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Maceration, Soxhlet Extraction, and Steam Hydrodistillation of Anethum graveolens Seeds: Phytochemical Profiles and Cytotoxic Effects
Open accessAbstract
Dill, Anethum graveolens L., is a culinary plant used in traditional medicine to treat gastrointestinal conditions; its seeds have been previously reported to exhibit antimicrobial, anti-inflammatory, hypolipidemic, anticancer and antioxidant activities. This study aimed to assess dill seed extracts and essential oil in terms of chemical composition and cytotoxic effects, combined with an exploration of the potential underlying mechanisms. Four hydroethanolic seed extracts were prepared using Soxhlet extraction or maceration and the essential oil (AGEO) was prepared by steam distillation. Extracts were physicochemically analysed via LC-MS and spectrophotometry while the AGEO composition was determined through GC-MS. Their cytotoxic effects were assessed in A375, PANC-1 and SK-OV-3 cancer cells, while HaCaT keratinocytes were used as non-cancerous control. The mechanistic investigations included immunofluorescence assay, high-resolution respirometry, network pharmacology and molecular docking. Ferulic acid and fisetin were identified as major components of the hydroethanolic extracts, while D-limonene and carvotanacetone were the main compounds in the AGEO. All tested products were revealed to induce dose-dependent cytotoxicity (with IC50 values ranging between 619.8 and 810.2 μg/mL), diminished OXPHOS efficiency and a mitochondrial uncoupling effect. Morphological changes consistent with apoptosis were recorded, particularly in PANC-1 and A375 cells. Network pharmacology suggested a multitarget mechanism for EO that involved STAT3 and HSP90AA1, while the molecular docking calculations indicated that D-limonene and carvotanacetone could be accommodated within the ATP-binding pocket of Hsp90α.
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TL;DR
Hydroethanolic extracts and essential oil from *Anethum graveolens* seeds showed dose-dependent cytotoxicity in A375, PANC-1, and SK-OV-3 cancer cells, with AGEO generally strongest and PANC-1 most sensitive. Mechanistic assays indicated reduced OXPHOS efficiency, mitochondrial uncoupling/depolarization, apoptosis-like morphology, and computational support for HSP90α as a candidate AGEO target.
Key findings
- Extraction yields for hydroethanolic extracts ranged from 8.8% to 9.4%; AGEO yield was much lower. LC-MS showed A2 (maceration, 60% ethanol) had the highest total targeted polyphenolics (~6.03 µg/mg d.e.), while A3 (Soxhlet, 40% ethanol) was highest for sinapic acid (~3.27 µg/mg d.e.) and A4 (Soxhlet, 60% ethanol) for chlorogenic acid (1.99 µg/mg d.e.).
- Spectrophotometric contents ranged from 169.4 ± 0.5 to 177.3 ± 0.5 mg GAE/g extract for TPC, 5.91 ± 0.02 to 12.85 ± 0.05 mg QE/g extract for TFC, and 4.43 ± 0.03 to 9.05 ± 0.06 mg CE/g extract for CTC; A4 had the highest TPC, TFC, and CTC.
- AGEO contained carvotanacetone (60.41%) and D-limonene (38.70%), cumulatively 99.11% of the chromatographic area. Cytotoxicity was dose-dependent; IC50 values ranged from 619.8 to 810.2 µg/mL, with PANC-1 A2 IC50 = 619.8 µg/mL, A1 = 763.1 µg/mL, A4 = 810.2 µg/mL, and SK-OV-3 A2 = 776.1 µg/mL. Highest selectivity indices were PANC-1 A2 (3.138), A1 (3.117), and A4 (2.613).
- Mechanistic assays showed decreased OXPHOS and increased State 2/State 4 respiration consistent with mitochondrial uncoupling; JC-1 indicated depolarization, strongest in PANC-1 with AGEO (0.522 ± 0.06 vs. control 1). Network pharmacology identified 35 common targets and a 28-node PPI network with STAT3 and HSP90AA1 as key; docking gave D-limonene and carvotanacetone −7.2 kcal/mol each versus co-crystallised EOR −8.6 kcal/mol.
Methods
- Four hydroethanolic seed extracts were prepared by maceration or Soxhlet extraction using 40% or 60% ethanol (A1–A4), and AGEO was obtained by steam hydrodistillation from a single authenticated batch of dried *A. graveolens* seeds collected in Timiș county in September 2024.
- Phytochemical profiling used LC-MS for 18 polyphenols, spectrophotometric assays for TPC/TFC/CTC, and GC-MS for AGEO. Biological testing used A375, PANC-1, SK-OV-3 cancer cells and HaCaT keratinocytes with Alamar Blue viability, immunofluorescence, high-resolution respirometry, and JC-1 MMP assays.
- Statistical analysis used one-way ANOVA with Dunnett’s test for viability and two-way ANOVA with Bonferroni’s post-test for respirometry; network pharmacology used GeneCards, SwissTargetPrediction, PharmMapper, STRING, Cytoscape/CytoHubba, and Metascape, followed by AutoDock Vina/PyRx docking against Hsp90α (PDB 6LR9).
Limitations
- The paper states that variable chemical composition depending on plant origin, developmental stage, harvesting, processing, and storage limits extrapolation to other *A. graveolens* preparations; it also states that computational analyses are predictive and require experimental validation, that direct Hsp90α involvement was not experimentally demonstrated, and that apoptotic markers and comparisons of carvotanacetone/D-limonene with AGEO are still needed.
- The text makes evident additional constraints: extractions were performed once, GC-MS identification/quantification did not use authentic reference standards, and AGEO concentrations were expressed as volume percentages, so they cannot be directly compared with mass concentrations of dry extracts.
- The mechanistic mitochondrial experiments used 720 µg/mL for A2/A4 and 0.05% AGEO, doses near or above the observed IC50 range; the paper itself notes that lower doses should be tested to clarify dose–response and distinguish pharmacological mechanisms from high-dose toxicity.
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113 Own result · 38 Check pending · 38 Own interpretation · 15 Own method · 10 No full text of the sources · 5 Not checked yet
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“while on SK-OV-3 cell line, only A2 had an IC50 value under the tested concentration range (776.1 μg/mL)—Table 7.”
§ Maceration, Soxhlet Extraction, and Steam Hydrodistillation of Anethum graveolens Seeds: Phytochemical Profiles and Cytotoxic Effects > 2. Results > 2.5. The Effects of A. graveolens Extracts and AGEO on Cell Viability
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“On the PANC-1 cell line, all the tested extracts inhibited cell proliferation at 720 and 1000 μg/mL, the latter causing the following drop in cell viability: 39.01% ± 6.50 (A1), 28.37 ± 6.00 (A2), 49.03% ± 4.54 (A3) and 30.80% ± 6.22 (A4).”
§ Maceration, Soxhlet Extraction, and Steam Hydrodistillation of Anethum graveolens Seeds: Phytochemical Profiles and Cytotoxic Effects > 2. Results > 2.5. The Effects of A. graveolens Extracts and AGEO on Cell Viability
plainstates a cause- Asserted: passes — The authors put it forward as true.
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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“LC-MS analysis of A. graveolens L. seed extracts via SIM in the negative ion mode revealed a profile dominated mainly by phenolic acids.”
§ Maceration, Soxhlet Extraction, and Steam Hydrodistillation of Anethum graveolens Seeds: Phytochemical Profiles and Cytotoxic Effects > 2. Results > 2.2. LC-MS Polyphenolic Profile of A1–A4
plain- Asserted: passes — The authors put it forward as true.
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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The calculated IC50 values reveal that on PANC-1 cell line, A1 (763.1 μg/mL), A2 (619.8 μg/mL) and A4 (810.2 μg/mL) were the most active extracts, while on SK-OV-3 cell line, only A2 had an IC50 value under the tested concentration range (776.1 μg/mL)—Table 7.”
§ Maceration, Soxhlet Extraction, and Steam Hydrodistillation of Anethum graveolens Seeds: Phytochemical Profiles and Cytotoxic Effects > 2. Results > 2.5. The Effects of A. graveolens Extracts and AGEO on Cell Viability
plain- Asserted: passes — The authors put it forward as true.
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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In PANC-1 cells, treatment with A2, A4 and AGEO altered mitochondrial respiration in a similar pattern to that observed in the A375 cell line, thus suggesting the presence of the uncoupling effect together with inhibition of active respiration.”
§ Maceration, Soxhlet Extraction, and Steam Hydrodistillation of Anethum graveolens Seeds: Phytochemical Profiles and Cytotoxic Effects > 2. Results > 2.7. Evaluation of Mitochondrial Respiration
plainstates a cause- Asserted: passes — The authors put it forward as true.
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Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
§ Maceration, Soxhlet Extraction, and Steam Hydrodistillation of Anethum graveolens Seeds: Phytochemical Profiles and Cytotoxic Effects > 2. Results > 2.9. Target Identification and Network Pharmacology of AGEO in Pancreatic Cancer
“Based on the network topology characteristics summarised in Table 7, STAT3 and HSP90AA1 were selected as potential primary targets for AGEO”
§ Maceration, Soxhlet Extraction, and Steam Hydrodistillation of Anethum graveolens Seeds: Phytochemical Profiles and Cytotoxic Effects > 2. Results > 2.9. Target Identification and Network Pharmacology of AGEO in Pancreatic Cancer
“Among the highly connected nodes in the network, depicted in blue and dark green boxes, were STAT3, HSP90AA1, TNF, PPARG and TLR4 (Figure 11A and Table 9)”
Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
“The remaining 28 nodes were used to construct a protein–protein interaction network with 160 edges”
§ Maceration, Soxhlet Extraction, and Steam Hydrodistillation of Anethum graveolens Seeds: Phytochemical Profiles and Cytotoxic Effects > 2. Results > 2.9. Target Identification and Network Pharmacology of AGEO in Pancreatic Cancer
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Cited by
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Metadata
- Authors
- Goldiș, C., Racoviceanu, R., Negrea-Ghiulai, R., Mioc, A., Prodea, A., Atyim, E., Maksimovic, T., Lukinich-Gruia, A. T., Pricop, M.-A., & Șoica, C.
- Affiliations
- Victor Babeș University of Medicine and Pharmacy Timișoara (RO)
- Faculty of Pharmacy, “Victor Babeș” University of Medicine and Pharmacy, 2 Eftimie Murgu Square, 300041 Timisoara, Romania
- Spitalul Clinic Judeţean de Urgenţă "Pius Brînzeu" Timişoara (RO)
- Politehnica University of Timisoara (RO)
- Corresponding author
- Racoviceanu, Roxana
- ORCID
- Venue
- Molecules
- Publisher
- MDPI AG
- Published
- 2026-09-20
- Volume
- 31
- Issue
- 18
- Pages
- 3337
- ISSN
- 1420-3049 (ISSN-L)
- PMC
- PMC13609506
- OpenAlex
- W7213948843
- Cited by
- 0
- References
- 77
- FWCI
- 0.00
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- Molecules (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion)
- Funders
- “Victor Babes” University of Medicine and Pharmacy Timisoara
- Topics
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Plant chemical constituents analysis
- Life Sciences › Agricultural and Biological Sciences › Food Science › Essential Oils and Antimicrobial Activity
- Physical Sciences › Chemistry › Organic Chemistry › Phytochemical Studies and Bioactivities
- Keywords
- dill, dill seeds, essential oils, cytotoxicity, ferulic acid, fisetin, D-limonene, mitochondrial uncoupling, apoptosis, STAT3, HSP90AA1, anticancer activity, phytochemical composition, cancer cell lines
- MeSH terms
- Antineoplastic Agents, Phytogenic, Humans, Oils, Volatile, Plant Extracts, Seeds, Steam, Apoptosis, Anethum graveolens, Cell Line, Tumor, Distillation, Molecular Docking Simulation, Phytochemicals
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