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Metabolic Profiling Reveals Genotype-Associated Alterations in Carotenoid Content during Banana Postharvest Ripening
Abstract
Banana fruits have attracted considerable attention for health-promoting effects attributed to ubiquitous functional metabolites. However, genotype-dependent accumulation patterns of carotenoids in banana remain largely unclear. Here, we performed a systematic metabolomic investigation of 18 banana cultivars of the AAA, AAB, or ABB genome groups. Our results indicate that the levels of soluble sugars increase during postharvest ripening regardless of genotype, whereas amino acids (AAs) and tricarboxylic acid (TCA) cycle-derived organic acids display genotype-dependent patterns. The levels of AAs derived from the glycolytic pathway increased, whereas those derived from the TCA cycle significantly decreased during ripening. The carotenoid composition in banana pulp was genotype-specific, and the contents of α-carotene were the highest in AAA-genome bananas. Moreover, high α-carotene and β-carotene contents in banana were correlated with elevated levels of TCA cycle-derived AAs and decreased levels of glycolysis-derived AAs. Taken together, these findings provide a comprehensive understanding of genotype-associated carotenoid accumulation, thereby facilitating the breeding of future high carotenoid banana cultivars.
... Carotenoids accumulate in many flowers, fruits, vegetables, and grains, and their quantity and composition exhibit significant species-and variety-specific variations [15,20,[27][28][29][30][31][32]. Several studies have determined the carotenoid concentration in potato tubers, consisting of mainly violaxanthin, lutein, zeaxanthin, and antheraxanthin. ...
Individual carotenoids provide significant health benefits to humans, and potatoes are recognized as some of the most stable crops, distinguished by their substantial carotenoid content. To elucidate the accumulation patterns of individual carotenoids in potatoes, we quantified the carotenoid content in yellow- and white-fleshed genotypes across five developmental stages using LC-MS/MS. A total of 22 carotenoids were identified in yellow potatoes, whereas 18 were detected in white potatoes. The yellow-fleshed genotype was characterized by high levels of individual carotenoids and xanthophyll esters, with violaxanthin as the dominant component. The white-fleshed genotype exhibited low concentrations of individual carotenoids and xanthophyll esters, with lutein as the predominant compound. Notably, lutein, violaxanthin, zeaxanthin, antheraxanthin, neoxanthin, violaxanthin myristate, and lutein myristate were consistently detected during the developmental period in both genotypes. Violaxanthin myristate was identified as the dominant xanthophyll ester in both genotypes, showing an increasing trend throughout the tuber development stages. In contrast, xanthophyll esters maintained stable low levels in white-fleshed potatoes while exhibiting increasing types and contents in yellow-fleshed potatoes as the tubers continued to grow. Additionally, violaxanthin exhibited a significant correlation with b*, H°, and C*, suggesting that it may play an important role in forming yellow flesh.
... Although mammalian systems have adopted these tools in advance of plant research, key pioneering events in plant MSI research have greatly contributed to shaping this rapidly evolving discipline. Recently, the use of MSI techniques has shown immense promise for mapping pesticides and environmental pollutants within plant tissues Selby et al., 2024;Wu et al., 2020), as well as secondary metabolites involved in biosynthetic pathways of medicinal plants Li, Chen, et al., 2023;Yamamoto et al., 2019), key targets in breeding (Dong et al., 2020;Li et al., 2022;Lu et al., 2018;O'Neill & Lee, 2020;Sheng et al., 2023), plant growth and reproduction Feenstra et al., 2017;Salimath et al., 2021;Shiono et al., 2017;Shiono & Taira, 2020;Yin et al., 2022), and plant defense and ecological response (Crecelius et al., 2017;de Moraes Pontes et al., 2020;Dreisbach et al., 2021;Liu et al., 2021;Parker et al., 2023;Righetti et al., 2021). Additionally, some plant-specific challenges, such as the immense complexity of secondary metabolism, present considerable limitations that have yet to be fully addressed for state-of-the-art MSI techniques. ...
Mass spectrometry imaging (MSI) has become increasingly popular in plant science due to its ability to characterize complex chemical, spatial, and temporal aspects of plant metabolism. Over the past decade, as the emerging and unique features of various MSI techniques have continued to support new discoveries in studies of plant metabolism closely associated with various aspects of plant function and physiology, spatial metabolomics based on MSI techniques has positioned it at the forefront of plant metabolic studies, providing the opportunity for far higher resolution than was previously available. Despite these efforts, profound challenges at the levels of spatial resolution, sensitivity, quantitative ability, chemical confidence, isomer discrimination, and spatial multi‐omics integration, undoubtedly remain. In this Perspective, we provide a contemporary overview of the emergent MSI techniques widely used in the plant sciences, with particular emphasis on recent advances in methodological breakthroughs. Having established the detailed context of MSI, we outline both the golden opportunities and key challenges currently facing plant metabolomics, presenting our vision as to how the enormous potential of MSI technologies will contribute to progress in plant science in the coming years.
... The WI value represents the whiteness of the sample, which increased with the ripening of the banana during the ripening stage, but the lowest whiteness was observed on the ninth day of development, which was also affected by brown spots. Thus, evidence of the ripening course is visible, in which the color of the banana peel changes from green to yellow due to chlorophyll degradation with marginal changes in carotenoids and phenols, while brown spots appear during ripening stage 9, followed by softening of the pulp, which begins spoilage, leading to serious economic losses [5,7,8,10,21,46,47]. ...
Banana is a typical tropical fruit that is widely cultivated in tropical and subtropical regions, and consumed daily because of its superior nutritional profile. This study aimed to investigate the changes in bananas’ physicochemical properties and nutritional composition from ripening stages 1 to 9. The analysis included color (peel and flesh), texture, total starch content, resistant starch content, free sugar content, minerals (calcium, magnesium, potassium, sodium, iron, phosphorus), and vitamin content. At the same time, the browning enzyme activities of polyphenol oxidase (PPO), peroxidase, oxalic acid, and tannin were also measured. In this study, the banana peel gradually changed from dark green to light green to yellow during the ripening process, eventually appearing with many black spots, while the pulp color changed from white to light yellow and decreased in brightness. The pulp became softer with a prolonged ripening period, which was inconsistent with the time point of peel color change (ripening period 4–5). However, total starch content decreased from 74% to 31%, resistant starch originally at 32% gradually decreased to 5%, and free sugar content gradually increased with increasing ripening period. Vitamin C content and PPO activity decreased significantly during the ripening period, while other vitamins and minerals did not change significantly. In contrast, oxalic acid, tannin content, and peroxidase activity were negatively correlated with the ripening period. Therefore, this study reveals the changes in the nutrient content of bananas at different ripening stages, which may provide helpful information for researchers and producers to identify suitable raw materials for producing stable and high-nutrition-value banana-related foods during the peak raw material abundance seasons while reducing business risk.
... In the major banana-producing countries, the per capita consumption of Plantains ranges from 40 kg/year in the Democratic Republic of the Congo to 153 kg/year in Gabon (Akyeampong and Escalant, 1998). Plantain fruits have orange flesh and contain high levels of carotenoids (Sheng et al., 2023). Most Plantain varieties are resistant to Fusarium wilt caused by Fusarium oxysporum f. sp. ...
Bananas (Musa spp.) are one of the world’s most important fruit crops and play a vital role in food security for many developing countries. Most banana cultivars are triploids derived from inter- and intraspecific hybridizations between the wild diploid ancestor species Musa acuminate (AA) and M. balbisiana (BB). We report two haplotype-resolved genome assemblies of the representative AAB-cultivated types, Plantain and Silk, and precisely characterize ancestral contributions by examining ancestry mosaics across the genome. Widespread asymmetric evolution is observed in their subgenomes, which can be linked to frequent homologous exchange events. We reveal the genetic makeup of triploid banana cultivars and verify that subgenome B is a rich source of disease resistance genes. Only 58.5% and 59.4% of Plantain and Silk genes, respectively, are present in all three haplotypes, with >50% of genes being differentially expressed alleles in different subgenomes. We observed that the number of upregulated genes in Plantain is significantly higher than that in Silk at one-week post-inoculation with Fusarium wilt tropical race 4 (Foc TR4), which confirms that Plantain can initiate defense responses faster than Silk. Additionally, we compared genomic and transcriptomic differences among the genes related to carotenoid synthesis and starch metabolism between Plantain and Silk. Our study provides resources for better understanding the genomic architecture of cultivated bananas and has important implications for Musa genetics and breeding.
Fruit quality traits are determined to a large extent by their metabolome. The metabolite content of climacteric fruit changes drastically during ripening and postharvest storage and has been investigated extensively. However, the spatial distribution of metabolites and how it changes in time has received much less attention as fruit are usually considered as homogenous plant organs. Yet, spatio-temporal changes of starch, which is hydrolysed during ripening, has been used for ages as a ripening index. As vascular transport of water, and, hence, convective transport of metabolites slows down in mature fruit and even stalls after detachment, spatio-temporal changes in their concentration are likely affected by diffusive transport of gaseous molecules that act as substrate (O2), inhibitor (CO2) or regulator (ethylene, NO) of the metabolic pathways that are active during climacteric ripening. In this review we discuss such spatio-temporal changes of the metabolome and how they are affected by transport of metabolic gases and gaseous hormones. As there are currently no measurement techniques available to measure the metabolite distribution repeatedly by nondestructive means, we introduce reaction-diffusion models as an in silico tool to compute it. We show how the different components of such a model can be integrated and used to better understand the role of spatio-temporal changes of the metabolome in ripening and postharvest storage of climacteric fruit that is detached from the plant and discuss future research needs.
Banana is a good source of carotenoids, which are bioactive metabolites with health beneficial properties for human. However, the molecular mechanism of carotenoid accumulation in banana fruit is largely unclear. In this study, we found that high temperature elevated carotenoid production in banana pulp, which is presumably due to upregulation of a subset of carotenogenic genes as well as a carotenoid biosynthesis regulator MaSPL16. Moreover, an ethylene signaling component MaEIL9 was identified, whose transcript and protein contents were also induced by high temperature. In addition, MaEIL9 positively regulates transcription of MaDXR1, MaPDS1, MaZDS1 and MaSPL16 through directly targeting their promoters. Overexpression of MaEIL9 in tomato fruit substantially increased the expression of carotenoid formation genes and elevated carotenoid content. Importantly, transiently silencing MaEIL9 in banana fruit weakened carotenoid production caused by high temperature. Taken together, these results indicate that high temperature induces carotenoid production in banana fruit, at least in part, through MaEIL9-mediated activation of MaDXR1, MaPDS1, MaZDS1 and MaSPL16 expression.
Carotenoids are isoprenoid metabolites synthesized de novo in all photosynthetic organisms. Carotenoids are essential for plants with diverse functions in photosynthesis, photoprotection, pigmentation, phytohormone synthesis, and signaling. They are also critically important for humans as precursors of vitamin A synthesis and as dietary antioxidants. The vital roles of carotenoids to plants and humans have prompted significant progress toward our understanding of carotenoid metabolism and regulation. New regulators and novel roles of carotenoid metabolites are continuously revealed. This review focuses on current status of carotenoid metabolism and highlights recent advances in comprehension of the intrinsic and multi-dimensional regulation of carotenoid accumulation. We also discuss the functional evolution of carotenoids, the agricultural and horticultural application, and some key areas for future research.
Background
Banana fruits are rich in various high-value metabolites and play a key role in the human diet. Of these components, carotenoids have attracted considerable attention due to their physiological role and human health care functions. However, the accumulation patterns of carotenoids and genome-wide analysis of gene expression during banana fruit development have not been comprehensively evaluated.
Results
In the present study, an integrative analysis of metabolites and transcriptome profiles in banana fruit with three different development stages was performed. A total of 11 carotenoid compounds were identified, and most of these compounds showed markedly higher abundances in mature green and/or mature fruit than in young fruit. Results were linked to the high expression of carotenoid synthesis and regulatory genes in the middle and late stages of fruit development. Co-expression network analysis revealed that 79 differentially expressed transcription factor genes may be responsible for the regulation of LCYB (lycopene β-cyclase), a key enzyme catalyzing the biosynthesis of α- and β-carotene.
Conclusions
Collectively, the study provided new insights into the understanding of dynamic changes in carotenoid content and gene expression level during banana fruit development.
We obtained a new hybrid soybean (Hybrid) by hybridizing β-carotene-enhanced soybean (BCE; Glycine max L.) containing the phytoene synthase-2A-carotene desaturase gene and wild-type soybean (Wild; Glycine soja). To investigate metabolic changes between variants, we performed metabolic profiling of leaves (three growth stages) and seeds. Multivariate analyses revealed significant metabolic differences between genotypes in seeds and leaves, with seeds showing accumulation of phytosterols, tocopherols, and carotenoids (BCE only), indicating co-induction of the methylerythritol 4-phosphate and mevalonic acid pathways. Additionally, Hybrid produced intermediate levels of carotenoids and high levels of amino acids. Principal component analysis revealed metabolic discrimination between growth stages of soybean leaves and identified differences in leaf groups according to different genotypes at 8, 12, and 16 weeks, with Wild showing higher levels of environmental stress-related compounds relative to BCE and Hybrid leaves. The metabolic profiling approach could be a useful tool to identify metabolic links in various soybean cultivars.
Food safety issues caused by pesticide residue have exerted far-reaching impacts on human daily life, yet the available detection methods normally focus on surface residue rather than pesticide penetration to the internal area of foods. Herein, we demonstrated gold nanoparticle (AuNP)-immersed paper imprinting mass spectrometry imaging (MSI) for monitoring pesticide migration behaviors in various fruits and vegetables (i.e., apple, cucumber, pepper, plum, carrot, and strawberry). By manually stamping food tissues onto AuNP-immersed paper, this method affords the spatiotemporal visualization of insecticides and fungicides within fruits and vegetables, avoiding tedious and time-consuming sample preparation. Using the established MSI platform, we can track the migration of insecticides and fungicides into the inner region of foods. The results revealed that both the octanol-water partition coefficient of pesticides and water content of garden stuffs could influence the discrepancy in the migration speed of pesticides into food kernels. Taken together, this nanopaper imprinting MSI is poised to be a powerful tool because of its simplicity, rapidity, and easy operation, offering the potential to facilitate further applications in food analysis. Moreover, new perspectives are given to provide guidelines for the rational design of novel pesticide candidates, reducing the risk of food safety issues caused by pesticide residue.
Rice with panicle-blast resistance is needed for stable rice production. Although we have previously demonstrated that OsGF14b underlies a quantitative trait locus that positively regulates rice panicle blast resistance, the mechanism is still unknown. In this study, a multi-omics approach was used to investigate the possible downstream signaling pathway regulated by OsGF14b. OsGF14b both strongly activated the gibberellin biosynthetic pathway during pathogen infection and reprogrammed the lignin biosynthetic pathway. Reduced lignin accumulation was observed in glumes of OsGF14b-overexpressing plants in comparison with the wild type after pathogen inoculation. OsGF14b activated the auxin and jasmonic acid signaling pathways, but inactivated the salicylic acid signaling pathway. Auxin and jasmonic acid appeared to act independently on OsGF14b-mediated panicle blast resistance. The roles of gibberellin, lignin, and auxin were different from their roles in leaf blast, suggesting that different mechanisms underlie leaf and panicle blast resistance in rice. This study provides a comprehensive catalog of molecular changes that could be targets for future studies of rice panicle blast resistance.
Red peppers (Capsicum annuum) are rich in carotenoids and are widely grown and consumed all over the world. Today’s consumption patterns are characterized by periodical purchases of food and longer food storage periods, including raw fruits and vegetables, which could have a negative effect on healthy components. This study aims to investigate the individual carotenoid content in Lamuyo-variety red peppers in cool storage (7 °C) for three weeks. Carotenoid dcarotene (272), β-carotene (2167), β-cryptoxanthin (525), violaxanthin (770), capsanthin (9667), phytoene (348) and phytofluene (143). Carotenoid concentrations did not significantly vary after 21 days under household refrigeration conditions and thus the nutritional supply of provitamin A carotenoids and of carotenoids with eye health benefits such as lutein and zeaxanthin, as well as others with potential health benefits in humans such as capsanthin, violaxathin, phytoene and phytofluene.
Design and discovery of carrier-mediated modified pesticides are vital for reducing pesticide dosage and increasing utilization, yet it remains a great challenge due to limited insights into plant translocation mechanisms. Nanostructure/nanoparticle assisted laser desorption/ionization strategy has established itself as a preferential analytical tool for biological tissue analysis, whereas potential applications in plant sciences are hindered with regard to the inability to slice plant leaves and petals. Herein, we report gold nanoparticle (AuNP)-immersed paper imprinting mass spectrometry imaging (MSI) for the spatiotemporal visualization of pesticide translocation in plant leaves. This approach plays a dual role in preserving spatial information and improving ionization efficiency for pesticides regardless of imaging artifacts due to homogenous AuNP deposition. Using this MSI platform, we proposed the elaborate plant translocation mechanism of agrochemicals for the first time, which is currently poorly understood. The dynamic processes of carrier-mediated pesticides can be clearly visualized, including crossing of plasma membranes by transporters, translocation downward in stems through the phloem, diffusion to the xylem and, conversely, accumulation at margins of the treated leaves. Moreover, this AuNP-assisted paper imprinting method could be highly compatible with laser-based MSI instruments, expediting researches across a broad range of fields, especially in nanomaterial development and life sciences.
Banana (Musa acuminata, AAA group) is a representative climacteric fruit with essential nutrients and pleasant flavors. Control of its ripening determines both the fruit quality and the shelf life. NAC (NAM, ATAF, CUC2) proteins, as one of the largest superfamilies of transcription factors, play crucial roles in various functions, especially developmental processes. Thus, it is important to conduct a comprehensive identification and characterization of the NAC transcription factor family at the genomic level in M. acuminata. In this article, a total of 181 banana NAC genes were identified. Phylogenetic analysis indicated that NAC genes in M. acuminata, Arabidopsis, and rice were clustered into 18 groups (S1–S18), and MCScanX analysis disclosed that the evolution of MaNAC genes was promoted by segmental duplication events. Expression patterns of NAC genes during banana fruit ripening induced by ethylene were investigated using RNA-Seq data, and 10 MaNAC genes were identified as related to fruit ripening. A subcellular localization assay of selected MaNACs revealed that they were all localized to the nucleus. These results lay a good foundation for the investigation of NAC genes in banana toward the biological functions and evolution.
Ripening of the model fruit tomato (Solanum lycopersicum) is controlled by a transcription factor network including NAC (NAM, ATAF1/2, and CUC2) domain proteins such as No-ripening (NOR), SlNAC1, and SlNAC4, but very little is known about the NAC targets or how they regulate ripening. Here, we conducted a systematic search of fruit-expressed NAC genes and showed that silencing NOR-like1 (Solyc07g063420) using virus-induced gene silencing (VIGS) inhibited specific aspects of ripening. Ripening initiation was delayed by 14 days when NOR-like1 function was inactivated by CRISPR/Cas9 and fruits showed obviously reduced ethylene production, retarded softening and chlorophyll loss, and reduced lycopene accumulation. RNA-sequencing profiling and gene promoter analysis suggested that genes involved in ethylene biosynthesis (SlACS2, SlACS4), color formation (SlGgpps2, SlSGR1), and cell wall metabolism (SlPG2a, SlPL, SlCEL2, and SlEXP1) are direct targets of NOR-like1. Electrophoretic mobility shift assays (EMSA), chromatin immunoprecipitation-quantitative PCR (ChIP-qPCR), and dual-luciferase reporter assay (DLR) confirmed that NOR-like1 bound to the promoters of these genes both in vitro and in vivo, and activated their expression. Our findings demonstrate that NOR-like1 is a new positive regulator of tomato fruit ripening, with an important role in the transcriptional regulatory network.
Banana is an edible fruit and is herbaceous flowering plant belonging to the genus Musa and the family Musaceae. Banana is also eaten as cooked vegetable (and is then called plantains). All the edible banana fruits are seedless (parthenocarpic) and belong to two main species, Musa acuminata Colla and Musa balbisiana Colla. The hybrid from these two species Musa x paradisiaca L. is also available nowadays. Although banana is native to Indomalaya and Australia, Papua New Guinea was the first to domesticate this fruit. Banana has now spread to almost 135 countries around the world. As per 2016 data, nearly 28 per cent of the total world’s banana production comes from India and China. Cavendish group banana, being the main export item from the banana-exporting countries, usually refers to soft, sweet, and dessert banana in the Western countries, but the plantain bananas have firm, starchy fruit which is suitable for cooking as a vegetable. Banana is known to be rich not only in carbohydrates, dietary fibres, certain vitamins and minerals, but is also rich in many health-promoting bioactive phytochemicals. General composition including various bioactives and their health contributions has been reviewed in this paper.
Bananas (Musa spp.) are not only one of the most popular fruits but also important staple food in some tropic countries. Carotenoid in pulp of banana fruits is an important source of vitamin A. While little is reported on carotenoid in peel, which is traditionally recognized as byproduct of food industry and may be a potential source of antioxidant compounds. Here we determined and reported carotenoid content and composition in both peel and pulp of 36 different banana cultivars. A total of 3 and 8 carotenoid components were identified in pulp and peel, respectively. The results show that carotenoid ranged from 0.18 to 36.82 µg/g FW in pulp and 0.9 to 16.2 µg/g FW in peel. Vitamin A values as retinol activity equivalents (RAE) in the pulp ranged from 0.003 to 1.85 µg/g. This report represents the first comprehensive assessment of carotenoid profiling in both fruit peel and pulp of 36 white to orange types of bananas and demonstrates banana peel product is also a valuable source with great nutritional properties.
Carotenoids are a diverse group of pigments widely distributed in nature. The vivid yellow, orange, and red colors of many horticultural crops are attributed to the overaccumulation of carotenoids, which contribute to a critical agronomic trait for flowers and an important quality trait for fruits and vegetables. Not only do carotenoids give horticultural crops their visual appeal, they also enhance nutritional value and health benefits for humans. As a result, carotenoid research in horticultural crops has grown exponentially over the last decade. These investigations have advanced our fundamental understanding of carotenoid metabolism and regulation in plants. In this review, we provide an overview of carotenoid biosynthesis, degradation, and accumulation in horticultural crops and highlight recent achievements in our understanding of carotenoid metabolic regulation in vegetables, fruits, and flowers.
Carotenoids are critical for photosynthetic function in chloroplasts, and are essential for the formation of the prolamellar body in the etioplasts of dark-grown (etiolated) seedlings. They are also precursors for plant hormones in both types of plastids. Lutein is one of the most abundant carotenoids found in both plastids. In this study we examine the regulation of lutein biosynthesis and investigate the effect of perturbing carotenoid biosynthesis on the formation of the lattice-like membranous structure of etioplasts, the prolamellar body (PLB). Analysis of mRNA abundance in wildtype and lutein-deficient mutants, lut2 and ccr2, in response to light transitions and herbicide treatments demonstrated that the mRNA abundance of the carotenoid isomerase (CRTISO) and epsilon-cyclase (εLCY) can be rate limiting steps in lutein biosynthesis. We show that accumulation of tetra-cis-lycopene and all-trans-lycopene correlates with the abundance of mRNA of several carotenoid biosynthetic genes. Herbicide treatments that inhibit carotenoid biosynthetic enzymes in wildtype and ccr2 etiolated seedlings were used to demonstrate that the loss of the PLB in ccr2 mutants is a result of perturbations in carotenoid accumulation, not indirect secondary effects, as PLB formation could be restored in ccr2 mutants treated with norflurazon.
Metabolism has an essential role in biological systems. Identification and quantitation of the compounds in the metabolome is defined as metabolic profiling, and it is applied to define metabolic changes related to genetic differences, environmental influences and disease or drug perturbations. Chromatography-mass spectrometry (MS) platforms are frequently used to provide the sensitive and reproducible detection of hundreds to thousands of metabolites in a single biofluid or tissue sample. Here we describe the experimental workflow for long-term and large-scale metabolomic studies involving thousands of human samples with data acquired for multiple analytical batches over many months and years. Protocols for serum- and plasma-based metabolic profiling applying gas chromatography-MS (GC-MS) and ultraperformance liquid chromatography-MS (UPLC-MS) are described. These include biofluid collection, sample preparation, data acquisition, data pre-processing and quality assurance. Methods for quality control-based robust LOESS signal correction to provide signal correction and integration of data from multiple analytical batches are also described.
Carotenoids are a diverse group of colourful pigments naturally found in plants, algae, fungi and bacteria. They play essential roles in development, photosynthesis, root-mycorrhizal interactions and the production of phytohormones, such as abscisic acid and strigolactone. Carotenoid biosynthesis is regulated throughout the life cycle of a plant with dynamic changes in composition matched to prevailing developmental requirements and in response to external environmental stimuli. There are key regulatory nodes in the pathway that control the flux of metabolites into the pathway and alter flux through the pathway. The molecular nature of the mechanisms regulating carotenoid biosynthesis, including evidence for metabolite feedback, transcription and epigenetic control as well as their accumulation, storage and degradation will be the focus of this review.
Carotenoids and tocopherols are the two most abundant groups of lipid-soluble antioxidants in chloroplasts. In addition to their many functional roles in photosynthetic organisms, these compounds are also essential components of animal diets, including humans. During the past decade, a near complete set of genes required for the synthesis of both classes of compounds in photosynthetic tissues has been identified, primarily as a result of molecular genetic and biochemical genomics-based approaches in the model organisms Arabidopsis thaliana and Synechocystis sp. PCC6803. Mutant analysis and transgenic studies in these and other systems have provided important insight into the regulation, activities, integration, and evolution of individual enzymes and are already providing a knowledge base for breeding and transgenic approaches to modify the types and levels of these important compounds in agricultural crops.
The accumulation of carotenoids in higher plants is regulated by the environment, tissue type and developmental stage. In Brassica napus leaves, beta-carotene and lutein were the main carotenoids present while petals primarily accumulated lutein and violaxanthin. Carotenoid accumulation in seeds was developmentally regulated with the highest levels detected at 35-40 days post anthesis. The carotenoid biosynthesis pathway branches after the formation of lycopene. One branch forms carotenoids with two beta rings such as beta-carotene, zeaxanthin and violaxanthin, while the other introduces both beta- and epsilon-rings in lycopene to form alpha-carotene and lutein. By reducing the expression of lycopene epsilon-cyclase (epsilon-CYC) using RNAi, we investigated altering carotenoid accumulation in seeds of B. napus. Transgenic seeds expressing this construct had increased levels of beta-carotene, zeaxanthin, violaxanthin and, unexpectedly, lutein. The higher total carotenoid content resulting from reduction of epsilon-CYC expression in seeds suggests that this gene is a rate-limiting step in the carotenoid biosynthesis pathway. epsilon-CYC activity and carotenoid production may also be related to fatty acid biosynthesis in seeds as transgenic seeds showed an overall decrease in total fatty acid content and minor changes in the proportions of various fatty acids.
In tomato (Solanum lycopersicum), phytoene synthase-1 (PSY-1) is the key biosynthetic enzyme responsible for the synthesis of fruit carotenoids. To further our understanding of carotenoid formation in tomato fruit, we characterized the effect of constitutive expression of an additional tomato Psy-1 gene product. A quantitative data set defining levels of carotenoid/isoprenoid gene expression, enzyme activities, and metabolites was generated from fruit that showed the greatest perturbation in carotenoid content. Transcriptional upregulation, resulting in increased enzyme activities and metabolites, occurred only in the case of Psy-1, Psy-2, and lycopene cyclase B. For reactions involving 1-deoxy-d-xylulose5-phosphate synthase, geranylgeranyl diphosphate synthase, phytoene desaturase, zeta-carotene desaturase, carotene isomerase, and lycopene beta-cyclase, there were no correlations between gene expression, enzyme activities, and metabolites. Perturbations in carotenoid composition were associated with changes in plastid type and with chromoplast-like structures arising prematurely during fruit development. The levels of >120 known metabolites were determined. Comparison with the wild type illustrated that key metabolites (sucrose, glucose/fructose, and Glu) and sectors of intermediary metabolism (e.g., tricarboxylic [corrected] acid cycle intermediates and fatty acids) in the Psy-1 transgenic mature green fruit resembled changes in metabolism associated with fruit ripening. General fruit developmental and ripening properties, such as ethylene production and fruit firmness, were unaffected. Therefore, it appears that the changes to pigmentation, plastid type, and metabolism associated with Psy-1 overexpression are not connected with the ripening process.
Black wolfberry is a commonly cultivated woody plant in China, and is rich in nutrients that are beneficial for human. To characterize the endogenous metabolite differences among black wolfberry fruits grown in different geographical regions, mass spectrometry-based metabolomic and lipidomic analyses were performed in black wolfberry grown in nine locations throughout five provinces in China, from which 204 primary and specialized metabolites, and 267 lipids were identified in their fruits. Three samples from Alxa Left Banner, Jinta, and Minqin showed dramatically altered metabolite profiles, displaying higher levels of phenolic acids, soluble sugars and flavonoids, but lower levels of tricarboxylic acid cycle intermediates and aromatic amino acids. Moreover, the lipid profile of the Alxa Left Banner sample was strikingly distinct from all other samples, with high levels of monogalactosyl diacylglycerol and sulfoquinovosyl diacylglycerol, which are positively correlated with their anti-inflammatory capacities. These findings thus prompt for further studies on black wolfberry fruit for their health benefits.
Ultra-high-performance liquid chromatography high-resolution mass spectrometry (UHPLC–HRMS) variants currently represent the best tools to tackle the challenges of complexity and lack of comprehensive coverage of the metabolome. UHPLC offers flexible and efficient separation coupled with high-sensitivity detection via HRMS, allowing for the detection and identification of a broad range of metabolites. Here we discuss current common strategies for UHPLC–HRMS-based metabolomics, with a focus on expanding metabolome coverage.
The present study was conducted to evaluate effect of ethephon and acetylene treatments on phenolics, flavonoids and antioxidant activity of banana flesh and their bioaccessibility. Total phenolics, flavonoids and antioxidant activity (DPPH, ABTS, FRAP) were measured at different phases of simulated gastrointestinal digestion of banana treated with 1000 ppm ethephon and 1000 ppm acetylene against natural ripening. The results revealed that inducing ripening lowers the content of phenolics, flavonoids and antioxidant activity considerably in the fresh fruit. Bioavailability of phenolics, flavonoids and FRAP activity were increased significantly (p < 0.05) after gastric digestion regardless of the treatment. The release of polyphenols and flavonoids during gastric digestion in treated banana was more significant than in naturally ripened banana. Recovery of polyphenols after dialysis was significantly high in naturally ripened banana. Dialyzable flavonoids, DPPH and ABTS activities of dialyzed fractions were not significantly affected by ethephon or acetylene treatments.
The variations of volatile phenolic metabolites in the artificially ripened banana fruits (Musa Sapientum) were studied. The fruits were artificially ripened using calcium carbide, ethephon (Ethrel) solution, and ethylene gas. The phenolics were extracted from the fruit pulp and analysed using gas chromatography-mass spectrometry. The obtained phenolic compounds were quantified with reference to the concentration of 1,3-dihydroxybenzene. The results showed significant upregulation of sinapic acid, butylated hydroxytoluene, α-hydroxycinnamic acid, isoferulic acid, and downregulation of 2,4,6-trimethoxytoluene and elemicin in the fruits ripened with calcium carbide. Ripening with ethylene gas showed upregulation of vanillic acid and the ripening by dipping the fruits in ethephon solution showed up-regulation of 2,4,6-trimethoxytoluene, p-coumaric acid and downregulation of isoferulic acid. The data was subjected to analysis of variance (ANOVA) and principal component analysis (PCA). The ANOVA analysis statistically certified the significance of variation among different ripening methods and the quantified phenolic metabolites. The results obtained from the PCA studies indicate that the artificially ripened banana fruits with ethylene and ethephon exhibited distinct patterns as compared to that of the naturally ripened ones.
Due to the ongoing prevalence of vitamin A deficiency (VAD) in developing countries there has been a large effort towards increasing the carotenoid content of staple foods via biofortification. Common strategies used for carotenoid biofortification include altering flux through the biosynthesis pathway to direct synthesis to a specific product, generally β-carotene, or via increasing the expression of genes early in the carotenoid biosynthesis pathway. Recently, carotenoid biofortification strategies are turning towards increasing the retention of carotenoids in plant tissues either via altering sequestration within the cell or via downregulating enzymes known to cause degradation of carotenoids. To date, little attention has focused on increasing the stability of carotenoids, which may be a promising method of increasing carotenoid content in staple foods.
Carotenoids are a kind of bioactive compounds that exhibit health-promoting properties for humans, but their regulation in bananas during fruit ripening remains largely unclear. Here we found that total carotenoid content continued to be elevated along the course of banana ripening and peaked at the ripening stage followed by a decrease, which is presumably caused by the transcript abundances of carotenoid biosynthetic genes MaLCYB1.1 and MaLCYB1.2. Moreover, a ripening-inducible transcription factor MaSPL16 was characterized, which was shown to be a nuclear protein with transactivation activity. Transient transformation of MaSPL16 in banana fruit led to enhanced transcript levels of MaLCYB1.1 and MaLCYB1.2, and hence the total carotenoid accumulation. Importantly, MaSPL16 stimulated the transcription of MaLCYB1.1 and MaLCYB1.2 through directly binding to their promoters. Collectively, our findings indicate that MaSPL16 behaves as an activator to modulate banana carotenoid biosynthesis, which may provide a new target for molecular improvement of the nutritional and bioactive qualities of agricultural crops that accumulate carotenoids.
Bioactive amines are found in food and can be relevant for the assessment of fruits shelf life and nutritional quality. The pulp and peel of 20 banana and plantain were analyzed and the bioactive amine content varied according to the genotype, ripening stage, fruit tissue and thermal processing. In most of the analyzed genotypes, tyramine, histamine, dopamine, serotonin, spermidine, and spermine were decreased during the ripening process in the pulps. By contrast, there was an increase in putrescine level. In many genotypes of plantains, the serotonin and dopamine contents in pulp decreased until stage 5 and increased at stage 7. Peels contain higher levels of serotonin, dopamine, histamine and tyramine than pulps. Additionally, thermal processing affects the content of amines present in fruit. Boiling with the peel should be preferred in domestic preparations, regardless of the genotype used.
To gain a better understanding concerning factors underlying carotenoid metabolism in banana pulp we investigated the carotenoid profile, metabolome, proteome and relative expression levels of carotegenesis-associated genes of fruit pulp in the two banana varieties ON and GN, with ON being characterized of high carotenoid accumulation. Results showed that high carotenoid content in banana pulp was well correlated with the relative expression of carotenogenesis-associated genes and the abundance of the corresponding proteins. An elevated accumulation of sugar metabolism-related compounds and a decreased amino acid accumulation were also observed in ON. Additionally proteins involved in the glycolytic pathway were more highly abundant in ON suggesting that this supports the higher accumulation of carotenoid in this genotype. We suggest that up-regulated expression of carotenogenesis-associated genes alongside elevated carbohydrate accumulation contribute to high carotenoid content in banana pulp, implying that a multi-target approach is necessary in order to improve carotenoid content in banana.
This study is part of an extensive project that evaluated the effects of a natural ecosystem on a healthy banana crop and the quality of its fruit. In particular, the study examined the influence of the maintenance of natural biodiversity (Atlantic forest) near a conventional banana crop on the metabolic profiling of ripe banana fruits. Results revealed differences between ripe fruits harvested from plants near the natural forest (Near-NF) and fruits harvested in areas distant from the natural forest (Distant-NF). A total of 76 metabolites were identified in ripe banana fruits. Bananas harvested from Near-NF plot showed increased levels of γ-aminobutyric acid and reduced levels of putrescine compared with fruits from Distant-NF plot. Furthermore, fatty acids profile suggests that ripe fruits harvested from Near-NF plot had increased nutritional quality compared with fruits from Distant-NF plot. Multivariate statistical analysis revealed that these metabolites, which potentially influence the nutritional/sensorial quality of ripe fruits, also contributed to distinguishing fruits harvested from Near-NF and Distant-NF plots. Collectively, the results suggest that the natural biodiversity surrounding a crop area could benefit ripe banana nutritional/sensorial quality. The maintenance of natural forest fragments thus appears to be a promising tool for increasing the quality of fruit crops.
Banana is considered as a potential source of valuable nutraceutical bioactive compounds. In addition, It is not only considered as a fruit crop but also serve as high energy containing food in various parts of the world. The aim of this review is to evaluate the historical record, uses, biological activities, chemical composition and compositional changes during ripening behavior in both edible and non-edible part of banana. Recent studies have shown that banana as raw materials riches in valuable bioactive compounds including as vitamins, phytosterols, biogenic amines, phenolics, carotenoids, volatile compounds, minerals, starch and carbohydrates which are highly required in the diet as they play important role in the maintenance of human health and well-being. This paper also covers the potential biological activities of banana such as antidiarrheal, antiulcerative, antimicrobial, antioxidant, hypoglycemic, wound healing, antilithiatic and anticancer activity. It also provide an outline of chemical constituents of banana named vitamins, polyphenols, steroids, triterpenes and amines, carotenoids, starch and carbohydrates, volatile compounds and mineral contents. Furthermore, with the development of fruit these beneficial bioactive compounds vary rapidly because of adopting atmospheric and standardized post and pre-harvested conditions. The mainstream sectors responsible for these changes are highlighted.
Carotenoids are a diverse group of tetraterpenoid pigments that play indispensable roles in plants and animals. The biosynthesis of carotenoids in plants is strictly regulated at the transcriptional and post-transcriptional levels in accordance with inherited genetic signals, developmental requirements, and in response to external environmental stimulants. The alteration in the biosynthesis of carotenoids under the influence of external environmental stimulants, such as high light, drought, salinity, and chilling stresses, has been shown to significantly influence the nutritional value of crop plants. In addition to these stimulants, several pre- and post-harvesting cultivation practices significantly influence the carotenoid composition and contents. Thus, this review discusses how various environmental stimulants and pre- and post-harvesting factors can be positively modulated for the enhanced biosynthesis and accumulation of carotenoids in the edible parts of crop plants, such as leaves, roots, tubers, flowers, fruit, and seeds. In addition, future research directions in this context are identified.
Seed storability is an important trait for crop breeding, however, the mechanism underlying seed storability remains largely unknown. Here, a mass spectrometry-based comparative metabolomic study was performed for rice seeds before and after 24-month natural storage between two hybrid rice cultivars, IIYou 998 (IIY) with low storability and BoYou 998 (BY) with relative high storability. A total of 48 metabolites among 90 metabolite peaks detected were conclusively identified, and most of them are involved in the primary metabolism. During the 24-month storage, 19 metabolites with significant changes in abundance were found in the storage-sensitive IIY seeds, but only 8 in the BY seeds, most of which are free amino acids and soluble sugars. The observed changes of the metabolites in IIY seeds that are consistent with our protoemics results are likely to be involved in its sensitivity to storage. Levels of all identified 18 amino acid-related metabolites and most sugar-related metabolites were significantly higher in IIY seeds both before and after storage. However the level of raffinose was lower in IIY seeds before and after storage, and did not change significantly throughout the storage period in both two cultivars, suggesting its potential role in seed storability. Taken together, these results may help to improve our understanding of seed storability.
Banana flour has been recognized as functional ingredient, owing to its healthy nutritional pattern. Nevertheless, unripe and ripe banana flours show different characteristics and scarce information is available about changes undergone during banana ripening. This study evaluates the changes on physiochemical (chemical composition, hydration properties, rheological properties and structural characteristic) and nutritional (resistant starch content, phenolic compound and antioxidant activity) characteristics of banana flour at the initial four ripening stages. The significant increase in protein content and decrease in carbohydrate and apparent amylose content, besides the reduction in pasting properties, between 2nd and 3rd stages suggested a climacteric peak during ripening. Between those stages, a significant decrease in total and resistant starch was produced together with an increase in total phenolic content and antioxidant activity. Therefore, the knowledge of the physicochemical and nutritional characteristics of banana flour at each ripening stage allows better selection depending on the industrial application.
Banana is a tropical fruit widely accepted by people over the world. Its chemical composition is critical for its organoleptic properties and nutritional value. In this work, the metabolite changes during postharvest senescence were investigated using NMR spectroscopy. The 1D and 2D NMR spectroscopic information revealed the primary and secondary metabolites in banana fruit, including organic acids, amino acids, carbohydrates and phenolics. Bananas at five senescence stages showed similar chemical profiles, but the levels of the individual compounds varied to a large extent. The principal metabolites responsible for postharvest senescence of banana were valine, alanine, aspartic acid, choline, acetate, glucose, malic acid, gallic acid and dopamine. At stage V, ethanol was present due to the conversion of glucose. Salsolinol was generated due to the conversion of dopamine. This was a characteristic marker for the postharvest senescence of banana fruit.
Banana is a very popular fruit in the world market and is consumed as staple food in many countries. It is grown worldwide and constitutes the fifth most important agricultural food crop in terms of world trade. It has been classified into the dessert or sweet bananas and the cooking bananas or plantains. It is either eaten raw or processed, and also as a functional ingredient in various food products. Banana contains several bioactive compounds, such as phenolics, carotenoids, biogenic amines and phytosterols, which are highly desirable in the diet as they exert many positive effects on human health and well-being. Many of these compounds have antioxidant activities and are effective in protecting the body against various oxidative stresses. In the past, bananas were effectively used in the treatment of various diseases, including reducing the risk of many chronic degenerative disorders. In the present review, historical background, cultivar classification, beneficial phytochemicals, antioxidant activity and health benefits of bananas are discussed.
The plant metabolome is the readout of plant physiological status and is regarded as the bridge between the genome and the phenome of plants. Unraveling the natural variation and the underlying genetic basis of plant metabolism has received increasing interest from plant biologists. Enabled by the recent advances in high-throughput profiling and genotyping technologies, metabolite-based genome-wide association study (mGWAS) has emerged as a powerful alternative forward genetics strategy to dissect the genetic and biochemical bases of metabolism in model and crop plants. In this review, recent progress and applications of mGWAS in understanding the genetic control of plant metabolism and in interactive functional genomics and metabolomics are presented. Further directions and perspectives of mGWAS in plants are also discussed.
Copyright © 2015 Elsevier Ltd. All rights reserved.
Carotenoids are mostly C40 terpenoids, a class of hydrocarbons that participate in various biological processes in plants, such as photosynthesis, photomorphogenesis, photoprotection, and development. Carotenoids also serve as precursors for two plant hormones and a diverse set of apocarotenoids. They are colorants and critical components of the human diet as antioxidants and provitamin A. In this review, we summarize current knowledge of the genes and enzymes involved in carotenoid metabolism and describe recent progress in understanding the regulatory mechanisms underlying carotenoid accumulation. The importance of the specific location of carotenoid enzyme metabolons and plastid types as well as of carotenoid-derived signals is discussed.
Copyright © 2015 The Author. Published by Elsevier Inc. All rights reserved.
Bananas and plantains (Musa spp.) are a staple food for millions of impoverished people and as such are an important source of vitamins and micronutrients. To evaluate the potential of Musa spp. to meet dietary micronutrients requirements, we have screened 171 different genotypes for fruit provitamin A carotenoids (pVACs) contents, and a subset of 47 genotypes for macro- and micro-mineral (iron and zinc) contents using standardised sampling and analytical protocols. The results indicate that there is substantial variability in mean fruit pulp pVACs contents between cultivars, and that cultivars with a high fruit pVACs content are widely distributed across the different genome groups but only at a low frequency. The introduction of such high pVACs cultivars has much potential for improving the vitamin A nutritional status of Musa-dependent populations at modest and realistic fruit-consumption levels. In contrast, fruit pulp mineral micronutrient contents (iron and zinc), were low and showed limited inter-cultivar variability, even for genotypes grown under widely-differing environments and soil types. Results are discussed within the framework of the development of strategies to improve the nutritional health and alleviation of micronutrient deficiencies within Musa-consuming population groups.
Green bananas were harvested at the full three-quarter stage, conditioned in polyethylene and stored for 35 days at 16 ± 1 °C and 85% relative humidity. Peel colour changed with time. The yellow colour ideal for consumption was achieved at 21 days and, after 28 days, brown specks started to appear. There was a significant increase in the pulp-to-peel ratio. The green fruit had high starch and low soluble sugars levels. Starch levels decreased significantly throughout ripening. At the seventh day of storage sucrose was prevalent, however, fructose and glucose levels increased while sucrose remained constant, decreasing after 28 days. Starch loss followed a first order reaction. Formation of glucose and fructose followed zero order kinetics with higher rate for fructose. The bioactive amines detected were putrescine, spermidine and serotonin. Serotonin decreased significantly after the 14th day of storage. Putrescine levels were similar up to 21 days and decreased significantly thereafter.
Sweetpotato (Ipomoea batatas Lam.) is an important industrial crop and source of food that contains useful components, including antioxidants such as carotenoids. β-Carotene hydroxylase (CHY-β) is a key regulatory enzyme in the beta-beta-branch of carotenoid biosynthesis and it catalyzes hydroxylation into both β-carotene to β-cryptoxanthin and β-cryptoxanthin to zeaxanthin. To increase the β-carotene content of sweetpotato through the inhibition of further hydroxylation of β-carotene, the effects of silencing CHY-β in the carotenoid biosynthetic pathway were evaluated. A partial cDNA encoding CHY-β was cloned from the storage roots of orange-fleshed sweetpotato (cv. Shinhwangmi) to generate an RNA interference-IbCHY-β construct. This construct was introduced into cultured cells of white-fleshed sweetpotato (cv. Yulmi). Reverse transcription-polymerase chain reaction analysis confirmed the successful suppression of IbCHY-β gene expression in transgenic cultured cells. The expression level of phytoene synthase and lycopene β-cyclase increased, whereas the expression of other genes showed no detectable change. Down-regulation of IbCHY-β gene expression changed the composition and levels of carotenoids between non-transgenic (NT) and transgenic cells. In transgenic line #7, the total carotenoid content reached a maximum of 117 μg/g dry weight, of which β-carotene measured 34.43 μg/g dry weight. In addition, IbCHY-β-silenced calli showed elevated β-cryptoxanthin and zeaxanthin contents as well as high transcript level P450 gene. The 2,2-diphenyl-1-picrylhydrazyl radical scavenging activity (DPPH) in transgenic cells was more than twice that in NT cells. RNA-IbCHY-β calli increased abscisic acid (ABA) content, which was accompanied by enhanced tolerance to salt stress. In addition, the production of reactive oxygen species measured by 3,3'-diaminobenzidine (DAB) staining was significantly decreased in transgenic cultured cells under salt stress. Taken together, the present results indicate that down-regulation of IbCHY-β increased β-carotene contents and total carotenoids in transgenic plant cells and enhanced their antioxidant capacity.
The effect of fruit maturation on changes in carotenoids, flavonoids, total soluble reducing equivalents, phenolic acids, ascorbic acid, and antioxidant activity (AOX) in different pepper types (Capsicum annuum, Capsicum frutescens, and Capsicum chinese) was determined. Generally, the concentration of these chemical constituents increased as the peppers reached maturity. Peppers contained high levels of L-ascorbic acid and carotenoids at maturity, contributing 124-338% of the RDA for vitamin C and 0.33-336 RE/100 g of provitamin A activity, respectively. Levels of phenolic acids, capxanthin, and zeaxanthin generally increased during maturation, whereas the level of lutein declined. Flavonoid concentrations varied greatly among the pepper types analyzed and were negatively correlated to AOX under the conditions of the beta-carotene-linoleic assay. Model systems were used to aid in understanding the relationship between flavonoids and AOX. Significant increases in AOX were observed in pepper juice models in response to increasing dilution factors and the presence of EDTA, indicating a pro-oxidant effect due to metal ions in the system. In vitro models demonstrated that increasing levels of flavonoids in combination with constant levels of caffeic and ascorbic acid gave a resultant AOX that was either additive of the two compounds or competitive in their ability to scavenge peroxyl radicals. The model systems were in good agreement with the chemical composition of the pepper cultivars and reflected the interactions affecting AOX. More research is needed to understand the complex interactions that occur among various antioxidants present in pepper extracts.
Carotenoids are lipid-soluble pigments found in many vegetable crops that are reported to have the health benefits of cancer and eye disease reduction when consumed in the diet. Research shows that environmental and genetic factors can significantly influence carotenoid concentrations in vegetable crops, and that changing cultural management strategies could be advantageous, resulting in increased vegetable carotenoid concentrations. Improvements in vegetable carotenoid levels have been achieved using traditional breeding methods and molecular transformations to stimulate biosynthetic pathways. Postharvest and processing activities can alter carotenoid chemistry, and ultimately affect bioavailability. Bioavailability data emphasize the importance of carotenoid enhancement in vegetable crops and the need to characterize potential changes in carotenoid composition during cultivation, storage and processing before consumer purchase.