Quinoa (Chenopodium quinoa Willd.) is a nutrient-rich grain native to South America and eaten worldwide as a healthy food, sometimes even referred to as a ”superfood”. Like quinoa grains, quinoa greens (green leaves, sprouts, and microgreens) are also rich in nutrients and have health promoting properties such as being antimicrobial, anticancer, antidiabetic, antioxidant, antiobesity, and cardio-beneficial. Quinoa greens are gluten-free and provide an excellent source of protein, amino acids, essential minerals, and omega-3 fatty acids. Quinoa greens represent a promising value-added vegetable that could resolve malnutrition problems and contribute to food and nutritional security. The greens can be grown year-round (in the field, high tunnel, and greenhouse) and have short growth durations. In addition, quinoa is salt-, drought-, and cold-tolerant and requires little fertilizer and water to grow. Nevertheless, consumption of quinoa greens as leafy vegetables is uncommon. To date, only a few researchers have investigated the nutritional properties, phytochemical composition, and human health benefits of quinoa greens. We undertook a comprehensive review of the literature on quinoa greens to explore their nutritional and functional significance to human health and to bring awareness to their use in human diets.

Quinoa (Chenopodium quinoa Willd.) is an herbaceous annual crop of the amaranth family (Amaranthaceae). It is increasingly cultivated for its nutritious grains, which are rich in protein and essential amino acids, lipids, and minerals. Quinoa exhibits a high tolerance towards various abiotic stresses including drought and salinity, which supports its agricultural cultivation under climate change conditions. The use of quinoa grains is compromised by anti-nutritional saponins, a terpenoid class of secondary metabolites deposited in the seed coat; their removal before consumption requires extensive washing, an economically and environmentally unfavorable process; or their accumulation can be reduced through breeding. In this study, we analyzed the seed metabolomes, including amino acids, fatty acids, and saponins, from 471 quinoa cultivars, including two related species, by liquid chromatography - mass spectrometry. Additionally, we determined a large number of agronomic traits including biomass, flowering time, and seed yield. The results revealed considerable diversity between genotypes and provide a knowledge base for future breeding or genome editing of quinoa.© 2022. The Author(s).

Chenopodium quinoa, a halophytic crop belonging to the Amaranthaceae, has remarkable resistance to harsh growth conditions and produces seed with excellent nutritional value. This makes it a suitable crop for marginal soils. However, to date most of the commercial cultivars are susceptible to preharvest sprouting (PHS). Meanwhile, understanding of the PHS regulatory mechanisms is still limited. Abscisic acid (ABA) has been demonstrated to be tightly associated with seed dormancy and germination regulation in many crops. Whether ABA metabolism pathway could be manipulated to prevent PHS in quinoa is worth investigating. In the present study, we tested the inhibitory effects of exogenous ABA on quinoa seed germination. By RNA-seq analysis we investigated the global gene expression changes during seed germination, and obtained 1066 ABA-repressed and 392 ABA-induced genes. Cis-elements enrichment analysis indicated that the promoters of these genes were highly enriched in motifs "AAAAAAAA" and "ACGTGKC (K = G/T)", the specific binding motifs of ABI3/VP1 and ABI5. Transcription factor annotation showed that 13 genes in bHLH, MADS-box, G2-like and NF-YB, and five genes in B3, bZIP, GATA and LBD families were specifically ABA-repressed and -induced, respectively. Furthermore, expression levels of 53 key homologs involved in seed dormancy and germination regulation were markedly changed. Hence, we speculated that the 18 transcription factors and the homologs were potential candidates involved in ABA-mediated seed dormancy and germination regulation, which could be manipulated for molecular breeding of quinoa elites with PHS tolerance in future.Copyright © 2020 Elsevier Masson SAS. All rights reserved.

水稻穗发芽是指水稻在收获前如遇连续阴雨天气或高温潮湿环境条件，往往诱导籽粒在穗上发芽的现象。穗发芽导致水稻种子活力和品质下降，给水稻生产带来巨大损失。对于水稻抗穗发芽育种，除了扩大对穗发芽突变体的筛选外，挖掘和克隆一些控制穗发芽的新基因并解析其穗发芽调控机制，是水稻抗穗发芽育种的重要工作。穗发芽过程中，水稻的淀粉酶活性增强、可溶性糖含量升高，且水稻籽粒中植物激素 ABA 和 GA 的含量及二者的平衡是决定穗发芽的关键。OsVP1等一些关键基因通过ABA信号通路控制水稻种子休眠，GA则可通过激活GA相关转录因子等调控种子萌发。本文从水稻穗发芽的内部生理因素及环境条件、水稻穗发芽的遗传机制、水稻穗发芽的分子机制和水稻穗发芽的性状改良这四方面进行综述，以期阐述穗发芽的整体调控机制，为水稻抗穗发芽品种的选育提供理论参考。

Quinoa is a high-value, nutritious crop that performs well in variable environments, marginal soils, and in diverse crop rotations. Quinoa’s many attributes make it an ideal crop for supporting human health in global communities and economies. To date, quinoa research has largely focused on traits in adult plants important for enhancing plant phenotypic plasticity, abiotic stress, disease resistance, and yield. Fewer studies have evaluated quinoa seed dormancy and suggest that most modern quinoa varieties have weak or no seed dormancy, and a narrow window of seed viability post-harvest. In other crops, diminished seed dormancy is a major risk factor for preharvest sprouting (PHS; germination on the panicle due to rain prior to harvest) and may also pose a similar risk for quinoa.

针对西北地区藜麦栽培土壤限制问题,研究盐碱胁迫对藜麦种子萌发及抗性相关酶特性的影响,探讨藜麦对盐碱土壤的适应机制,为藜麦在盐碱地的栽培实践提供理论依据。将中性盐(NaCl、Na₂SO₄)和碱性盐(NaHCO₃、NaCO₃)按不同比例混合模拟出20种混合盐碱条件对藜麦种子进行胁迫,分析盐碱胁迫下藜麦种子的发芽率、发芽势、发芽指数和抗氧化酶活性及同工酶表达。结果表明:5种盐碱胁迫均引起藜麦种子发芽率,发芽势,发芽指数降低,且随着盐浓度的增加,萌发受到显著抑制(P<0.05),对藜麦种子萌发影响较大。A,B处理组各浓度盐碱胁迫下超氧化物歧化酶(superoxide dismutase, SOD)和谷光甘肽还原酶(glutathione reductase, GR)活性均高于CK;C,D,E处理组SOD活性均低于CK,但GR活性在50 mmol·L^-1盐碱浓度时高于CK。过氧化物酶(peroxidase, POD)活性在盐碱浓度为50 mmol·L^-1时最高,随盐浓度增加POD活性比CK降低4倍,过氧化氢酶(catalase, CAT)活性随盐碱浓度的增加而下降。SDS-PAGE分析发现,盐碱胁迫可诱导SOD,POD,CAT,GR同工酶出现新的条带,但随着盐碱浓度的增加,酶带的表达量降低。上述结果说明,盐碱胁迫抑制藜麦种子萌发,其中Na₂SO₄和NaHCO₃对藜麦种子萌发抑制效应明显。本试验条件下,藜麦的耐盐碱阈值为50 mmol·L^-1,其中盐碱浓度是影响藜麦种子萌发的主要因素,盐组分值对藜麦种子萌发影响次之。

The economic and health significance of quinoa is steadily growing on a global scale. Nevertheless, the primary obstacle to achieving high yields in quinoa cultivation is pre-harvest sprouting (PHS), which is intricately linked to seed dormancy. However, there exists a dearth of research concerning the regulatory mechanisms governing PHS. The regulation of seed germination by various plant hormones has been extensively studied. Consequently, understanding the mechanisms underlying the role of endogenous hormones in the germination process of quinoa seeds and developing strategies to mitigate PHS in quinoa cultivation are of significant research importance. This study employed the HPLC-ESI-MS/MS internal standard and ELISA method to quantify 8 endogenous hormones. The investigation of gene expression changes before and after germination was conducted using RNA-seq analysis, leading to the discovery of 280 differentially expressed genes associated with the regulatory pathway of endogenous hormones. Additionally, a correlation analysis of 99 genes with significant differences identified 14 potential genes that may act as crucial “transportation hubs” in hormonal interactions. Through the performance of an analysis on the modifications in hormone composition and the expression of associated regulatory genes, we posit a prediction that implies the presence of a negative feedback regulatory mechanism of endogenous hormones during the germination of quinoa seeds. This mechanism is potentially influenced by the unique structure of quinoa seeds. To shed light on the involvement of endogenous hormones in the process of quinoa seed germination, we have established a regulatory network. This study aims to offer innovative perspectives on the breeding of quinoa varieties that exhibit resistance to PHS, as well as strategies for preventing PHS.

Endogenous abscisic acid (ABA) levels are regulated by both biosynthesis and catabolism of the hormone. ABA 8′-hydroxylase is considered to be the key catabolic enzyme in many physiological processes. We have previously identified that four members of the Arabidopsis (Arabidopsis thaliana) CYP707A gene family (CYP707A1 to CYP707A4) encode ABA 8′-hydroxylases, and that the cyp707a2 mutants showed an increase in ABA levels in dry and imbibed seeds. In this study, we showed that the cyp707a1 mutant accumulated ABA to higher levels in dry seeds than the cyp707a2 mutant. Expression analysis showed that the CYP707A1 was expressed predominantly during mid-maturation and was down-regulated during late-maturation. Concomitantly, the CYP707A2 transcript levels increased from late-maturation to mature dry seed. Phenotypic analysis of single and double cyp707a mutants indicates that the CYP707A1 is important for reducing ABA levels during mid-maturation. On the other hand, CYP707A2 is responsible for the regulation of ABA levels from late-maturation to germination. Moreover, CYP707A1 and CYP707A3 were also shown to be involved in postgermination growth. Spatial expression analysis suggests that CYP707A1 was expressed predominantly in embryo during mid-maturation, whereas CYP707A2 expression was detected in both embryo and endosperm from late-maturation to germination. Our results demonstrate that each CYP707A gene plays a distinct role during seed development and postgermination growth.

Silencing of CqGID1s from quinoa resulted in severely dwarfed plants, whereas CqGID1 overexpression significantly increased GA sensitivity and plant height in Arabidopsis thaliana. Gibberellin (GA) is an important phytohormone that regulates seed germination and growth, and the GIBBERELLIN-INSENSITIVE DWARF1 (GIDI) is a key mediator of GA. In this study, we identified three quinoa GID1 genes: the expression level of CqGID1c was low during the germination of quinoa seeds, whereas those of CqGID1b1 and CqGID1b2 were high, suggesting that CqGID1b1 and CqGID1b2 may play important roles in the germination of quinoa seeds. The silencing of CqGID1s in quinoa resulted in severe plant dwarfism, whereas CqGID1-overexpressing Arabidopsis had significantly increased plant heights. Overexpression of CqGID1s increased the sensitivity of plants to GA. CqGID1s-overexpressed Arabidopsis showed a significant increase in root length, hypocotyl length, seed germination rate, internode number, and flowering time. Both overexpression and silencing of CqGID1s caused changes in the endogenous hormone contents and the expression of genes related to GA biosynthesis and degradation, suggesting that GA-mediated plant growth and development is influenced by its signaling, biosynthesis, and degradation genes. Overall, our study identified and investigated quinoa CqGID1s, established a foundation for understanding the role of GID1 in plant growth and development, and provided a theoretical basis for elucidating the mechanism by which the GA signaling pathway regulates seed germination and plant height in quinoa.© 2025. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.

Seed vigor is an important trait closely related to improved seed quality and long-term germplasm conservation, and it gradually decreases during storage, which has become a major concern for agriculture. However, the underlying regulatory mechanisms of seed vigor loss in terms of genes remain largely unknown in quinoa. Here, two cultivars of quinoa seeds with different storage performance, Longli No.4 (L4) and Longli No.1 (L1), were subjected to transcriptome sequencing to decipher the pathways and genes possibly related to vigor loss under artificial aging. Multispectral imaging features and germination phenotypes showed significantly less seed vigor loss in L1 than in L4, indicating L1 seeds having stronger aging resistance and storability. Totally, 272 and 75 differentially expressed genes (DEGs) were, respectively, identified in L4 and L1 during aging. Transcriptomic analysis further revealed the differences in metabolic pathways, especially, flavonoid biosynthesis, TCA cycle, and terpenoid backbone biosynthesis were significantly enriched in L4 seeds, while carbon metabolism in L1 seeds, which involved key genes such as CHS, CHI, AACT, ENO1, IDH, NADP-ME, and HAO2L. It indicated that the adverse effects on flavonoids and terpenoids induced by aging might be the significant reasons for more vigor loss in storage sensitive seeds, whereas storage tolerant seeds had a stronger ability to maintain carbon metabolism and energy supply. These findings elucidated the underlying molecular mechanism of seed vigor loss in quinoa, which also provided novel insights into improving seed vigor through modern molecular breeding strategies.

Starch and sucrose metabolism plays a crucial role in the formation and development of bulbs in bulbous plants. However, these mechanisms remain unclear and unexplored in the corms of Freesia hybrida. Herein, we investigated the dynamics of the major form of carbohydrates and related enzyme activities and profiled the transcriptome of freesia corms at four developmental stages with the aim to reveal the relation between the expression of genes involved in the metabolism of carbohydrates and the accumulation of carbohydrates in corm developmental stages for further exploring the mechanism on the starch and sucrose metabolism regulating the formation and development of corms in F. hybrida. The content of starch, sucrose and soluble sugars followed an overall upward trend across the corm developmental stages. Activities of the adenosine diphosphoglucose pyrophosphorylase, starch branching enzyme and β-amylase generally followed the pattern of the starch and sucrose levels. Activities of sucrose phosphate synthase increased from corm formation till the initial swelling stage and subsequently reached a plateau. Activities of invertase and sucrose synthase peaked at the later rapid swelling stage. These suggested that the starch and sucrose dynamics paralleled corm swelling under the action of metabolic enzymes. A total of 100,999 unigenes were assembled in the transcriptomic analysis, and 44,405 unigenes of them were annotated. Analysis based on Clusters of Orthologous Groups suggested that carbohydrate transport and metabolism (9.34% of the sequences) was prominent across the corm developmental process. In total 3,427 differentially expressed genes (DEGs) were identified and the enrichment analysis detected starch and sucrose metabolism as a critical pathway in corm development, especially at the rapid swelling stage. Further, DEGs encoding key carbohydrate-metabolizing enzymes were identified and correlated to enzyme activities and carbohydrate accumulation. The results construct a valuable resource pool for further molecular-level studies, which are helpful for metabolic regulation of carbohydrates and improvement in F. hybrida.

Two possible sources of resistance to pre-harvest sprouting were evaluated in quinoa. They showed dormancy at harvest and significant variations in dormancy level in response to environmental conditions experienced during seed development. The aims of this work were to evaluate the importance of seed coats in the regulation of dormancy in this species, to investigate possible mechanisms of action and to assess association of seed coat properties with changes in dormancy level caused by the environment. Accessions Chadmo and 2-Want were grown under field conditions on different sowing dates during 2 years. Seed coats were manipulated and seed germination was evaluated at different temperatures. Seed coat perforation before incubation led to faster dormancy loss in both accessions. This effect decreased with delayed sowing date, and seeds expressed a level of dormancy not imposed by coats. This suggests the presence of embryo dormancy in the genus Chenopodium. Seeds of the accession 2-Want had a significantly thinner seed coat at later sowing dates, associated with a decreasing coat-imposed dormancy, but this pattern was not detected in Chadmo. The seed coat acts as a barrier to the release of endogenous abscisic acid (ABA) in quinoa, suggested by the increase in germination and a higher amount of ABA leached from perforated seeds. ABA is able to leach from seeds with an intact seed coat, suggesting that differences in seed coat thickness may allow the leakage of different amounts of ABA. This mechanism may contribute to the observed differences in dormancy level, either between sowing dates or between accessions.

The demand for plant-based proteins has increased remarkably over the last decade. Expanding the availability and variety of plant-based protein options has shown positive potential. This study aimed to investigate the qualitative and quantitative changes in amino acids of yellow and red quinoa seeds (YQ and RQ) during a 9-day germination period. The results showed that the germination process led to an increase in the total amino acids by 7.43% and 14.36% in the YQ and RQ, respectively. Both varieties exhibited significant (p < 0.05) increases in non-essential and essential amino acids, including lysine, phenylalanine, threonine, and tyrosine. The content of non-essential amino acids nearly reached the standard values found in chicken eggs. These results were likely attributed to the impact of the germination process in increasing enzymes activity and decreasing anti-nutrient content (e.g., saponins). A linear relationship between increased seeds’ hydration and decreased saponins content was observed, indicating the effect of water absorption in changing the chemical composition of the plant. Both sprouts showed positive germination progression; however, the sprouted RQ showed a higher germination rate than the YQ (57.67% vs. 43.33%, respectively). Overall, this study demonstrates that germination is a promising technique for enhancing the nutritional value of quinoa seeds, delivering sprouted quinoa seeds as a highly recommended source of high-protein grains with notable functional properties.

Bud dormancy is an evolved trait that confers adaptation to harsh environments, and affects flower differentiation, crop yield and vegetative growth in perennials. ABA is a stress hormone and a major regulator of dormancy. Although the physiology of bud dormancy is complex, several advancements have been achieved in this field recently by using genetics, omics and bioinformatics methods. Here, we review the current knowledge on the role of ABA and environmental signals, as well as the interplay of other hormones and sucrose, in the regulation of this process. We also discuss emerging potential mechanisms in this physiological process, including epigenetic regulation.