa Willd (Chenopodiaceae) is a rustic crop, able to grow in marginal areas. Altiplano and Coastal/Lowlands are two representative zones of quinoa cultivation in South America with contrasting soil fertility and edaphoclimatic conditions. In the present work, we hypothesize that the ecotypes of Quinoa from Altiplano (landrace Socaire) and from Coastal/Lowland (landrace Faro) have developed differential adaptive responses in order to survive under conditions of low availability of N in their respective climatic zones of Altiplano and Lowlands. In order to understand intrinsic differences for N competence between landraces, seed metabolite profile and germinative capacity were studied. Additionally, in order to elucidate the mechanisms of N uptake and assimilation at limiting N conditions during establishment, germinated seeds of both landraces were grown at either sufficient nitrate (HN) or low nitrate (LN) supply. We studied the photosynthetic performance, protein storage, root morphometrical parameters, activity and expression of N-assimilating enzymes, and the expression of nitrate transporters of roots in plants submitted to the different treatments. RESULTS: Seeds from Socaire landrace presented higher content of free N-related metabolites and faster seed germination rate compared to Faro landrace. Seedlings of both ecotypes presented similar physiological performance at HN supply, but at LN supply their differences were exalted. At LN, Socaire plants showed an increased root biomass (including a higher number and total length of lateral roots), a differential regulation of a nitrate transporter (a NPF6.3-like homologue) belonging to the Low Affinity Transport System (LATS), and an upregulation of a nitrate transporter (a NRT2.1-like homologue) belonging to the High Affinity nitrate Transport System (HATS) compared to Faro. These responses as a whole could be linked to a higher amount of stored proteins in leaves, associated to an enhanced photochemical performance in Altiplano plants, in comparison to Lowland quinoa plants. CONCLUSIONS: These differential characteristics of Socaire over Faro plants could involve an adaptation to enhanced nitrate uptake under the brutal unfavorable climate conditions of Altiplano.
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Chenopodium quinoa/metabolismo , Nitrógeno/metabolismo , Plantones/metabolismo , Semillas/metabolismo , Proteínas de Transporte de Anión/genética , Proteínas de Transporte de Anión/metabolismo , Chenopodium quinoa/genética , Chenopodium quinoa/crecimiento & desarrollo , Chile , Ecotipo , Regulación de la Expresión Génica de las Plantas , Germinación , Glutamato-Amoníaco Ligasa/metabolismo , Nitrato-Reductasa/metabolismo , Transportadores de Nitrato , Nitratos/metabolismo , Filogenia , Proteínas de Plantas/genética , Proteínas de Plantas/metabolismo , Raíces de Plantas/metabolismo , Plantones/crecimiento & desarrollo , Semillas/fisiologíaRESUMEN
The interaction of the alternative oxidase (AOX) pathway with nutrient metabolism is important for understanding how respiration modulates ATP synthesis and carbon economy in plants under nutrient deficiency. Although AOX activity reduces the energy yield of respiration, this enzymatic activity is upregulated under stress conditions to maintain the functioning of primary metabolism. The in vivo metabolic regulation of AOX activity by phosphorus (P) and nitrogen (N) and during plant symbioses with Arbuscular mycorrhizal fungi (AMF) and Rhizobium bacteria is still not fully understood. We highlight several findings and open questions concerning the in vivo regulation of AOX activity and its impact on plant metabolism during P deficiency and symbiosis with AMF. We also highlight the need for the identification of which metabolic regulatory factors of AOX activity are related to N availability and nitrogen-fixing legume-rhizobia symbiosis in order to improve our understanding of N assimilation and biological nitrogen fixation.
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Proteínas Mitocondriales/metabolismo , Micorrizas/fisiología , Oxidorreductasas/metabolismo , Proteínas de Plantas/metabolismo , Plantas/microbiología , Rhizobium/fisiología , Adenosina Trifosfato/metabolismo , Carbono/metabolismo , Regulación de la Expresión Génica de las Plantas , Nitrógeno/metabolismo , Fósforo/metabolismo , Plantas/metabolismo , Transducción de Señal , Estrés Fisiológico , SimbiosisRESUMEN
Leaf respiration and photosynthesis will respond differently to an increase in temperature during night, which can be more relevant in sensitive ecosystems such as Antarctica. We postulate that the plant species able to colonize the Antarctic Peninsula - Colobanthus quitensis (Kunth) Bartl. and Deschampsia antarctica Desv. - are able to acclimate their foliar respiration and to maintain photosynthesis under nocturnal warming to sustain a positive foliar carbon balance. We conducted a laboratory experiment to evaluate the effect of time of day (day and night) and nocturnal warming on dark respiration. Short (E0 and Q10 ) and long-term acclimation of respiration, leaf carbohydrates, photosynthesis (Asat ) and foliar carbon balance (R/A) were evaluated. The results suggest that the two species have differential thermal acclimation respiration, where D. antarctica showed more thermosensitivity to short-term changes in temperature than C. quitensis. Experimental nocturnal warming affected respiration at daytime differentially between the two species, with a significant increase of R10 and Asat in D. antarctica, while no changes on respiration were observed in C. quitensis. Long thermal treatments of the plants indicated that nocturnal but not diurnal respiration could acclimate in both species, and to a greater extent in C. quitensis. Non-structural carbohydrates were related with respiration in C. quitensis but not in D. antarctica, suggesting that respiration in the former species is likely controlled by total soluble sugars and starch during day and night, respectively. Finally, foliar carbon balance was differentially improved under warming conditions in Antarctic plants by different mechanisms, with C. quitensis deploying respiratory acclimation, while D. antarctica increased its Asat.
Asunto(s)
Aclimatación , Carbono/metabolismo , Caryophyllaceae/fisiología , Poaceae/fisiología , Oscuridad , Ecosistema , Fotosíntesis/fisiología , Hojas de la Planta/fisiología , TemperaturaRESUMEN
MAIN CONCLUSION: Water deficit stress followed by re-watering during grain filling resulted in the induction of the ornithine pathway and in changes in Quinoa grain quality. The genetic diversity of Chenopodium quinoa Willd. (Quinoa) is accompanied by an outstanding environmental adaptability and high nutritional properties of the grains. However, little is known about the biochemical and physiological mechanisms associated with the abiotic stress tolerance of Quinoa. Here, we characterized carbon and nitrogen metabolic changes in Quinoa leaves and grains in response to water deficit stress analyzing their impact on the grain quality of two lowland ecotypes (Faro and BO78). Differences in the stress recovery response were found between genotypes including changes in the activity of nitrogen assimilation-associated enzymes that resulted in differences in grain quality. Both genotypes showed a common strategy to overcome water stress including the stress-induced synthesis of reactive oxygen species scavengers and osmolytes. Particularly, water deficit stress induced the stimulation of the ornithine and raffinose pathways. Our results would suggest that the regulation of C- and N partitioning in Quinoa during grain filling could be used for the improvement of the grain quality without altering grain yields.
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Carbono/metabolismo , Chenopodium quinoa/fisiología , Nitrógeno/metabolismo , Agua/fisiología , Chenopodium quinoa/genética , Deshidratación , Grano Comestible/genética , Grano Comestible/fisiología , Variación Genética , Hojas de la Planta/genética , Hojas de la Planta/fisiología , Estrés FisiológicoRESUMEN
The Antarctic Peninsula has rapidly warmed up in past decades, and global warming has exhibited an asymmetric trend; therefore, it is interesting to understand whether nocturnal or diurnal warming is the most relevant for plant cold deacclimation. This study aimed to evaluate the effect of diurnal and nocturnal warming on Antarctic vascular plant's freezing resistance under laboratory conditions. This was studied by measuring the lethal temperature for 50% of tissue (LT50), ice nucleation temperature (INT), and freezing point (FP) on Deschampsia antarctica and Colobanthus quitensis plants. Additionally, soluble carbohydrates content and dehydrin levels were analyzed during nocturnal and diurnal temperatures increase. Nocturnal warming led to a 7 °C increase in the LT50 of D. antarctica and reduced dehydrin-like peptide expression. Meanwhile, C. quitensis warmed plants reduce their LT50 to about 3.6 °C. Both species reduce their sucrose content by more than 28% in warming treatments. Therefore, nocturnal warming leads to cold deacclimation in both plant species, while C. quitensis plants are also cold-deacclimated upon warm days. This suggests that even when the remaining freezing resistance of both species allows them to tolerate summer freezing events, C. quitensis can reach its boundaries of freezing vulnerability in the near future if warming in the Antarctic Peninsula progress.
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Desert shrubs are keystone species for plant diversity and ecosystem function. Atriplex clivicola and Atriplex deserticola (Amaranthaceae) are native shrubs from the Atacama Desert that show contrasting altitudinal distribution (A. clivicola: 0-700 m.a.s.l.; A. deserticola: 1500-3000 m.a.s.l.). Both species possess a C4 photosynthetic pathway and Kranz anatomy, traits adaptive to high temperatures. Historical records and projections for the near future show trends in increasing air temperature and frequency of heat wave events in these species' habitats. Besides sharing a C4 pathway, it is not clear how their leaf-level physiological traits associated with photosynthesis and water relations respond to heat stress. We studied their physiological traits (gas exchange, chlorophyll fluorescence, water status) before and after a simulated heat wave (HW). Both species enhanced their intrinsic water use efficiency after HW but via different mechanisms. A. clivicola, which has a higher LMA than A. deserticola, enhances water saving by closing stomata and maintaining RWC (%) and leaf Ψmd potential at similar values to those measured before HW. After HW, A. deserticola showed an increase of Amax without concurrent changes in gs and a significant reduction of RWC and Ψmd. A. deserticola showed higher values of Chla fluorescence after HW. Thus, under heat stress, A. clivicola maximizes water saving, whilst A. deserticola enhances its photosynthetic performance. These contrasting (eco)physiological strategies are consistent with the adaptation of each species to their local environmental conditions at different altitudes.
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Chenopodium quinoa Willd. is a native species that originated in the High Andes plateau (Altiplano) and its cultivation spread out to the south of Chile. Because of the different edaphoclimatic characteristics of both regions, soils from Altiplano accumulated higher levels of nitrate (NO3-) than in the south of Chile, where soils favor ammonium (NH4 +) accumulation. To elucidate whether C. quinoa ecotypes differ in several physiological and biochemical parameters related to their capacity to assimilate NO3- and NH4 +, juvenile plants of Socaire (from Altiplano) and Faro (from Lowland/South of Chile) were grown under different sources of N (NO3- or NH4 +). Measurements of photosynthesis and foliar oxygen-isotope fractionation were carried out, together with biochemical analyses, as proxies for the analysis of plant performance or sensitivity to NH4 +. Overall, while NH4 + reduced the growth of Socaire, it induced higher biomass productivity and increased protein synthesis, oxygen consumption, and cytochrome oxidase activity in Faro. We discussed that ATP yield from respiration in Faro could promote protein production from assimilated NH4 + to benefit its growth. The characterization of this differential sensitivity of both quinoa ecotypes for NH4 + contributes to a better understanding of nutritional aspects driving plant primary productivity.
RESUMEN
BACKGROUND: Ecotypes of Colobanthus quitensis Kunt Bartl (Cariophyllaceae) from Andes Mountains and Maritime Antarctic grow under contrasting photoinhibitory conditions, reaching differential cold tolerance upon cold acclimation. Photoinhibition depends on the extent of photodamage and recovery capability. We propose that cold acclimation increases resistance to low-temperature-induced photoinhibition, limiting photodamage and promoting recovery under cold. Therefore, the Antarctic ecotype (cold hardiest) should be less photoinhibited and have better recovery from low-temperature-induced photoinhibition than the Andean ecotype. Both ecotypes were exposed to cold induced photoinhibitory treatment (PhT). Photoinhibition and recovery of photosystem II (PSII) was followed by fluorescence, CO2 exchange, and immunoblotting analyses. RESULTS: The same reduction (25%) in maximum PSII efficiency (Fv/Fm) was observed in both cold-acclimated (CA) and non-acclimated (NA) plants under PhT. A full recovery was observed in CA plants of both ecotypes under dark conditions, but CA Antarctic plants recover faster than the Andean ecotype.Under PhT, CA plants maintain their quantum yield of PSII, while NA plants reduced it strongly (50% and 73% for Andean and Antarctic plants respectively). Cold acclimation induced the maintenance of PsaA and Cyt b6/f and reduced a 41% the excitation pressure in Antarctic plants, exhibiting the lowest level under PhT. xCold acclimation decreased significantly NPQs in both ecotypes, and reduced chlorophylls and D1 degradation in Andean plants under PhT.NA and CA plants were able to fully restore their normal photosynthesis, while CA Antarctic plants reached 50% higher photosynthetic rates after recovery, which was associated to electron fluxes maintenance under photoinhibitory conditions. CONCLUSIONS: Cold acclimation has a greater importance on the recovery process than on limiting photodamage. Cold acclimation determined the kinetic and extent of recovery process under darkness in both C. quitensis ecotypes. The greater recovery of PSII at low temperature in the Antarctic ecotype was related with its ability to maintain PsaA, Cyt b6/f and D1 protein after photoinhibitory conditions. This is probably due to either a higher stability of these polypeptides or to the maintenance of their turnover upon cold acclimation. In both cases, it is associated to the maintenance of electron drainage from the intersystem pool, which maintains QA more oxidized and may allow the synthesis of ATP and NADPH necessaries for the regeneration of ribulose 1,5-bisphosphate in the Calvin Cycle. This could be a key factor for C. quitensis success under the harsh conditions and the short growing period in the Maritime Antarctic.
Asunto(s)
Aclimatación , Caryophyllaceae/química , Frío , Oscuridad , Fotosíntesis , Complejo de Proteína del Fotosistema II/química , Regiones Antárticas , Dióxido de Carbono/química , Caryophyllaceae/fisiología , Chile , Clorofila/química , Citocromos b6/química , Ecotipo , Fluorescencia , Especificidad de la Especie , Almidón/química , Proteínas de las Membranas de los Tilacoides/químicaRESUMEN
Projected rises in atmospheric CO2 concentration and minimum night-time temperatures may have important effects on plant carbon metabolism altering the carbon balance of the only two vascular plant species in the Antarctic Peninsula. We assessed the effect of nocturnal warming (8/5 °C vs. 8/8 °C day/night) and CO2 concentrations (400 ppm and 750 ppm) on gas exchange, non-structural carbohydrates, two respiratory-related enzymes, and mitochondrial size and number in two species of vascular plants. In Colobanthus quitensis, light-saturated photosynthesis measured at 400 ppm was reduced when plants were grown in the elevated CO2 or in the nocturnal warming treatments. Growth in elevated CO2 reduced stomatal conductance but nocturnal warming did not. The short-term sensitivity of respiration, relative protein abundance, and mitochondrial traits were not responsive to either treatment in this species. Moreover, some acclimation to nocturnal warming at ambient CO2 was observed. Altogether, these responses in C. quitensis led to an increase in the respiration-assimilation ratio in plants grown in elevated CO2. The response of Deschampsia antarctica to the experimental treatments was quite distinct. Photosynthesis was not affected by either treatment; however, respiration acclimated to temperature in the elevated CO2 treatment. The observed short-term changes in thermal sensitivity indicate type I acclimation of respiration. Growth in elevated CO2 and nocturnal warming resulted in a reduction in mitochondrial numbers and an increase in mitochondrial size in D. antarctica. Overall, our results suggest that with climate change D. antarctica could be more successful than C. quitensis, due to its ability to make metabolic adjustments to maintain its carbon balance.
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"Memory imprint" refers to the process when prior exposure to stress prepares the plant for subsequent stress episodes. Seed priming is a strategy to change the performance of seedlings to cope with stress; however, mechanisms associated with the metabolic response are fragmentary. Salinity is one of the major abiotic stresses that affect crop production in arid and semiarid areas. Chenopodium quinoa Willd. (Amaranthaceae) is a promising crop to sustain food security and possesses a wide genetic diversity of salinity tolerance. To elucidate if the metabolic memory induced by seed halo-priming (HP) differs among contrasting saline tolerance plants, seeds of two ecotypes of Quinoa (Socaire from Atacama Salar, and BO78 from Chilean Coastal/lowlands) were treated with a saline solution and then germinated and grown under different saline conditions. The seed HP showed a more positive impact on the sensitive ecotype during germination and promoted changes in the metabolomic profile in both ecotypes, including a reduction in carbohydrates (starch) and organic acids (citric and succinic acid), and an increase in antioxidants (ascorbic acid and α-tocopherol) and related metabolites. These changes were linked to a further reduced level of oxidative markers (methionine sulfoxide and malondialdehyde), allowing improvements in the energy use in photosystem II under saline conditions in the salt-sensitive ecotype. In view of these results, we conclude that seed HP prompts a "metabolic imprint" related to ROS scavenger at the thylakoid level, improving further the physiological performance of the most sensitive ecotype.
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Quinoa is a strategic crop due to its high N content and its adaptability to adverse conditions, where most of the soils are deficient of nitrogen (N). The central question in this review was the following: How can quinoa yield low levels of nitrogen in the soils of Altiplano? This question was unraveled based on different factors: (1) fertilization effect on productivity, (2) fertilization limits, (3) uptake and assimilation of nitrogen parameters, (4) monoculture practice effect, and (5) possible sources and strategies. One hundred eleven articles of different scientific platforms were revised and data were collected. Information from articles was used to calculate the partial factor productivity for nitrogen (PFPN), the apparent use efficiency of N (APUEN), available nitrogen (AN), and nitrogen content harvested in grains (HarvN). Quinoa responds positively to fertilization, but differences in yield were found among irrigated and rainfed conditions. Quinoa can produce 1850 kg grains ha-1 with 50 kg N ha-1 under irrigated conditions, and 670 kg grains ha-1 with 15 kg N ha-1 in rainfed conditions. Quinoa increases seed yield and HarvN increases N fertilization, but decreases nitrogen efficiency. In Altiplano, without nitrogen fertilizer, the quinoa yield relies on between 500 and 1000 kg ha-1, which shows that in the soil, there are other nitrogen sources.
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The neurulation process is regulated by a large amount of genetic and environmental factors that determine the establishment, folding, and fusion of the neural plate to form the neural tube, which develops into the main structure of the central nervous system. A recently described factor involved in this process is glutamate. Through NMDA ionotropic receptor, glutamate modifies intracellular Ca2+ dynamics allowing the oriented cell migration and proliferation, essentials processes in neurulation. Glutamate synthesis depends on the mitochondrial enzyme known as glutaminase 1 (GLS1) that is widely expressed in brain and kidney. The participation of GLS 1 in prenatal neurogenic processes and in the adult brain has been experimentally established, however, its participation in early stages of embryonic development has not been described. The present investigation describes for the first time the presence and functionality of GLS1 in Xenopus laevis embryos during neurulation. Although protein expression levels remains constant, the catalytic activity of GLS1 increases significantly (~66%) between early (stage 12) and middle to late (stages 14-19) neurulation process. Additionally, the use of 6-diazo-5-oxo-L-norleucine (L-DON, competitive inhibitor of glutamine-depend enzymes), reduced significantly the GLS1 specific activity during neurulation (~36%) and induce the occurrence of neural tube defects involving its possible participation in the neural tube closure in Xenopus laevis embryos.
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Hymenoglossum cruentum (Hymenophyllaceae) is a poikilohydric, homoiochlorophyllous desiccation-tolerant (DT) epiphyte fern. It can undergo fast and frequent dehydration-rehydration cycles. This fern is highly abundant at high-humidity/low-light microenvironments within the canopy, although rapid changes in humidity and light intensity are frequent. The objective of this research is to identify genes associated to desiccation-rehydration cycle in the transcriptome of H. cruentum to better understand the genetic dynamics behind its desiccation tolerance mechanism. H. cruentum plants were subjected to a 7 days long desiccation-rehydration process and then used to identify key expressed genes associated to its capacity to dehydrate and rehydrate. The relative water content (RWC) and maximum quantum efficiency (F v/F m) of H. cruentum fronds decayed to 6% and 0.04, respectively, at the end of the desiccation stage. After re-watering, the fern showed a rapid recovery of RWC and F v/F m (ca. 73% and 0.8, respectively). Based on clustering and network analysis, our results reveal key genes, such as UBA/TS-N, DYNLL, and LHC, orchestrating intracellular motility and photosynthetic metabolism; strong balance between avoiding cell death and defense (CAT3, AP2/ERF) when dehydrated, and detoxifying pathways and stabilization of photosystems (GST, CAB2, and ELIP9) during rehydration. Here we provide novel insights into the genetic dynamics behind the desiccation tolerance mechanism of H. cruentum.
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Quinoa has been highlighted as a promising crop to sustain food security. The selection of physiological traits that allow identification genotypes with high Nitrogen use efficiency (NUE) is a key factor to increase Quinoa cultivation. In order to unveil the underpinning mechanisms for N-stress tolerance in Quinoa, three genotypes with similar phenology, but different NUE were developed under high (HN) or low (LN) nitrogen conditions. N metabolism processes and photosynthetic performance were studied after anthesis and in correlation with productivity to identify principal traits related to NUE. We found that protein content, net photosynthesis and leaf dry-mass were determinant attributes for yield at both HN and LN conditions. Contrastingly, the enhancement of N related metabolites ([Formula: see text], proline, betacyanins) and processes related with re-assimilation of [Formula: see text], including an increment of glutamine synthetase activity and up-regulation of CqAMT1,1 transporter expression in leaves, were negatively correlated with grain yield at both N conditions. Biochemical aspects of photosynthesis and root biomass were traits exclusively associated with grain yield at LN. The impact of N supply on seed quality is discussed. These results provide new insights towards the understanding the N metabolism of Quinoa.
Asunto(s)
Chenopodium quinoa/genética , Chenopodium quinoa/metabolismo , Genotipo , Nitrógeno/metabolismo , Biomasa , Clorofila/metabolismo , Fenotipo , Fotosíntesis , Carácter Cuantitativo Heredable , SemillasRESUMEN
Drought during senescence has become more common in Mediterranean climates in recent years. Chenopodium quinoa Willd has been identified as tolerant to poor soil conditions and drought. Previous observations have found that sufficient nitrogen (N) supply mitigates yield losses under terminal drought conditions. However, there is no understanding of the mechanisms behind this effect. We hypothesized that N up-regulates both photosynthetic and photoprotective elements during drought-induced senescence, alleviating the negative impact of drought on yield. The role of N supply and terminal drought on photoprotection was tested using three Chilean quinoa genotypes from different climatic zones: Faro, UdeC9, and BO78. Plants were grown under high nitrogen (HN) or low nitrogen (LN) conditions and subjected to terminal drought at the onset of senescence. Photosynthetic and photochemical and non-photochemical processes were evaluated at both the onset of drought and after 15 days of drought conditions. N supplementation modified most of the physiological parameters related to photochemical dissipation of energy, photosynthesis, and yield in quinoa. In contrast, water restriction did not affect photosynthesis in quinoa, and its effect on yield was dependent on the genotype. A significant interaction N × G was observed in photosynthesis, relative water content, protein content, Fv/Fm, and chlorophylls. In general, Faro was able to maintain higher levels of these attributes under LN conditions than UdeC9 and BO78. In addition, the interacting effects of N × W regulated the level of most pigments in quinoa as well as the photoprotective induction of non-photochemical quenching (NPQ) during senescence. During terminal drought at LN conditions, Faro presented a larger NPQ induction under drought conditions than UdeC9 and BO78, which was supported by a larger zeaxanthin content and de-epoxidation state of the xanthophyll pool. Interestingly, BO78 did not induce NPQ in response to drought-induced senescence but instead enhanced the content of betacyanins. This response needs to be researched in future works. Finally, we observed that LN supply reduced the correlationship between the de-epoxidation state of the xanthophyll cycle and NPQ. This could be an indication that N supply not only compromised the capacity for photosynthetic performance in quinoa plants, but also affected the plasticity of thermal dissipation, restricting further changes during drought-induced senescence.
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Quinoa cultivation has been expanded around the world in the last decade and is considered an exceptional crop with the potential of contributing to food security worldwide. The exceptional nutritional value of quinoa seeds relies on their high protein content, their amino acid profile that includes a good balance of essential amino acids, the mineral composition and the presence of antioxidants and other important nutrients such as fiber or vitamins. Although several studies have pointed to the influence of different environmental stresses in certain nutritional components little attention has been paid to the effect of the agroecological context on the nutritional properties of the seeds what may strongly impact on the consumer food's quality. Thus, aiming to evaluate the effect of the agroecological conditions on the nutritional profile of quinoa seeds we analyzed three quinoa cultivars (Salcedo-INIA, Titicaca and Regalona) at different locations (Spain, Peru and Chile). The results revealed that several nutritional parameters such as the amino acid profile, the protein content, the mineral composition and the phytate amount in the seeds depend on the location and cultivar while other parameters such as saponin or fiber were more stable across locations. Our results support the notion that nutritional characteristics of seeds may be determined by seed's origin and further analysis are needed to define the exact mechanisms that control the changes in the seeds nutritional properties.