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Title: Borneol: Antimicrobial and Antibacterial Applications and Mechanistic Insights
Borneol's Antimicrobial Activity in Plant Essential Oils and Individual Testing
Borneol Content Variation in Plant Essential Oils and Associated Antimicrobial Activity
Borneol is a widespread oxygenated monoterpene component in plant essential oils (EOs), with content varying significantly across plant species, populations of the same species, phenological stages, harvesting times, and environmental conditions. Below is a consolidated summary of borneol content reported in EOs from diverse plant species and sources:
Borneol Content in Plant Essential Oils Across Species and Sources
| Plant Species/Sample Description | Borneol Content | Citation(s) |
|---|---|---|
| Thymus satureioides EO (Tata site) | 41.3% | [1] |
| Thymus satureioides EO (Azrou site) | 31.7% | [1] |
| Tetraclinis articulata EO | 14.83% | [2] |
| Dittrichia viscosa EO | 9.3% | [3] |
| Rosmarinus officinalis EO | 26.48 wt.% | [4] |
| Plectranthus barbatus EO | 20.7% | [5] |
| Clerodendrum urucurana stem bark EO | 14.7% | [6] |
| Thymus EOs | 0.3–41.3% | [1] |
| Callicarpa ornifolia EO | 2.9% | [7] |
| Thymus satureioides EO | 27% | [8] |
| Crude EO (unspecified species) | 2.7% | [9] |
| Artemisia judaica EO (endo-borneol) | 5.72% | [10] |
| Origanum vulgare subsp. glandulosum EO (early vegetative stage) | 2.38% | [11] |
| Thymus maroccanus EO (post-flowering leaves) | 16.3% | [12] |
| Thymus maroccanus EO (other stages/parts) | 0.2–16.3% | [12] |
| Satureja satureioides EO | 12.4% | [13] |
| Satureja cuneifolia EO samples | 12.9–24% | [14] |
| EO (unspecified species, major component) | 21.83% | [15] |
| Poldokhtar EO | 7.1% | [16] |
| Oil type III (unspecified species) | 2.6% | [17] |
| T. satureioides EO | 34.26% | [18] |
| E. fimbriobracteata EO | 8.1% | [19] |
| Dried leaf EO (unspecified species) | 3.1% | [20] |
| T. parthenium leaf EO | 2.9% | [21] |
| T. punctatum leaf EO | 2.1% | [21] |
| S. veneris EO | 6.2% | [22] |
| A. gypsicola EO (post-flowering stage-8 pm) | 22.62% | [23] |
| Thymus arduini EO | 5.4% | [24] |
| EO (unspecified source, with camphor 14.8% and eucalyptol 12.9%) | 10.8% | [25] |
| Xylopia hypolampra stem bark EO | 7.8% | [26] |
| R. officinalis EO | 10.39% | [27] |
| Thymus satureioides EO | 21.2% | [28] |
| Lavandula angustifolia EO | 4.2% | [29] |
| Lavandin EO | 6.1% | [29] |
| EOBD | 11.07% | [30] |
| Lavender EO LA 2019 | 15.67% | [31] |
| Lavender EO LA 2020 | 19.35% | [31] |
| Siddha variety EO | 5.2% | [32] |
| R. beesianus EO | 15.0% | [33] |
| Lavender EO (50 mg·dm⁻³ gold nanoparticles, 'Blue River') | 25.75% | [34] |
| Lavender EO (50 mg·dm⁻³ gold nanoparticles, 'Elegance Purple') | 32.17% | [34] |
| Lavender EO (50 mg·dm⁻³ gold nanoparticles, Munstead) | 13.38% | [34] |
| Kadsura longipedunculata EO | 6.05% | [35] |
| X. strumarium EO | 11.6% | [35] |
| Mandarin peel oil (immature stage) | 1.22% | [36] |
| Kandovan population EO (before flowering) | 13.50% | [37] |
| A. herba-alba EO | 3.3% | [38] |
| Sage EOs | 2.12–18.38% | [39] |
| Achillea odorata subsp. pectinata EO (AOpEO) | 11.33% | [40] |
| EO 3 (unspecified source, (+)-borneol) | 5.80% | [41] |
| Sat. macrosiphonia EO (Poldokhtar, Lorestan Province) | 16.60% | [42] |
| Sat. macrosiphonia EO (Kabirkooh, Ilam Province) | 8.20% | [42] |
| A. vulgaris leaf EO | 8.06% | [43] |
| T. satureioides aerial part EO | 18.1% | [44] |
| S. officinalis EO | 7.6% | [45] |
| T. argyrophyllum var. argyrophyllum flower EO | 12.0% | [46] |
| T. argyrophyllum var. argyrophyllum stem EO | 15.0% | [46] |
| A. ligustica EO | 6.2% | [47] |
| S. subspicata EO | 2.11% | [48] |
| S. corsica EO | 26.2% | [49] |
| B. socotrana EO | Major component | [50] |
| EO (unspecified species) | 2.34% | [51] |
| D. graveolens EO | 18.7% | [52] |
| M. communis L. EO | 27.15% | [53] |
| A. canescens EO | 6.9% | [54] |
| R. beesianus EO | 15.0% | [55] |
| Thyme oil | Main component | [56] |
| O. vulgare L. EO | 6.52% | [57] |
| A. millefolium L. flower EO | 12.4% | [58] |
| T. satureioides EO | 0–27.7% | [59] |
| T. satureioides Coss EO | 21.56% | [60] |
| T. satureioides Coss HE | 25.04% | [60] |
| H. chrysotricha EO | 32.7% | [61] |
| Lavender OE | 5.34% | [62] |
| S. kitaibelii EO | Dominant compound | [63] |
Factors Influencing Borneol Content
Environmental factors and phenological stages modulate borneol accumulation: copper sulphate treatment increases borneol levels in some EOs [64], while nano Zn oxide treatment boosts borneol in A. millefolium EO [65]. Harvest timing affects content—e.g., A. gypsicola EO harvested at post-flowering stage (8 pm) reaches 22.62% borneol [23], and T. polium EO contains 8.24% borneol in winter-collected aerial parts versus 0% in autumn samples [66]. Phenological stages also impact levels: T. parthenium EO has reduced borneol in flowering versus pre-flowering stages [67], and sage EO borneol peaks in the vegetative period (May–June) [39].
Borneol and EO Antimicrobial Activity
Variation in borneol content correlates with EO antimicrobial efficacy. High-borneol Thymus EOs (e.g., 41.3% in T. satureioides from Tata) exhibit bactericidal activity against Gram-negative bacteria like Enterobacter cloacae and Acinetobacter baumannii, sometimes exceeding antibiotic potency [1]. D. viscosa EO (9.3% borneol) acts against Gram-negative (E. coli, P. aeruginosa, K. pneumoniae), Gram-positive (S. aureus) bacteria, fungi (C. albicans), and yeasts (S. cerevisiae) with MICs 0.101–3.25 mg/mL [3]. R. officinalis EO (26.48 wt.% borneol) inhibits Streptococcus equi zooepidemicus, S. aureus, and E. coli below 7.50 mg/mL [4], while P. barbatus EO (20.7% borneol) shows MICs 0.137–0.55 mg/mL against most microbes [5]. C. urucurana stem bark EO (14.7% borneol) is more potent than leaf EO, with MICs 1.25 mg/mL against S. epidermidis and E. coli [6].
Further associations include: T. satureioides EO (34.26% borneol) inhibits mycobacterial strains with inhibition zones 18.5–28.0 mm and MICs 0.015–0.062% (v/v) [18]; A. gypsicola EO (up to 22.62% borneol) has inhibition zones up to 42.3 mm for fungi [23]; and winter-collected T. polium EO (8.24% borneol) is more active than autumn samples (0% borneol) with MIC 5 µl/ml [66]. Recent studies link borneol to broad-spectrum activity: T. arduini EO (5.4% borneol) inhibits bacteria (MICs 6.25–37.50 mg/mL) and fungi (7.81–25.00 mg/mL) [24]; EOBD (11.07% borneol) has inhibition zones 36.40±1.70 mm (S. aureus) and 28.47±1.44 mm (P. aeruginosa) [30]; and lavender EO LA 2020 (19.35% borneol) is more effective than LA 2019 (15.67% borneol) against E. coli and B. subtilis [31].
Borneol interacts synergistically with other components (e.g., carvacrol, thymol, camphor) to enhance activity [1]. A PLS model identifies borneol as a variable important in projection (VIP>1) for C. wenyujin EO antimicrobial properties [68], and chemometric analyses highlight its role in S. kitaibelii EO activity [63].
Individual Borneol’s Antimicrobial Properties
Borneol itself exhibits antimicrobial activity [69][3][70], with bacteriostatic effects against E. coli at 0.5 μl/ml and bactericidal effects against L. monocytogenes (1 μl/ml) and E. coli (2 μl/ml) [71]. Isolated borneol from A. millefolium EO has MICs 2.5±0.05–25.0±0.11 μg/mL (bacteria) and 3.0±0.35–4.5±0.65 μg/mL (fungi) [65]. Its mechanism involves membrane disruption: lipophilic borneol impairs nutrient transfer, metabolic functions, and growth by disrupting cell walls, dissipating ion gradients, and limiting hydrophobic compound diffusion through lipopolysaccharide layers [3][32][33]. Recent insights link borneol to S. aureus membrane invagination and altered cytoplasmic density [49], though it may be inactive against P. aeruginosa [36].
Individual Borneol’s Antimicrobial Efficacy Against Gram-Positive, Gram-Negative, and Fungal Pathogens
Borneol is a monoterpene with documented antimicrobial activity, recognized in traditional Chinese and Indian medicine alongside its derivatives for this property [72][73]. When evaluated individually or as a component of essential oils, borneol exhibits differential efficacy against Gram-positive, Gram-negative, and fungal pathogens, with stronger activity typically observed against Gram-positive bacteria. It also inhibits bacterial adherence to inert substrates at subinhibitory concentrations (MIC/2) [74], and its molecular mechanisms include disrupting bacterial membrane integrity—limiting hydrophobic compound diffusion through lipopolysaccharide layers, dissipating ion gradients, and impairing essential cellular processes to induce cell death—underlying broad-spectrum activity against Gram-positive, Gram-negative, and multi-drug resistant bacteria [32][33]. While borneol has a 31% binding probability to the transient receptor potential cation channel subfamily M member 8 (TRPM8) (its main target for topical analgesia), the role of TRPM8 in its antimicrobial mechanisms requires further exploration [72][73].
Borneol and Borneol-Containing Essential Oil Antimicrobial Activity Against Pathogens
| Pathogen Category | Pathogen Strain | Borneol Form/Source | Key Result | Citation(s) |
|---|---|---|---|---|
| Gram-Positive Bacteria | Listeria monocytogenes 4b | Isolated borneol | Identical inhibitory and bactericidal concentrations | [75] |
| Listeria innocua | Isolated borneol | MIC = 5 mg/ml | [76] | |
| Multiple Gram-positive strains | Isolated borneol | Ranked among active compounds, following thymol/carvacrol | [77] | |
| Staphylococcus aureus, Bacillus subtilis | Isolated borneol | MIC = 6.49 mM (less potent than trans-o-coumaric/trans-cinnamic acid) | [78] | |
| Xanthomonas arboricola pv. pruni (Xap) | Rosemary essential oil component | Responsible for antibacterial effect | [79] | |
| MRSA, vancomycin-resistant E. faecalis | Isolated borneol | Exhibited activity | [35] | |
| S. aureus, E. faecalis | Rosemary essential oil component | Active, but low levels may limit contribution | [80] | |
| L. innocua, B. cereus | Sat. macrosiphonia essential oil (8.20–16.60% borneol) | Largest inhibition zones (30.00 ± 3.46 mm for L. innocua via MAHD) | [42] | |
| B. cereus, S. aureus | A. vulgaris essential oil (8.06% borneol) | MIC = 5 μg/mL | [43] | |
| M. luteus, B. cereus | T. satureioides essential oil (18.1% borneol) | MIC = 2.25–18.4 mg/mL; inhibition zones >20 mm | [44] | |
| S. aureus | I. graveolens essential oil (26.2% borneol) | MIC = 5 mg·ml⁻¹; bactericidal in 2 hours; membrane/cytoplasmic changes | [49] | |
| B. cereus | T. argyrophyllum var. argyrophyllum essential oil (15.0% borneol) | Activity matched chloramphenicol (125 μg/mL) | [46] | |
| E. faecalis, S. aureus, N. asteroides, C. jeikeium | A. ligustica essential oil (6.2% borneol) | Significant activity | [47] | |
| Gram-Negative Bacteria | P. aeruginosa, E. coli | Isolated borneol | MIC = 6.1–6.75 mM (low efficacy) | [78] |
| Klebsiella | Isolated borneol | Weak activity; MIC = 1250 µg/mL | [81] | |
| Xap (G2 vs. Xap3 isolates) | Isolated borneol | More effective against XapG2; MIC = 50–400 µg/mL/nL/mL | [79] | |
| E. coli | Isolated borneol | Moderate activity; inactive against P. aeruginosa | [36] | |
| Sal. typhimurium | Sat. macrosiphonia essential oil (8.20–16.60% borneol) | Smallest inhibition zones (13.17 ± 1.53 mm via MAHD) | [42] | |
| S. typhimurium, E. coli | A. vulgaris essential oil (8.06% borneol) | MIC = 40 μg/mL | [43] | |
| E. coli, P. aeruginosa | D. graveolens essential oil (18.7% borneol) | Slight activity; MIC = 569.4–18220.8 μg/mL | [52] | |
| P. aeruginosa, E. coli | A. ligustica essential oil (6.2% borneol) | Modest activity | [47] | |
| Fungal Pathogens | C. neoformans | Plectranthus barbatus essential oil (20.7% borneol) | Inhibited growth (major component) | [5] |
| Dermatophytes (M. canis, M. gypseum, T. mentagrophytes, T. rubrum) | (–)-borneol (essential oil component) | Part of group with broad activity | [82] | |
| Sclerotium rolfsii | Isolated borneol; LA 2020 lavender oil | Inhibited growth; higher borneol content correlated with greater efficacy | [31] | |
| General fungal pathogens | Isolated borneol | Reported antifungal activity | [83] | |
| C. acutatum, B. cinerea | S. officinalis essential oil (7.6% borneol) | Reduced mycelial growth; MIC = 1800 ppm against C. acutatum | [45] | |
| C. albicans | A. canescens essential oil (6.9% borneol) | Weak activity | [54] |
Mechanistic Insights into Borneol’s Antimicrobial Action: Membrane Disruption and Biofilm Inhibition
Borneol, a monoterpene component of plant essential oils (EOs), exhibits well-documented antimicrobial and antibiofilm activity, primarily mediated by disrupting microbial membrane integrity [32, 163]. Its effects have been evaluated across multiple microbial species, with key findings summarized below:
Borneol’s Antimicrobial and Antibiofilm Activity Across Microorganisms
| Microorganism Group | Specific Microorganism(s) | Activity Type | Concentration/Dosage | Key Outcome | Citation(s) |
|---|---|---|---|---|---|
| Candida albicans | C. albicans | Biofilm inhibition | 0.01% | Inhibits biofilm formation | [84] |
| Candida albicans | C. albicans | Biofilm reduction | 0.005% | Reduces biofilm formation by over 80% | [84] |
| Bacteria (via Siddha GEO) | Not specified (broad-spectrum) | Membrane disruption | Part of Siddha GEO formulation | Impairs membrane function (limits hydrophobic diffusion, dissipates ion gradients) leading to cell death | [32] |
| Bacteria (via R. beesianus EO) | Gram-positive (S. aureus, E. faecalis, B. subtilis); Gram-negative (E. coli, P. aeruginosa, P. vulgaris) | Broad-spectrum antibacterial | 15.0% component of R. beesianus EO; MIC 3.13–6.25 mg/mL | Demonstrates broad-spectrum activity against Gram-positive and Gram-negative bacteria | [33] |
| Multi-drug resistant bacteria | Not specified | Antibacterial capability | Not specified | Effective via membrane disruption mechanism | [33] |
As a constituent of EOs like Siddha GEO and R. beesianus EO, borneol works alone or alongside other monoterpenes to target microbial membranes: in Siddha GEO, it limits hydrophobic compound diffusion through the lipopolysaccharide layer and dissipates ion gradients [32], while its 15.0% presence in R. beesianus EO contributes to the oil’s broad-spectrum activity against both Gram-positive and Gram-negative bacteria [33]. This membrane disruption mechanism is central to borneol’s role in enhancing the antimicrobial properties of plant EOs [163, 164].
Borneol’s Synergistic and Seasonal/Extraction-Dependent Antimicrobial Contributions
Borneol contributes to the antimicrobial activity of plant essential oils (EOs) both as a joint component and through synergistic interactions with other terpenes, with its content and associated efficacy modulated by factors including extraction method, harvest season, and cultivation conditions. Its role in enhancing antimicrobial effects is further supported by its identification as a major active compound in EOs containing terpene alcohols (e.g., linalool, terpineol), where authentic standards confirmed its contribution to inhibitory effects[85]. Notably, while individual borneol testing via broth dilution did not yield antimicrobial results[86], its presence in complex EO mixtures enhances overall activity.
Borneol Content and Associated Antimicrobial Activity Across Plant EOs and Modulating Factors
| Plant Species | Modulating Factor | Borneol Content | Antimicrobial Activity Outcome | Citation |
|---|---|---|---|---|
| Rosemary | Extraction method (SC-CO₂ vs HD vs OM) | SC-CO₂: 18.79%; HD: 8.52%; OM: 3.75% | SC-CO₂ oil had 1.5–2.5-fold larger inhibition zones against bacteria (E. coli, S. typhimurium, B. subtilis, S. aureus) and yeasts (C. albicans, C. tropicalis) vs HD oil | [87] |
| Teucrium polium (TP) | Harvest season (winter vs autumn) | Winter: 8.24%; Autumn: absent | Winter EO showed greater antibacterial activity (MIC: 5 µl/ml) vs autumn EO (MIC: 10 µl/ml) against B. subtilis, S. aureus, P. aeruginosa | [66] |
| Lavender | Sample year (LA 2020 vs LA 2019) | LA 2020: 19.35%; LA 2019: 15.67% | LA 2020 EO had stronger antibacterial (E. coli DH5α, B. subtilis PY79) and antifungal (S. rolfsii) activity, with concentration-dependent bactericidal effects and greater mycelial growth reduction | [31] |
| Sage | Seasonal period (vegetative: May–June) | Peak: 18.38% | High borneol content (alongside α-thujone, camphor) linked to antibacterial activity | [39] |
| Lavender cultivars | In vitro propagation + AuNP supplementation (50 mg·dm⁻³) | Control: 13.38–32.17%; With AuNPs: 16.46% | (Activity outcome not specified in original text) | [34] |
These findings collectively highlight that higher borneol content—whether achieved via optimized extraction, targeted harvest timing, or cultivation adjustments—correlates with improved antimicrobial performance of EOs against a range of bacterial and fungal strains.
Borneol's Mechanisms of Antimicrobial Action
Borneol exerts broad-spectrum antimicrobial effects through interconnected mechanisms targeting bacterial cell membrane integrity, specific molecular interactions, antiadhesion/antibiofilm properties, and synergistic enhancement of conventional agents [6, 164]. A primary mechanism involves disrupting bacterial cell membranes, which increases permeability, ion leakage, and loss of essential cellular components—impairing osmotic balance and enzyme function. Borneol also interacts with key molecular targets and modulates bacterial adhesion and biofilm formation, with synergistic effects that further enhance its efficacy.
Borneol-Induced Bacterial Membrane Damage and Associated Effects
| Borneol Formulation | Bacterial Strain(s) | Observed Effect | Citation |
|---|---|---|---|
| Borneol | Listeria monocytogenes, Pseudomonas aeruginosa | Enhanced membrane porosity | [88] |
| Borneol-containing essential oils (BEO) | Staphylococcus aureus, Bacillus subtilis | Increased electric conductivity via leakage of Na⁺, K⁺, and Ca²⁺ ions | [89] |
| 2× MIC BEO | S. aureus | 15-fold higher protein leakage compared to controls | [89] |
| BEO | S. aureus, B. subtilis | Leakage of nucleic acids (measured by A₂₆₀ absorbance) and water-soluble proteins | [89] |
| Borneol | S. aureus, B. subtilis | Deformed, shriveled cells with broken cell walls/blurred outlines (via SEM/TEM), indicating irreversible cytoplasmic membrane damage | [89] |
Beyond membrane disruption, borneol targets specific molecular sites: computational studies identify human carbonic anhydrases I, II, IV (36% binding probability) and transient receptor potential cation channel subfamily M member 8 (31% binding probability) as high-affinity targets [72]. In bacteria, it forms hydrogen bonds with glycine (47) and alanine (51) residues of tyrosyl-tRNA synthetase (TyrRS) from Escherichia coli, S. aureus, and P. aeruginosa—potentially inhibiting the aminoacylation process critical for protein synthesis [90]; this hydrogen bonding capacity also contributes to activity against Gram-positive bacteria [91]. Additionally, borneol exhibits antiadhesion and antibiofilm properties, reducing bacterial attachment and biofilm formation [92]. Synergistically, it enhances the efficacy of conventional antibiotics and other compounds (e.g., terpinen-4-ol, limonene) by penetrating membranes and impairing efflux pump activity, improving co-administered agents’ antibacterial effects [92]. Collectively, these mechanisms underpin borneol’s activity against Gram-positive, Gram-negative, and multi-drug resistant bacteria [33].
Borneol's Synergistic Interactions with Other Compounds and Formulations
Borneol exhibits weak intrinsic antimicrobial activity, with minimum inhibitory concentration (MIC) values ≥4 mg mL⁻¹ categorizing it as a weakly active compound against tested microbes [93]. However, its synergistic interactions with other antimicrobial agents enhance overall antibacterial efficacy through distinct mechanistic contributions: borneol can penetrate bacterial membranes and impair efflux pump activity, facilitating the entry of co-administered compounds with stronger antimicrobial potential [92]. Synergistic effects extend to combinations with cyclic monoterpenes, monoterpene phenols, essential oils (EOs), conventional antibiotics, and formulated compounds, though rare antagonism has been reported.
Borneol Synergistic Combinations and Their Antimicrobial Effects
| Combination Type | Partner Compound/EO | Key Details | Target Microorganism(s) | Citation(s) |
|---|---|---|---|---|
| Cyclic monoterpene | γ-terpinene/α-terpinene | Synergistic at multiple ratios; higher proportions of γ/α-terpinene optimal | Candida albicans, Candida tropicalis | [93] |
| Monoterpene phenol (EO component) | Carvacrol, thymol (Thymus satureioides EO) | Borneol contributes to greater bacterial growth inhibition via synergism | Bacteria (unspecified) | [94] |
| EO blend | Thymus vulgaris EO + Myrtus communis EO (27.15% borneol) | Synergism observed | Salmonella typhimurium (FICI=0.498) | [53] |
| Conventional antibiotic + EO | Chloramphenicol + Daucus graveolens EO (18.7% borneol) | 60.0% synergistic interactions; strongest effect on Proteus mirabilis ATCC 12453 | Various bacteria (including Proteus mirabilis) | [52] |
| Conventional antibiotic + EO | (Implied: antibiotic context) + Lavandula angustifolia leafy stalk EO (13.0–19.7% borneol) | Synergism; FICI as low as 0.076 for 'Blue River' cultivar | Staphylococcus aureus (including MRSA) | [95] |
| Formulated polymer | Borneol-based polymer + desorbed camphor | Synergism enhances antifouling performance via hydrolysis-released borneol | (Antifouling context; microbes unspecified) | [96] |
| Formulated nanoparticle | ZnO nanoparticles + rosemary EO (contains borneol) | Increased inhibition zones vs. standalone rosemary EO; EO sensitizes cells to ZnO | Staphylococcus aureus | [97] |
| Rare antagonism | Thymol + borneol | Antagonistic interaction reported | Moraxella cattarhalis | [93] |
For instance, combinations of γ-terpinene or α-terpinene (cyclic monoterpenes that disrupt membrane lipid bilayers) with borneol showed synergistic effects against fungal pathogens [93], while borneol in Thymus satureioides EO synergizes with carvacrol and thymol to boost bacterial inhibition [94]. Formulations like ZnO nanoparticles loaded with rosemary EO (containing borneol) demonstrate enhanced efficacy against S. aureus by sensitizing bacterial cells to the nanoparticles [97]. Notably, the combination of thymol and borneol is an exception, showing antagonism against Moraxella cattarhalis [93].
Borneol Extraction and Quantification Methods in Plant Materials
Borneol extraction and quantification in plant materials depend on species-specific standardized methods and analytical techniques. Key details of these protocols, including target species, extraction approaches, analytical instruments, and calibration parameters, are summarized below.
Borneol Extraction and Analytical Protocols for Selected Plant Species
| Target Species | Extraction Method/Standardization | Analytical Technique & Key Parameters | Calibration/Quantification Details | Citation |
|---|---|---|---|---|
| Alpinia villosum | Volatile oil determination per Chinese Pharmacopoeia 2020 (General Rule 2204) | GC-MS (Agilent 7890B-5977B) with borneol, bornyl acetate, camphor as standards | N/A (standards used for quantification) | [98] |
| Artemisia argyi (AAEO) | N/A | GC (Agilent 6890N; HP-INNOWax column: 30 m × 0.25 mm × 0.5 μm); oven program: 110°C (8 min) → 158°C (25°C/min, 8 min) → 230°C (25°C/min, 9 min); injector 250°C, detector 280°C | Internal standard: cyclohexanone; borneol standard curve: Y = 4.2625X + 0.0028, R² = 1.0000 | [99] |
| Traditional Chinese Medicines (TCMs) | N/A | N/A | Calibration curve method; methanol-dissolved standards (5.0 mg mL⁻¹ stock); borneol curve: Y = 7.86X + 0.53, R² = 0.9997 (Y = peak area ratio to internal standard, X = concentration) | [100] |
| Cinnamomum burmannii | Soxhlet extraction; solid-liquid ratio (fresh leaf: anhydrous ethanol): 1:60 (g/mL); 80°C, 6 h; essential oil diluted to 100 mL | 1. GC (Agilent 7890A; Agilent 19091N-113 column: 30m × 320 μm × 0.25 μm); carrier gas: N₂ (2 mL/min); oven program: 70°C (1 min) → 100°C (3°C/min) → 250°C (15°C/min, 1 min); injection: 1.0 µL splitless, port 220°C, detection 230°C 2. GC-MS (Shimadzu QP2010 PLUS; SH-RXI-5SILMS column: 30 m × 0.25 mm × 0.25 μm); same oven program; port 280°C, EI ion source 200°C, connection line 250°C, scan range m/z 29–500 |
N/A | [101] |
| Cinnamomum camphora chvar. Borneol (fresh leaves) | Neutral cellulase-assisted steam distillation (NCSD) vs. steam distillation (SD) | N/A | Borneol content: NCSD (11.66 ± 0.22%) vs. SD (10.69 ± 0.16%) | [102] |
Borneol content varies across plant species, organs, and growth conditions. For example, in A. villosum varieties A11 and A12, borneol levels are similar [98], while in rosemary, supercritical CO₂ (SC-CO₂) extraction yields the highest borneol content (18.79%), followed by hydrodistillation (HD, 8.52%) and organic solvent extraction (OM, 3.75%) [87]. Roots are the richest vegetative organ for borneol in certain plants [103]. Artemisia gypsicola shows peak borneol accumulation at the post-flowering stage-8 pm (22.62%), with high levels also at pre-flowering stage-6 am (18.22%) and full flowering stage-6 am (14.98%) [23]. Among Satureja cuneifolia oil samples, borneol contents are 24% (Sc-A), 19% (Sc-B), and 12.9% (Sc-C) [14]. Other species have varying borneol levels: A. grandifolia oil (5.2%), Tetradenia macrophyllum oil (9.1%) [104], Ocimum basilicum essential oils (OEOs, 0.26–1.35%) [105], Artemisia herba-alba essential oil (3.3%) [38], sage essential oil samples (2.12–18.38%) [39], Satureja montana essential oil (5.0%), and its August-harvested hydrosols (20.4%, four times the essential oil level) [106]. Thymus species oil contains 6.1% borneol [107], while a specimen from the Cobar Peneplain bioregion has borneol as the dominant component (31.7%) [108]. C. burmannii chemotypes differ in D-borneol content: high chemotypes have ~seven times more D-borneol than low chemotypes, accounting for 66.21% vs. 55.99% of the essential oil [101]. Sage essential oils show peak borneol content during the vegetative period (May-June) [39].
Biosynthetic pathways regulate borneol content across species. In A. argyi, (+)-borneol is produced from geranyl diphosphate (GPP) by bornyl diphosphate synthase (AarTPS89), which is then converted to (+)-camphor by bornyl dehydrogenases (AarBDH4 and AarBDH5) [109]. A. villosum has eight genes for bornyl acetate synthesis and two for camphor synthesis; Wv_032842 (upregulated in A12) may drive differential bornyl acetate content despite similar borneol levels between varieties [98]. In C. burmannii, 76 candidate isoforms encode terpenoid biosynthesis enzymes, including 4 for monoterpenoid (Ko00902) and 8 for diterpenoid (Ko00904) pathways; KEGG annotation links D-borneol biosynthesis to potential plant-pathogen interaction pathways [101]. C. camphora chvar. Borneol produces borneol via pinene bioconversion, and neutral cellulase pre-treatment in NCSD enhances release by hydrolyzing cell walls to access intracellular borneol [102]. In OEOs, linalool and borneol show a strong positive correlation (R = 0.98) [105].
Environmental and temporal factors influence borneol accumulation. Copper sulphate treatment increases borneol content in some plant oils [64]. Root-stem samples of one species show peak borneol synthesis at 35.85% (pre-flowering stage) and 44.39% (6 am) [110], with significant positive linear correlations between borneol and 1,8-cineole (r = 0.409**), camphor (r = 0.364**), terpinen-4-ol (r = 0.380**), and verbenone (r = 0.276**) at p ≤ 0.01 [110]. Mavandi et al. found that COM and VC increased lavender borneol content compared to controls [111]. Seasonal changes slightly alter borneol enantiomer composition, with (S)-borneol as the major enantiomer in some cases [106].
Borneol's Antimicrobial Efficacy Against Resistant Bacterial Strains
Borneol is a constituent of essential oils (EOs) and extracts with demonstrated activity against clinically and foodborne relevant resistant bacterial strains, though its efficacy may depend on synergistic interactions with other components in complex mixtures. In silico molecular docking studies have linked the antibacterial activity of borneol-containing EOs to targeting enzymes involved in microbial metabolism, cell wall biosynthesis, and nucleic acid repair; notably, sesquiterpenes (e.g., β-caryophyllene, α-humulene) in these EOs showed higher binding affinity to the key bacterial enzyme isoleucyl-tRNA synthetase than monoterpenes [112]. However, pure borneol alone exhibited limited activity, with MIC values above 128 µg/mL (0.557–0.842 mM) against tested strains [113].
Borneol-Containing Mixtures and Their Activity Against Resistant Bacterial Strains
| Source of Borneol-Containing Mixture | Borneol Context | Target Resistant Bacterial Strains | Key Activity Metrics | Citation |
|---|---|---|---|---|
| Thymus maroccanus EOs | Up to 16.3% borneol content | Antibiotic-resistant Klebsiella pneumoniae (resistant to cefixime and gentamycin) | Inhibition zones: 21.3–23.7 mm | [12] |
| Lavandin EOs (cv. ‘Alba’ and ‘Sumiens’) | High borneol content (notable in cv. ‘Sumiens’) | Salmonella strains (70% resistant to chloramphenicol) | cv. ‘Sumiens’ showed notable activity; MIC values contributed to inhibition | [114] |
| Volatile-enriched fraction | One of the most abundant monoterpenes | Helicobacter pylori strains | 6.3–7.1 log CFU reduction | [115] |
| Isolated borneol | Pure compound | Multidrug-resistant Gram-negative pathogens (K. pneumoniae, Escherichia coli, Acinetobacter baumannii, Enterobacter cloacae) | Potent activity: MIC of 0.47 mg/L against K. pneumoniae; more active than gentamicin against other strains | [116] |
| Combretum spp. leaf extracts | Present in relative abundance | Methicillin-resistant Staphylococcus aureus (MRSA); extended-spectrum β-lactamase (ESBL)-producing E. coli and K. pneumoniae | Inhibitory activity against all target strains | [117] |
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