Metabolomics reveals significant variations in metabolites and correlations regarding the maturation of walnuts (Juglans regia L.)

ABSTRACT The content of walnut metabolites is related to its nutritive value and physiological characteristics, however, comprehensive information concerning the metabolome of walnut kernels is limited. In this study we analyzed the metabolites of walnut kernels at five developmental stages from filling to ripening using GC-MS-based untargeted metabolomics; of a total 252 peaks identified, 85 metabolites were positively identified. Further statistical analysis revealed that these 85 metabolites covered different types of metabolism pathways. PCA scores revealed that the metabolic compositions of the embryo are different at each stage, while the metabolic composition of the endotesta could not be significantly separated into distinct groups. Additionally, 7225 metabolite-metabolite correlations were detected in walnut kernel by a Pearson correlation coefficient approach; during screening of the calculated correlations, 463 and 1047 were determined to be significant with r2≥0.49 and had a false discovery rate (FDR) ≤0.05 in endotesta and embryo, respectively. This work provides the first comprehensive metabolomic study of walnut kernels and reveals that most of the carbohydrate and protein-derived carbon was transferred into other compounds, such as fatty acids, during the maturation of walnuts, which may potentially provide the basis for further studies on walnut kernel metabolism. Summary: This work provides a comprehensive metabolomic study of walnut kernels and reveals most of the carbohydrate and protein-derived carbons were transferred into other compounds during the maturation of walnuts.

• Publication year

  2016

• Venue

  Biology Open

• Publication date

  2016-05-23

• Fields of study

  Agricultural and Food Sciences, Medicine, Biology

• Identifiers
  DOI 10.1242/bio.017863PMID 27215321PMCID 4920193
• External record

  Open on Semantic Scholar

• Source metadata

  Semantic Scholar, PubMed

• No claims are published for this paper.

• No concepts are published for this paper.

• Content of carotenoids, tocopherols, sterols, triterpenic and aliphatic alcohols, and volatile compounds in six walnuts (Juglans regia L.) varieties.

  2015cited by this paper
• Integration of metabolomics data into metabolic networks

  2015cited by this paper
• Amino Acid Composition, Molecular Weight Distribution and Gel Electrophoresis of Walnut (Juglans regia L.) Proteins and Protein Fractionations

  2014cited by this paper
• Metabolic engineering of microorganisms to produce omega-3 very long-chain polyunsaturated fatty acids.

  2014cited by this paper
• Seasonal variation of the main individual phenolics and juglone in walnut (Juglans regia) leaves

  2014cited by this paper
• Chemical composition, molecular weight distribution, secondary structure and effect of NaCl on functional properties of walnut (Juglans regia L) protein isolates and concentrates

  2014cited by this paper
• Comparative metabolomics in primates reveals the effects of diet and gene regulatory variation on metabolic divergence

  2014cited by this paper
• Ameliorating Effects of Exogenously Applied Proline on Seed Composition, Seed Oil Quality and Oil Antioxidant Activity of Maize (Zea mays L.) under Drought Stress

  2013cited by this paper
• HPLC determination of phenolic acids, flavonoids and juglone in walnut leaves.

  2013cited by this paper
• Jasmonates: biosynthesis, perception, signal transduction and action in plant stress response, growth and development. An update to the 2007 review in Annals of Botany.

  2013cited by this paper
• Relationship between asparagine metabolism and protein concentration in soybean seed

  2012cited by this paper
• Composition, Structure and Functional Properties of Protein Concentrates and Isolates Produced from Walnut (Juglans regia L.)

  2012influential reference
• Fatty acids, tocopherols, selenium and total carotene of pistachios (P. vera L.) from Diyarbakır (Southestern Turkey) and walnuts (J. regia L.) from Erzincan (Eastern Turkey)

  2012cited by this paper
• Preparation of fatty acid methyl esters from hazelnut, high-oleic peanut and walnut oils and evaluation as biodiesel

  2012cited by this paper
• Dynamic metabonomic responses of tobacco (Nicotiana tabacum) plants to salt stress.

  2011cited by this paper
• Subfunctionalization of Cellulose Synthases in Seed Coat Epidermal Cells Mediates Secondary Radial Wall Synthesis and Mucilage Attachment1[C][W][OA]

  2011cited by this paper
• Metabolic profiling of strawberry (Fragaria x ananassa Duch.) during fruit development and maturation.

  2011cited by this paper
• Walnut (Juglans regia L.): genetic resources, chemistry, by-products.

  2010cited by this paper
• CELLULOSE SYNTHASE 9 Serves a Nonredundant Role in Secondary Cell Wall Synthesis in Arabidopsis Epidermal Testa Cells 1 [ C ] [ W ] [ OA ]

  2010cited by this paper
• CELLULOSE SYNTHASE9 Serves a Nonredundant Role in Secondary Cell Wall Synthesis in Arabidopsis Epidermal Testa Cells1[C][W][OA]

  2010cited by this paper
• Seed coat mucilage cells of Arabidopsis thaliana as a model for plant cell wall research

  2010cited by this paper
• Oxidative stability of tree nut oils.

  2008cited by this paper
• Fatty acid unsaturation, mobilization, and regulation in the response of plants to stress

  2008cited by this paper
• Jasmonates: an update on biosynthesis, signal transduction and action in plant stress response, growth and development.

  2007cited by this paper
• Metabolomic alterations in elicitor treated Silybum marianum suspension cultures monitored by nuclear magnetic resonance spectroscopy.

  2007cited by this paper
• Integrated Analysis of Metabolite and Transcript Levels Reveals the Metabolic Shifts That Underlie Tomato Fruit Development and Highlight Regulatory Aspects of Metabolic Network Behavior1[W]

  2006cited by this paper
• A walnut diet improves endothelial function in hypercholesterolemic subjects: a randomized crossover trial, E. Ros, I. Núñez, A. Pérez-Heras, M. Serra, R. Gilabert, E. Casals, R. Deulofeu, in: Circulation, 109. (2004), 1609

  2005cited by this paper
• Soluble methionine enhances accumulation of a 15 kDa zein, a methionine-rich storage protein, in transgenic alfalfa but not in transgenic tobacco plants.

  2005cited by this paper
• A gas chromatography/electron ionization-mass spectrometry-selected ion monitoring method for determining the fatty acid pattern in food after formation of fatty acid methyl esters.

  2005cited by this paper
• A regulatory role for proline metabolism in stimulating Arabidopsis thaliana seed germination

  2004cited by this paper
• A Walnut Diet Improves Endothelial Function in Hypercholesterolemic Subjects: A Randomized Crossover Trial

  2004cited by this paper
• Process for the integrated extraction, identification and quantification of metabolites, proteins and RNA to reveal their co‐regulation in biochemical networks

  2004cited by this paper
• Antifungal Activities of Four Fatty Acids against Plant Pathogenic Fungi

  2004cited by this paper
• Metabolic engineering of fatty acid biosynthesis in plants.

  2002influential reference
• Oxylipin profiling in pathogen-infected potato leaves.

  2002cited by this paper
• Functionalization of cellulose

  1998cited by this paper
• 3-Methylcrotonyl-coenzyme A carboxylase is a component of the mitochondrial leucine catabolic pathway in plants

  1998cited by this paper
• Asparagine metabolism-key to the nitrogen nutrition of developing legume seeds.

  1975cited by this paper
• A rapid method of total lipid extraction and purification.

  1959cited by this paper

• Targeted lipidomics meets transcriptomics: how cinobufagin rewires fatty acid, sphingolipid, and glycerophospholipid metabolism to combat hepatoma cell growth.

  2026cites this paper
• Targeted metabolomics to quantitatively profile changes in amino acids and phenolics at every step of amahewu production from two Zea Mays L. maize types (white and yellow)

  2026cites this paper
• Effect of Different Organic Amendment Supply on Young Bearing Walnut Trees Nutritional Status and Soil Fertility

  2026cites this paper
• Integrated transcriptome and metabolome analyses reveal that SlEIL4 represses SlALDH2B4 to regulate GABA accumulation in tomato fruit

  2025cites this paper
• Unraveling the influence of lipids on aroma formation during microwave roasting of hazelnuts based on lipidomics

  2025cites this paper
• Defense dynamics in walnut (Juglans regia L.) fruit during walnut husk fly (Rhagoletis completa Cresson) infestation: an integrative, multi-level analysis

  2025cites this paper
• Metabolome and Metagenome Signatures Underlying the Differential Resistance of Percocypris pingi, Crucian Carp, and Yellow Catfish to Ichthyophthirius multifiliis Infection

  2025cites this paper
• Transcriptomic and metabolomic analyses unravel the different pathogenic mechanisms of Ustilaginoidea virens in indica and japonica rice

  2025cites this paper
• Metabolomic Analysis of Cauda Epididymal Fluid in Yaks and Cattle

  2025cites this paper
• The effects of dietary energy level on the growth performance of yaks (Bos grunniens) were studied based on omics technique

  2025cites this paper
• Enhancing Sheep Rumen Function, and Growth Performance Through Yeast Culture and Oxalic Acid Supplementation in a Hemicellulose-Based Diet

  2025cites this paper
• Alterations in serum metabolomics predict drug-resistant epilepsy

  2025cites this paper
• Polyphenol Changes and Antioxidant Effects of Roasted Gurguéia (Dipteryx lacunifera Ducke) Nut during Simulated Digestion In Vitro

  2025cites this paper
• Evaluation of biochemical profiles in walnut (Juglans regia L.) genotypes and predictive modeling using advanced machine learning algorithms

  2025cites this paper
• MD‐2‐related lipid‐recognition protein‐like is involved in mosquito reproduction by affecting lipid metabolism

  2025cites this paper
• The response mechanism of high pH and alkalinity interactive stress on immune system and energy metabolism pathway of Litopenaeus vannamei.

  2025cites this paper
• Multi-omic serum analysis reveals ferroptosis pathways and diagnostic molecular signatures associated with Moyamoya diseases

  2025cites this paper
• Lipidomics and biochemical profiling of adult Yili horses in a 26 km endurance race: exploring metabolic adaptations

  2025cites this paper
• Metabolite profiling of foxtail millet varieties from Qingyang using GC–MS and LC–MS analysis

  2025cites this paper
• Polysaccharide from Atractylodes macrocephala Koidz. alleviates pyrotinib-induced diarrhea through regulating cAMP/LKB1/AMPK/CFTR pathway and restoring gut microbiota and metabolites.

  2025cites this paper
• Integrated metabolome and transcriptome analysis of fulvic acid relieves nitrate stress-induced damage in spinach (Spinacia oleracea L.) by regulating multiple defense pathways

  2025cites this paper
• Gut microbiome and plasma metabolome alterations in ileostomy and after closure of ileostomy

  2025cites this paper
• Dynamic metabolic profiling of sea cucumbers (Apostichopus japonicus) under predation stress.

  2025cites this paper
• Identification of GDP as a small inhibitory molecule in HepG2 cells by non‑targeted metabolomics analysis

  2025cites this paper
• Transcriptome and Metabolome Analysis of Organic and Chemical Fertilizer Effects on Highland Barley Growth and Nutrient Utilization

  2025cites this paper
• Enhanced methanol-xylose co-utilization strategy in Komagataella phaffii.

  2025cites this paper
• The potential of three endemic Alocasia species in Indonesia: Insights from advancing in vitro propagation methods and comparative metabolites analysis

  2025cites this paper
• Metabolomic and transcriptomic analysis of Panax Notoginseng root: Mechanistic insights into the effects of flower bud removal on yield and phytochemical composition

  2025cites this paper
• Integrated Transcriptomic, Proteomic, and Metabolomic Analyses Revealed Molecular Mechanism for Salt Resistance in Soybean (Glycine max L.) Seedlings

  2024cites this paper
• Effects of polystyrene microplastics on the growth and metabolism of highland barley seedlings based on LC-MS

  2024cites this paper
• Acute stress response in gill of Pacific white shrimp Litopenaeus vannamei to high alkalinity

  2024cites this paper
• Mechanism of Atractylenolide Ⅲ alleviating pyrotinib-induced diarrhea by regulating AMPK/CFTR pathway through metabolite of gut microbiota

  2024cites this paper
• Melatonin alleviates heat stress-induced spermatogenesis dysfunction in male dairy goats by regulating arachidonic acid metabolism mediated by remodeling the gut microbiota

  2024cites this paper
• Identification of the serum metabolomic profile for acute ischemic preconditioning in athletes

  2024cites this paper
• Hormonal, biochemical, and genetic regulations of walnut fruit development and ripening: an integrated perspective

  2024influential citation
• Leguminous Green Manure Intercropping Promotes Soil Health in a Citrus (Citrus reticulata) Orchard

  2024cites this paper
• Acute stress response in hepatopancreas of Pacific white shrimp Litopenaeus vannamei to high alkalinity

  2024cites this paper
• Effects of Methionine on Milk Performance and Milk Constituents of Lactating Donkeys

  2024influential citation
• Metabolic characteristics of ischaemic preconditioning induced performance improvement in Taekwondo athletes using LC‒MS/MS-based plasma metabolomics

  2024influential citation
• Combined transcriptomic and metabolomic analysis of the mechanism by which Bacillus velezensis induces resistance to anthracnose in walnut

  2024cites this paper
• Dietary curcumin supplementation enhances growth performance and anti-inflammatory functions by modulating gut microbiota, microbiota-derived metabolites, and expression of inflammation-related genes in broilers.

  2024cites this paper
• Transcriptomic and coexpression network analyses revealed the regulatory mechanism of Cydia pomonella infestation on the synthesis of phytohormones in walnut husks

  2024cites this paper
• Dissection of Targeting Molecular Mechanisms of Celastrol-induced Nephrotoxicity via A Combined Deconvolution Strategy of Chemoproteomics and Metabolomics

  2024influential citation
• Co-expression analyses reveal key Cd stress response-related metabolites and transcriptional regulators in Kentucky bluegrass.

  2024cites this paper
• The flavonoids in root regulated rhizosphere microbiome of Angelica dahurica var. formosana in genuine producing area

  2024cites this paper
• Comparative metabolomics and transcriptomics provide new insights into florpyrauxifen-benzyl resistance in Echinochloa glabrescens

  2024cites this paper
• Effects of Low-Salinity Stress on Histology and Metabolomics in the Intestine of Fenneropenaeus chinensis

  2024cites this paper
• Lupeol improves bile acid metabolism and metabolic dysfunction-associated steatotic liver disease in mice via FXR signaling pathway and gut-liver axis.

  2024cites this paper
• Hippo-YAP/TAZ-ROS signaling axis regulates metaflammation induced by SelenoM deficiency in high-fat diet-derived obesity

  2024cites this paper
• Differential metabolite analysis of the pharmacodynamic differences between different ratios of Dahuang ()-Taoren () herb pair.

  2024cites this paper
• Alterations of intestinal mucosal barrier, cecal microbiota diversity, composition, and metabolites of yellow-feathered broilers under chronic corticosterone-induced stress: a possible mechanism underlying the anti-growth performance and glycolipid metabolism disorder

  2024cites this paper
• Comparative UHPLC-MS/MS-based untargeted metabolomics analysis, antioxidant, and anti-diabetic activities of six walnut cultivars

  2024cites this paper
• Magnesium Nutrient Application Induces Metabolomics and Physiological Responses in Mulberry (Morus alba) Plants

  2023cites this paper
• Analysis of microbial and metabolic diversity in Jiangshui from Northwest China

  2023cites this paper
• Transcriptome and Metabolome Reveal the Molecular Mechanism of Barley Genotypes Underlying the Response to Low Nitrogen and Resupply

  2023cites this paper
• Metabolomics analysis of follicular fluid in ovarian endometriosis women receiving progestin-primed ovary stimulation protocol for in vitro fertilization

  2023cites this paper
• Semen quality and seminal plasma metabolites in male rabbits (Oryctolagus cuniculus) under heat stress

  2023cites this paper
• Renal metabolomic profiling of large yellow croaker Larimichthys crocea acclimated in low salinity waters.

  2023cites this paper
• Metabolomic Analysis of the Response of Haloxylon ammodendron and Haloxylon persicum to Drought

  2023cites this paper
• Mechanism analysis of calcium-induced low freeze-drying survival of probiotic encapsulated in alginate

  2023cites this paper
• Leaf Biochemical and Kernel Metabolite Profiles as Potential Biomarkers of Water Deficit in Walnut (Juglans regia L.) cv. Chandler

  2023cites this paper
• The Effects of Altitude on Fruit Characteristics, Nutrient Chemicals, and Biochemical Properties of Walnut Fruits (Juglans regia L.)

  2023cites this paper
• Seasonal temperature variation in Zostera marina seedlings under ocean acidification

  2023cites this paper
• Postharvest drying of walnuts: Effect of drying methods on walnut quality

  2023cites this paper
• Transcriptomic and Metabolomic Profiling Reveals the Variations in Carbohydrate Metabolism between Two Blueberry Cultivars

  2023cites this paper
• Losartan ameliorates renal interstitial fibrosis through metabolic pathway and Smurfs-TGF-β/Smad.

  2022cites this paper
• Comparative transcriptomic and metabolomic analysis revealed molecular mechanism of two wheat near-isogenic lines response to nitrogen application.

  2022cites this paper
• Exogenous antibiotic resistance gene contributes to intestinal inflammation by modulating the gut microbiome and inflammatory cytokine responses in mouse

  2022cites this paper
• Hyperspectral detection of walnut protein contents based on improved whale optimized algorithm

  2022cites this paper
• Integrated Physiological and Metabolomic Analyses Reveal the Differences in the Fruit Quality of the Blueberry Cultivated in Three Soilless Substrates

  2022cites this paper
• Plasma metabolomics for the assessment of the progression of non-small cell lung cancer

  2022cites this paper
• Integrated Multi-Omic Characterization of the Detachment Process of Adherent Vero Cells with Animal-Based and Animal-Origin-Free Enzymes

  2022cites this paper
• Mechanisms of Ardisia japonica in the Treatment of Hepatic Injury in Rats Based on LC-MS Metabolomics

  2022cites this paper
• The Anti-Cancer Effects of Mitochondrial-Targeted Triphenylphosphonium–Resveratrol Conjugate on Breast Cancer Cells

  2022cites this paper
• Metabolomic and transcriptomic analysis of roots of tobacco varieties resistant and susceptible to bacterial wilt.

  2022cites this paper
• Integration of transcriptomic and metabolomic analysis of the mechanism of dietary N-carbamoylglutamate in promoting follicle development in yaks

  2022cites this paper
• fiSystematic transcriptomic and metabolomic analysis of walnut (Juglans regia L.) kernel to trace variations in antioxidant activity during ripening

  2022cites this paper
• A Comparison of Different Tissues Identifies the Main Precursors of Volatile Substances in Chicken Meat

  2022cites this paper
• Evaluating Starter Feeding on Ruminal Function in Yak Calves: Combined 16S rRNA Sequencing and Metabolomics

  2022cites this paper
• Untargeted metabolomics profiling in a mouse model of lung cancer treated with thermal ablation

  2022cites this paper
• A comprehensive analysis based on histopathology, metabolomics and transcriptomics reveals molecular regulatory mechanisms of the red claw crayfish (Cherax quadricarinatus) in response to acute hepatopancreatic necrosis disease (AHPND)

  2022cites this paper
• Altered Plasma Metabolic Profiles in Chinese Patients With Multiple Sclerosis

  2021cites this paper
• Integration of transcriptomics and metabolomics to identify key coumarin biosynthetic genes in Bupleurum chinense

  2021cites this paper
• Characterization and comparison of lipids in Camellia oleifera kernels of XL210 and XL1 based on LC-MS/MS

  2021cites this paper
• Metabolomic profiling of SQ-1-induced changes in starch metabolism in sterile anthers of wheat

  2021cites this paper
• Identification of Key Volatiles Differentiating Aromatic Rice Cultivars Using an Untargeted Metabolomics Approach

  2021cites this paper
• Effects of green-manure and tillage management on soil microbial community composition, nutrients and tree growth in a walnut orchard

  2021cites this paper
• Integrative metabolomic characterisation identifies altered portal vein serum metabolome contributing to human hepatocellular carcinoma

  2021cites this paper
• Olive Fruit Development and Ripening: Break on through to the “-Omics” Side

  2021cites this paper
• Untargeted Metabolomics Reveals Intestinal Pathogenesis and Self-Repair in Rabbits Fed an Antibiotic-Free Diet

  2021cites this paper
• Sensory and nutritional attributes of Persian walnut kernel influenced by maturity stage, drying method, and cultivar

  2021cites this paper
• Combined Transcriptomic and Metabolomic Analysis Reveals the Role of Phenylpropanoid Biosynthesis Pathway in the Salt Tolerance Process of Sophora alopecuroides

  2021cites this paper
• The effects of cultivar and climate zone on phytochemical components of walnut ( Juglans regia L . )

  2020cites this paper
• Metabolomics Analysis of the Deterioration Mechanism and Storage Time Limit of Tender Coconut Water during Storage

  2020cites this paper
• Comparative Proteomic Analysis of Walnut (Juglans regia L.) Pellicle Tissues Reveals the Regulation of Nut Quality Attributes

  2020cites this paper
• Metabolomics reveals the effect of valproic acid on MCF-7 and MDA-MB-231 cells

  2020cites this paper
• Food Profiling: Determination of the Geographical Origin of Walnuts by 1H NMR Spectroscopy Using the Polar Extract.

  2020cites this paper
• Characterization of iron walnut in different regions of China based on phytochemical composition

  2020influential citation
• Non-targeted metabolomics of quinoa seed filling period based on liquid chromatography-mass spectrometry.

  2020cites this paper
• Characterization of bioactive and bioenergy components from fresh walnut (Juglans regia) leaf

  2020cites this paper