지속가능한 다기능성 화장품 에몰리언트로서 쌍별귀뚜라미(Gryllus bimaculatus) 오일의 지방산 조성 및 항산화 특성 연구
Fatty Acid Profiling and Antioxidant Potential of Two-Spotted Cricket (Gryllus bimaculatus) Oil as a Sustainable Multifunctional Cosmetic Emollient
作为可持续多功能化妆品润肤剂的双斑蟋 (Gryllus bimaculatus)油脂肪酸组成 及抗氧化特性研究
Article information
Abstract
목적
식용 곤충은 우수한 생리활성을 지닌 지속가능한 바이오 자원으로 주목받고 있다. 본 연구는 자원 업사이클링 및 친환경적 화장품 소재 발굴을 위해 쌍별귀뚜라미(Gryllus bimaculatus) 유래 오일(GBO)의 항산화 및 항염증 활성을 분석하고, 지속 가능하며 친환경적인 화장품 에몰리언트 소재로서의 활용 가능성을 평가하고자 하였다.
방법
쌍별귀뚜라미를 절식, 세척, 건조한 후, 대기압 조건 및 120℃에서 스크류 프레스를 이용하여 오일을 압착 추출하였다. 쌍별귀뚜라미 오일의 지방산 조성은 GC/MS를 이용하여 분석하였고, 항산화 활성은 총 폴리페놀 함량(TPC), 총 플라보노이드 함량(TFC), DPPH 및 ABTS 라디칼 소거 활성을 측정하여 평가 하였다.
결과
세포생존율 및 항염증 활성은 다양한 농도의 쌍별귀뚜라미 오일을 처리한 RAW 264.7 세포에서 세포 생존율과 산화질소(NO) 생성 및 전염증성 세포전달물질 억제 능력을 통해 검증하였다. 쌍별귀뚜라미 오일은 9.7%의 수율을 보였으며, 풍부한 불포화 지방산 조성을 나타냈다. 쌍별귀뚜라미 오일의 6대 주요 지방산은 올레산(30.8%), 팔미트산(27.7%), 리놀레산(26.8%), 스테아르산(8.7%), 팔미톨레산(2.1%), 미리스틱산(1.02%)이었다. 쌍별귀뚜라미 오일은 농도 의존적인 방식으로 NO 생성과 전염증성 세포 전달물질을 유의미하게 감소시켰으며, 특히 높은 총 폴리페놀 함량과 함께 우수한 DPPH 라디칼 소거 활성을 나타내 높은 항산화력을 나타냈다.
결론
쌍별귀뚜라미 오일은 항산화 효능, 제형 안정화 효과 및 피부 생리학적 보호 기능을 제공하는 다기능성 화장품 성분으로서, 고기능성 코스메슈티컬 제재 적용을 위한 유망 바이오 소재가 될 수 있음을 시사한다.
Trans Abstract
Purpose
Recent research has recognized edible insects as sustainable, high-performance bioresources for advanced functional cosmetic applications. The present study aimed to analyze the antioxidant and anti-inflammatory activities of two-spotted cricket (Gryllus bimaculatus) oil (GBO) to support resource-circulating economic practices and evaluate its potential as a sustainable, eco-friendly cosmetic emollient.
Methods
The crickets were starved, washed, dried, and then their oil was extracted with a screw press at 120°C under atmospheric pressure. The fatty acid composition of GBO was analyzed using gas chromatography/mass spectrometry. The antioxidant activity was evaluated by measuring total polyphenol content (TPC), total flavonoid content (TFC), and DPPH and ABTS radical scavenging activities. Anti-inflammatory activity was assessed by examining the inhibitory effect on NO production and proinflammatory cytokine levels in RAW 264.7 cells at various concentrations of GBO.
Results
The extracted GBO showed a yield of 9.7% and a rich profile of unsaturated fatty acids. The six major fatty acids identified in GBO were oleic (30.8%), palmitic (27.7%), linoleic (26.8%), stearic (8.7%), palmitoleic (2.1%), and myristic (1.02%) acids. GBO attenuated NO production and proinflammatory cytokine levels in a concentration-dependent manner and exhibited high TPCs and robust DPPH scavenging.
Conclusion
As a multifunctional cosmetic ingredient that provides antioxidant efficacy, formulation-stabilizing benefits, and physiological protection, GBO represents a promising bioresource for advanced cosmeceutical applications.
Trans Abstract
目的
食用昆虫作为具有优异生物活性的可持续生物资源而备受关注。本研究旨在通过分析双斑蟋(Gryllus bimaculatus)衍生油(GBO)的抗氧化及抗炎活性，以实现资源升级再利用并开发环保型化妆品原料，从而评估 其作为可持续且环保的化妆品润肤剂原料的应用前景。
方法
将双斑蟋进行禁食、清洗和干燥处理后，在大气压 条件及120°C下，利用螺旋压榨机压榨提取得到油脂。利用GC/MS分析双斑蟋油的脂肪酸组成；通过测定总多酚 含量(TPC)、总黄酮含量(TFC)以及DPPH和ABTS自由基清除活性来评估其抗氧化活性。
结果
细胞存活率和抗炎 活性是通过在处理了不同浓度双斑蟋油的RAW 264.7细胞中，检测细胞存活率、一氧化氮(NO)生成量以及促炎 介质抑制能力来进行验证的。双斑蟋油的得率为9.7%，并表现出丰富的不饱和脂肪酸组成。双斑蟋油的六大主 要脂肪酸分别为：油酸(30.8%)、棕榈酸(27.7%)、亚油酸(26.8%)、硬脂酸(8.7%)、棕榈油酸(2.1%)和肉豆蔻酸 (1.02%)。双斑蟋油以浓度依赖性方式显著减少了NO的生成和促炎介质的释放，特别是其具有较高的总多酚含 量以及优异的DPPH自由基清除活性，展现出极高的抗氧化能力。
结论
双斑蟋油作为一种兼具抗氧化功效、配 方稳定效果和皮肤生理保护功能的多功能化妆品成分，有望成为应用于高功能性药妆制剂的极具前景的生物原料。
Introduction
Edible insects have attracted attention as sustainable alternatives for future food and bioresources in response to global population growth and resource depletion (Gebreyes & Teka, 2025). The two-spotted cricket (Gryllus bimaculatus) is one of the most abundant species of its kind; it is widely distributed across the tropical and subtropical regions of Asia, Africa, and southern Europe (Park et al., 2021). Due to its rich nutritional value, high reproductive rate, and eco-friendly rearing characteristics, this insect has been widely recognized and approved as a safe food ingredient by regulatory bodies, including the Ministry of Food and Drug Safety, South Korea (Lee et al., 2016; Kwon & Lee, 2026). With steady growth in the edible insect market, the potential of insect-derived components for high-value bioactive applications, extending beyond traditional entomophagy, has been recently investigated (Sinha et al., 2024; Guerrero et al., 2026; Liu et al., 2025; Martin et al., 2026).
Cutaneous disorders, including skin barrier disruption and accelerated aging, driven by climate change, environmental pollution, and the resulting oxidative stress, have risen globally (Haykal et al., 2025). In response to these issues, the demand for high-performance cosmeceuticals is rising, requiring the cosmetics industry to urgently identify and develop sustainable, natural alternatives to conventional ingredients. In this regard, lipids extracted from nutritionally validated edible two-spotted crickets are promising multifunctional emollient candidates. Human stratum corneum lipids consist of approximately 50% ceramides, 25% cholesterol, and 10-15% free fatty acids to maintain barrier integrity (Ananthapadmanabhan et al., 2013; Ahlström et al., 2024). GBO offers exceptional biomimetic compatibility due to its high content of palmitic, oleic, and linoleic acids, the predominant fatty acid species in human skin (Siqueira et al., 2025). This quantitative and qualitative alignment may help GBO effectively restore disrupted cutaneous lipid structures.
Oxidative stress and chronic inflammatory cascades, major drivers of cutaneous damage and accelerated aging, are induced by exogenous stimuli and trigger a deterioration in skin barrier function. As soon as the skin contacts these irritants, the cutaneous immune system overproduces nitric oxide (NO) and proinflammatory cytokines, disrupting the epidermal matrix and localized inflammation (Man et al., 2022). Therefore, an ideal multifunctional cosmetic agent must transcend from being a mere physical occlusive barrier for surface moisture retention and possess biological activities, including scavenging intracellular reactive oxygen species and modulating inflammatory mediators.
The stratum corneum (SC), a core component of the epidermal permeability barrier, comprises ceramides, cholesterol, and free fatty acids organized as a sophisticated lamellar liquid-crystal structure (Cui et al., 2010; Georgiev et al., 2026). Thus, a balanced ratio of saturated to unsaturated fatty acids is essential for restoring the barrier matrix. The abundant UFAs in GBO, along with their optimal balance with SFAs, may provide superior physicochemical configurations that reinforce the cutaneous lipid barrier and mitigate transepidermal water loss (TEWL). Furthermore, total bioactive compounds (e.g., TPC and TFC) inherent to these insect-derived lipids can protect against oxidative stress and subsequent skin barrier disruption (Romeo et al., 2020; Nip et al., 2024; Wang et al., 2024a).
Despite such biomimetic and therapeutic potential, research on two-spotted cricket oil (GBO) as a high-performance topical formulation or cosmeceutical ingredient remains limited. To validate the feasibility of GBO as a sustainable, multifunctional cosmeceutical, this study aimed to analyze its physicochemical properties, fatty acid profiles, bioactive composition, antioxidant capacities, and anti-inflammatory activities.
Materials and Methods
1. Oil Extraction from Two-spotted Crickets
Two-spotted crickets reared for 45 days were purchased from Baekmandori Agricultural Corporation (Korea) and fasted for 24 h (Ahn et al., 2015). The crickets were thoroughly washed with running tap water, rinsed with sterile distilled water, and dried in a convection oven at 60℃ for 24 h. For oil extraction, 3 kg of dried crickets were processed using a hot-press machine (Dongbang Electronic, Korea) at 120℃ at atmospheric pressure for 30 min. After centrifugation at 3000 RPM for 20 min at room temperature, the supernatants were collected in amber bottles. GBO was stored at 4℃ until further analysis.
2. Fatty Acid Composition
The total fatty acid profile of GBO was determined using a 7890B gas chromatography (GC) system equipped with a 5977B mass spectrometry (MS) detector (Agilent Technologies, USA). Samples were analyzed by Korea Bio Analysis Lab (Korea).
3. Determination of Total Phenolic and Flavonoid Contents
To evaluate bioactive components and antioxidant activities, GBO was dissolved in ethanol (EtOH; Merck, Germany). The control group received an identical final concentration of EtOH as a vehicle control to rule out any potential effects of the solvent on the experimental results. Total phenolic contents (TPC) of GBO were analyzed based on the method of Singleton & Rossi (1965). Briefly, 350 μL of GBO was mixed with 70 μL of 50% Folin-Ciocalteu reagent (Sigma-Aldrich, USA) for 3 min. After adding 350 μL of 2% Na2CO3 solution (Sigma-Aldrich) and incubation for 1 h, the OD750 was recorded with a VarioskanTM LUX microplate spectrophotometer (Thermo Fisher Scientific, Finland). TPC was quantified based on a gallic acid standard curve. Total flavonoid contents (TFC) were quantified according to Davis (1947) with minor modifications. Briefly, 70 μL of GBO was reacted with 700 μL of diethylene glycol and 7 μL of 1 N NaOH at 37℃ for 1 h. Absorbance was measured at 420 nm using a microplate spectrophotometer (Thermo Fisher Scientific, Finland), and TFC was calculated using a quercetin standard curve.
4. Determination of ABTS Radical Scavenging Activity
In our study, 2,2′-azinobis(3-ethylbenzothiazoline-6-sulfonic acid (ABTS) radical scavenging activity was assayed according to Fellegrini et al. (1999). Briefly, 7.4 mM ABTS (Sigma-Aldrich) and 2.6 mM potassium persulfate (Daejung Chemicals, Korea) were mixed in an equal volume and then incubated in the dark at room temperature for 24 h to generate ABTS+ radicals. Then, 10 μL of GBO was reacted with 190 μL of ABTS reagent for 10 min, with absorbance recorded at 732 nm using a microplate spectrophotometer (Thermo Fisher Scientific, Finland). ABTS radical scavenging activity was quantified using gallic acid as the standard.
5. Determination of DPPH Radical Scavenging Activity
The 1,1-diphenyl-2-picrylhydrazyl (DPPH) radical scavenging activity of GBO was determined using the method of Blois (1958) with minor modifications. Briefly, 100 μL of GBO was mixed with 0.15 mM DPPH solution (Sigma-Aldrich). After 30 min of incubation in the dark, absorbance was measured at 517 nm using a microplate spectrophotometer (Thermo Fisher Scientific, Finland). DPPH radical scavenging activity was quantified using gallic acid as the standard.
6. RAW 264.7 Cell Culture
RAW 264.7 mouse macrophage cell lines, obtained from the Korean Cell Line Bank (Seoul, Korea), were seeded into a 96-well plate at a density of 4.0×104 cells/well. After culture, the cells were supplemented with 10% fetal bovine serum (GibcoTM, USA) and 1% penicillin-streptomycin. The cells, upon reaching 80% confluence, were treated with 1.5625%-25% (v/v) GBO for all tests except cell viability determination. The cells were re-incubated at 37℃ for 24 h in a humidified atmosphere containing 5% CO2 in HeracellTM Vios 250i incubator (Thermo Fisher Scientific, Germany).
7. Determination of Cell Viability
Cell viability was assessed using the (3-(4,5-dimethylthiazol-2-yl)-5-(3-carboxymethoxyphenyl)-2-(4-sulfophenyl)-2H-tetrazolium) (MTS) assay. Briefly, 20 μL of MTS reagent was added to 100 μL of cells treated with 1.5625%-50% (v/v), followed by re-incubation at 37℃ for 1 h. Absorbance was recorded at 490 nm using a microplate spectrophotometer (Thermo Fisher Scientific, Finland).
8. Determination of Nitric Oxide Production
Anti-inflammatory activity was evaluated by measuring nitric oxide (NO) production in lipopolysaccharide-stimulated RAW 264.7 cells. When the cells reached 80% confluence, they were pretreated with 1.5625%-25% (v/v) GBO for 1 h and then stimulated with 100 ng/mL LPS for 24 h. The concentration of nitrite was determined by reacting Griess reagent (Intron Biotechnology, Korea) with 100 μL of the culture supernatant from each well. Absorbance was recorded at 540 nm using a microplate spectrophotometer (Thermo Fisher Scientific, Finland). NO production was quantified using a standard curve generated by sodium nitrite.
9. Measurement of Level of Proinflammatory Cytokines
To determine the levels of proinflammatory cytokines, 100 μL of RAW 264.7 cell culture supernatant was collected. The concentrations of TNF-α, IL-1β, and IL-6 in this supernatant were measured using commercial ELISA kits (Abcam, Cambridge, UK) according to the manufacturer’s instructions. The absorbance of each cytokine was recorded using a microplate spectrophotometer.
10. Statistical Analysis
All experiments were performed in triplicate. Statistical analyses were performed using SPSS S/W version 29.0 (IBM, USA). The TFC, TPC, ABTS, DPPH, and Nitrite production were compared using one-way analysis of variance. Tukey’s HSD post hoc tests were performed to test the equality of variances. A p≤0.05 was considered statistically significant.
Results and Discussion
1. Physicochemical Properties of GBO
GBO yield via the hot-press method was 9.7%. The density of GBO was 1.088 g/mL at room temperature. As a result of the color analysis, GBO exhibited a distinct yellowish hue, characterized by values of L* =79.57±0.01, a* =10.13±0.01, and a notably high b* =103.50±0.02 (Figure 1). The latter might imply a rich retention of lipophilic bioactive compounds (e.g., carotenoids, flavonoids, and tocopherols), which could potentially exhibit a synergistic antioxidant network that defends against oxidative stress and downregulates inflammatory mediators in the skin (Feng et al., 2022). This vivid natural coloration is advantageous for premium oils or nutrient-dense creams, where the formulation opacity can naturally accommodate the intense hue without compromising consumer preference. However, it may limit the dosage or require an additional decolorization process, such as activated carbon filtration or clay adsorption, to remove pigmented lipophilic components while using transparent toners or white emulsions. In contrast, as the L* value ranging from black (0) to white (+100) correlates with the level of skin pigmentation, it is the most sensitive of all trichromatic values to skin color changes (Huang et al., 2018). The relatively high L* value of GBO suggests that it could be easily incorporated into various formulations without remarkably compromising the aesthetics of the final product.
Schematic representation of the 3D CIELAB color space for GBO characterization, illustrating the L* axis (black to white lightness) alongside the chromatic a* (green to red) and b* (blue to yellow) axes.
2. Fatty Acid Profile of GBO
GC/MS analysis identified the total fatty acid profile of GBO, consisting of 39.41% SFAs and 60.59% UFAs (Table 1). The major fatty acids in GBO were oleic acid (C18:1, 30.81%) and linoleic acid (C18:2, 26.76%) among UFAs, and palmitic acid (C16:0, 27.71%) and stearic acid (C18:0, 8.69%) among SFAs. In addition to the major fatty acids, small amounts of palmitoleic (C16:1, 2.12%) and myristic (C14:0, 1.02%) acids were also present. In addition, trace levels of diverse fatty acids were also detected. Long-chain SFAs can serve as metabolic substrates for ceramide synthesis, thereby supporting the restoration of the skin barrier (Nip et al., 2024). Furthermore, these fatty acids are essential for synthesizing stable lamellar liquid-crystal structures within emulsions, which effectively mimic the multilamellar lipid organization of the human skin barrier (Cui et al., 2010; Georgiev et al., 2026). These structures not only improve the thermodynamic stability of the product but also reinforce the skin’s occlusive barrier, mitigating TEWL (Nip et al., 2024; Vitek et al., 2024). The predominance of UFAs, particularly oleic and linoleic acids, facilitates physiological skin benefits. Oleic acid has been recognized for its ability to regulate epidermal lipid metabolism, facilitate barrier recovery, and preserve SC hydration (Mank et al., 2016). Linoleic acid, the most abundant PUFA in the human epidermis, is an essential substrate for ceramide synthesis and maintaining the epidermal permeability barrier, offering potent immunomodulatory effects that suppress inflammatory responses (Wang et al., 2024a; Jiang et al., 2026). Recent studies have demonstrated its crucial role in the cosmeceutical field by demonstrating its efficacy in regulating acne and atopic dermatitis, alleviating psoriasis, and promoting wound healing (Li et al., 2020; Zhao et al., 2022; Wang et al., 2024b; Cheng et al., 2025). In spite of being in trace amounts, myristic acid could enhance the formulation’s tactile properties by improving spreading ability (Yang et al., 2020; Jitrangsri et al., 2026). Palmitoleic acid, a rare fatty acid in vegetable oils but common in insect lipids, correlates with accelerating skin regeneration and antimicrobial protection (Yang et al., 2020; Guerrero et al., 2026). Such balanced fatty acid profiles may position GBO as a high-performance cosmeceutical ingredient and a potential multifunctional emollient.
Fatty acid composition of the two-spotted cricket oil (GBO)
3. Antioxidant Properties of GBO
The antioxidant properties of GBO were evaluated by quantifying bioactive compounds, such as TPC and TFC, and by evaluating ABTS and DPPH radical scavenging activities. TPC and TFC of GBO were quantified to assess its bioactive potential. The TPC was 2.34±0.14 mg GAE/g, and the TFC was 0.63±0.06 mg QE/g. GBO exhibited ABTS and DPPH radical scavenging activities of 0.49±0.25 mg GAE/g and 3.58±0.07 mg GAE/g, respectively (Table 2). Although GBO was predominantly composed of UFAs susceptible to oxidative degradation, the rich TPC may act as endogenous stabilizers to preserve integrity (Romeo et al., 2020). The higher DPPH assay activity than the ABTS assay indicates the lipophilic nature of the GBO oil matrix, which may enhance the solubility and reactivity of hydrophobic antioxidants within the oil (Liu et al., 2025). The antioxidant properties of GBO could protect UFAs from lipid peroxidation and protect skin cells against oxidative damage.
Antioxidant properties of the two-spotted cricket oil (GBO).
4. Cell Viability and Cytotoxicity of GBO
Cell viability was evaluated using the MTS assay in RAW 264.7; no significant cytotoxicity was observed at 1.5625%-25% (v/v) GBO. Cell viability was maintained 93.0% at 25% (v/v) GBO, but decreased to 40.1% at 50% (v/v) GBO (LC50=43.73%± 1.82% [v/v]) (Figure 2). The low cytotoxicity may be attributed to the high TPC and DPPH scavenging radical scavenging activity (Masisi et al., 2021) as well as the biomimetic nature of the fatty acid composition in GBO, such as palmitic and oleic acids, which are naturally occurring components of the human skin barrier (Cui et al., 2010; Nip et al., 2024; Georgiev et al., 2026).
Cell viability evaluated by MTS assay using RAW264.7 cells treated with diverse concentrations of the two-spotted cricket oil (GBO).
*Values marked with the same letter are not significantly different, whereas those with different ones are significantly different according to ANOVA and Turkey’s HSD at p<0.05.
5. Anti-inflammatory Effects of GBO
To evaluate anti-inflammatory activity, NO production and proinflammatory cytokine levels were measured in LPS-stimulated RAW 264.7 cells. GBO treatment induced a concentration-dependent reduction of NO levels, decreasing from 73.37±0.40 to 12.27±2.86 μM at the highest concentration tested (EC50=7.30%±1.93%) (Figure 3A). A dose-dependent decline in NO production suggests that GBO suppresses the expression of key inflammatory response enzymes, such as inducible nitric oxide synthase (Cinelli et al., 2020). In addition, compared with the LPS-treated group, TNF-α, IL-1β, and IL-6 levels decreased markedly in a concentration-dependent manner at 1.5625% to 25% (v/v) GBO (Figure 3B, 3C, 3D). While IL-1β and IL-6 levels were suppressed significantly, TNF-α contents showed a relatively moderate decrease as the GBO concentration increased, indicating more specific action on downstream cytokine-induction pathways (e.g., JAK/STAT or NF-κB) rather than early-stage TNF-α (Alciato et al., 2010; Ahmad et al., 2015). This differential inhibitory effect is beneficial for the cosmeceutical application of GBO as it enables the alleviation of chronic inflammation without compromising the fundamental immune roles of TNF-α. Therefore, further studies utilizing protein expression analyses, such as Western blotting for JAK/STAT or NF-κB signaling, are required to identify these molecular mechanisms definitively. Such investigations will clarify the mechanism by which GBO selectively modulates downstream cytokine pathways.
Anti-inflammatory effects of two-spotted cricket oil (GBO) on RAW264.7 cells.
(A) Inhibition of nitric oxide (NO) production; (B–D) Inhibition of proinflammatory cytokine production, including TNF-α (B), IL-1β (C), and IL-6 (D).*Values marked with the same letter are not significantly different, whereas those with different ones are significantly different according to ANOVA and Turkey’s HSD at p<0.05.
Conclusion
The present study characterized the fatty acid composition of GBO, evaluated its antioxidant and anti-inflammatory activities, and determined its potential for cosmeceutical application as a multifunctional ingredient. SFAs rich in GBO, such as palmitic, stearic, and myristic acids, provided the structural stability necessary for physical barrier function. As long-chain SFAs, these fatty acids may promote the formation or stabilization of lamellar liquid-crystal structures that mimic the natural lipid barrier of human skin. Furthermore, UFAs, including oleic, linoleic, and palmitoleic acids, may contribute to the functional protection of the skin through their biomimetic compatibility, inhibition of NO production, and targeted suppression of proinflammatory cytokines. The high phenolic compound content and DPPH radical scavenging activities suggest that GBO can neutralize oxidative stress and preserve lipid matrix integrity. Regarding safety, the assessment of cell viability indicates that GBO exhibits low cytotoxicity and can exert its multifaceted bioactive benefits without adverse cellular responses. Taken together, the fatty acid profile and potent antioxidant and anti-inflammatory activities confirm that GBO can function as an eco-friendly, sustainable, and multifunctional bioactive emollient that may simultaneously strengthen the skin’s physical architecture and biologically alleviate cutaneous inflammation. Further studies, including in vivo skin irritation tests and long-term stability evaluations within actual cosmetic formulations, are necessary to validate its clinical safety and practical applicability in the cosmetics industry.
Notes
Acknowledgements
This study was supported by the 2026 RDA Fellowship Program (Project No. PJ01759602) of the National Institute of Agricultural Sciences, Rural Development Administration, Republic of Korea.
Author's contribution
SWY and SYL contributed to all aspects of the experiment and analysis and wrote the manuscript. JHL, JHL, and HSL assisted in experimental design, analysis, and forging collaborations with companies, other departments, and researchers. YYJ oversaw the project and assisted in the experimental design, supported/prepared all materials used in the experiments, and supervised the manuscript writing process.
Author details
Seung-Won Yi (Postdoctoral Fellow)/So-Young Lee (Postdoctoral Fellow)/Joon Ha Lee (Researcher)/Ji Hae Lee (Researcher)/Heui-Sam Lee (Senior Researcher)/You-Young Jo (Senior Researcher), Industrial Entomology Division, Department of Agricultural Biology, National Institute of Agricultural Sciences, Nongsaengmyeongro 166, Wanjugun, Jeollabukdo 55365, Korea.