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Tawfeek, M. E., Ali, H. M., Akrami, M., and Salem, M. Z. M. (2021). "Potential insecticidal activity of four essential oils against the rice weevil, Sitophilus oryzae (L.) (Coleoptera: Curculionidae)," BioResources 16(4), 7767-7783.

Abstract

Oils extracted from Cymbopogon citratus, Lantana camara, Artemisia camphorata, and Imperata cylindrica plants were used as potential insecticides against the rice weevil, Sitophilus oryzae (L.) (Coleoptera: Curculionidae). The phytochemical composition of the isolated oils was identified by gas chromatograph-mass spectrometry (GC-MS). Oil contact toxicities were evaluated against the adults of S. oryzae. The activities of acetylcholinesterase (AChE), alkaline phosphatase (ALP), and transaminases enzymes (AST) were measured. L. camara oil (LC50 = 9.81 mg/cm2) demonstrated the highest effect, followed by C. citratus oil (LC50 = 10.89 mg/cm2), A. camphorata EO (LC50 = 16.12 mg/cm2), and I. cylindrica oil (LC50= 36.85 mg/cm2) against the adults of S. oryzae. The inhibition percentages of AChE were 38.8, 41.7, 35.0, and 27.2%; ALP were 42.4, 49.3, 28.1, and 18.7%; AST were 33.9, 38.7, 20.8, and 11.8%; and ALT were 22.7, 30.5, 14.6, and 9.6% after treated S. oryzae with oils from C. citratus, L. camara, A. camphorata and I. cylindrica, respectively. The highest abundant compounds in C. citratus were geranial (25.95%), nerylacetal (8.85%), and neral (8.45%), in L. camara were caryophyllene (12.2%), and 3-elemene (8.89%), in A. camphorata were germacrene D-4-ol (20.83%), and borneol (19.47%), and in I. cylindrica were 5-phenylundecane (10.68%), and 6-phenyldodecane (8.70%).


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Potential Insecticidal Activity of Four Essential Oils against the Rice Weevil, Sitophilus oryzae (L.) (Coleoptera: Curculionidae)

Mohamed E. Tawfeek,a Hayssam M. Ali,b Mohammad Akrami,c and Mohamed Z. M. Salem d,*

Oils extracted from Cymbopogon citratus, Lantana camara, Artemisia camphorata, and Imperata cylindrica plants were used as potential insecticides against the rice weevil, Sitophilus oryzae (L.) (Coleoptera: Curculionidae). The phytochemical composition of the isolated oils was identified by gas chromatograph-mass spectrometry (GC-MS). Oil contact toxicities were evaluated against the adults of S. oryzae. The activities of acetylcholinesterase (AChE), alkaline phosphatase (ALP), and transaminases enzymes (AST) were measured. L. camara oil (LC50 = 9.81 mg/cm2) demonstrated the highest effect, followed by C. citratus oil (LC50 = 10.89 mg/cm2), A. camphorata EO (LC50 = 16.12 mg/cm2), and I. cylindrica oil (LC50= 36.85 mg/cm2) against the adults of S. oryzae. The inhibition percentages of AChE were 38.8, 41.7, 35.0, and 27.2%; ALP were 42.4, 49.3, 28.1, and 18.7%; AST were 33.9, 38.7, 20.8, and 11.8%; and ALT were 22.7, 30.5, 14.6, and 9.6% after treated S. oryzae with oils from C. citratus, L. camara, A. camphorata and I. cylindrica, respectively. The highest abundant compounds in C. citratus were geranial (25.95%), nerylacetal (8.85%), and neral (8.45%), in L. camara were caryophyllene (12.2%), and 3-elemene (8.89%), in A. camphorata were germacrene D-4-ol (20.83%), and borneol (19.47%), and in I. cylindrica were 5-phenylundecane (10.68%), and 6-phenyldodecane (8.70%).

Keywords: Oily extracts; Chemical composition; Contact toxicity; Sitophilus oryzae; Acetylcholinesterase; Aspartate transaminase; Alanine transaminase

Contact information: a: Department of Applied Entomology and Zoology, Faculty of Agriculture, 21545 El-Shatby, Alexandria University, Alexandria, Egypt; b: Botany and Microbiology Department, College of Science, King Saud University, P.O. Box 2455, Riyadh 11451, Saudi Arabia c: Department of Engineering, University of Exeter, Exeter EX4 4QF, UK; d: Forestry and Wood Technology Department, Faculty of Agriculture (El-Shatby), Alexandria University, Alexandria 21545, Egypt;

* Corresponding Author: zidan_forest@yahoo.com

INTRODUCTION

Stored-product pests cause critical misfortunes in weight and quality of the stored grains and cereal products (Fields and Korunic 2000; Neethirajan et al. 2007). The grain weevils (Curculionidae) are major pests of stored grains such as wheat, maize, and rice. The rice weevil, Sitophilus oryzae L. (Coleoptera: Curculionidae), is one of the most hazardous stored grain pests throughout the world (Pugazhvendan et al. 2009). Females deposit eggs into grain; larvae are legless and remain in the grain kernel for their entire duration. Feeding of S. oryzae larvae and adults can bring down grain weight by up to 75%, diminishing the dietary benefit and germination of the grains resulting in lower prices for seed grain (Dal Bello et al. 2000).

The protection of stored grain using synthetic pesticides is still the method of choice (Rattan 2010). However, many problems are associated with these chemicals, such as insect resistance, toxicity to mammals and other living organisms, toxic residues in stored products, increasing costs of application, and environmental contamination (Arthur et al. 2014). Therefore, there is interest in finding alternative ways for stored products protection. Natural pesticides with low mammalian toxicity are recommended for suppressing insect pests especially in storage (Parugrug and Roxas 2008).

Botanicals are plant-inferred materials that can be utilized as an important component in integrated pest management (IPM) for prevailing insect pests (Abdelsalam et al. 2019; Salem et al. 2020; Mosa et al. 2021; Salem et al. 2021; Moustafa et al. 2021). Botanical pesticides are biodegradable, have little or no deleterious impact on the environment and non-target living beings, cheap, easily produced, and may impede the advancement of resistance (Isman 2005; Rajendran and Sriranjini 2008). Essential oils (EOs) from different botanical parts have shown promising insecticidal activities. EOs from Cymbopogon citratus, C. nardus, and Eucalyptus citriodora have repellent effects against Anopheles arabiensis mosquitoes (Solomon et al. 2012). The EO composition of C. citratus EO showed the presence of neral, geranial, and β-pinene as main compounds, while in C. nardus they were citronellal, nerol, and citronellol; both oils showed low contact toxicity against Dinoderus porcellus L. (Coleoptera: Bostrichidae) (Loko et al. 2021). When tested against three stored grain insects, Sitophilus oryzae, Rhyzopertha dominica, and Tribolium castaneum, plant EOs could play an important role in control of stored-grain insects especially S. oryzae and could be recommended for use as a part of IPM program in stored grains (Tawfeek et al. 2017).

Acetylcholinesterase (AChE, EC 3.1.1.7) is a key enzyme that terminates nerve impulses by catalyzing the hydrolysis of neurotransmitter, acetylcholine, in the nervous system of various organisms. Its inhibition leads to paralysis and death (Zibaee 2011). Alkaline phosphatase (ALP, EC 3.1.3.1) is a set of hydrolytic enzymes, which hydrolyzes phosphomonoesters under alkaline conditions causing cytolysis of tissues during the insect development (Miao 2002). Aspartate aminotransferase (AST, EC 2.6.1.1) and alanine aminotransferase (ALT, EC 2.6.1.2) are also known as glutamate oxaloacetate transaminase (GOT) and glutamate pyruvate transaminase (GPT), respectively. The aminotransferases are important enzymes catalyzing amino acid catabolism and are critical in carbohydrate and protein metabolism (Zibaee et al. 2008). They are altered during various pathological and physiological activities (Etebari et al. 2005).

This work evaluated the insecticidal activities of the extracted oils from four plant species (Cymbopogon citratus, Lantana camara, Artemisia camphorata, and Imperata cylindrica) against Sitophilus oryzae (L.) adults. Their inhibitory effects on the enzyme activities of AChE, ALP, AST, and ALT were estimated.

EXPERIMENTAL

Insect Rearing

The rice weevil Sitophilus oryzae (L.) adults were gathered from infested wheat grains (Triticum aestivum L.). Insect rearing was performed at the laboratory of Applied Entomology and Zoology Department, Faculty of Agriculture (El-Shatby), Alexandria University, Alexandria, Egypt. The culture was maintained in plastic containers (26 × 30 × 20 cm) at 28 ± 2 °C and 65 ± 5 % relative humidity without exposure to any insecticide.

Extraction of Plant Oils

Fresh leaves collected from four plants and authenticated at the Department of Forestry and Wood Technology, Faculty of Agriculture, Alexandria University, Egypt, were used for the oil extraction. Oils from Cymbopogon citratus, Lantana camara, and Artemisia camphorata were extracted by the Clevenger method, where 100 g fresh weight from each material were cut to small pieces and put in 2-L flask containing 1000 mL of distilled water and subjected to hydrodistillation for 3 h to extract the essential oil (EO) (Salem et al. 2014, 2020). For Imperata cylindrica, 100 g were cut into small pieces, extracted by soaking in n-hexane solvent (200 mL) for 6 h under shaking, and filtered with Whatman No. 1 filter paper under suction pressure (Hamada et al. 2018). The n-hexane oily liquid extract (HOE) was separated and concentrated by evaporating the n-hexane solvent under vacuum using a rotary evaporator at 45 °C. The HOE was stored in brown tubes (Salem et al. 2019; Mohamed et al. 2020).

Contact Toxicity

The insecticidal activity of the four oils was tested against S. oryzae adults by the residual film technique (Qi and Burkholder 1981), under laboratory conditions at 12:12 h (light: dark photoperiod). Serial dilutions of tested oils from C. citratus, L. camara, A. camphorata, and I. cylindrica at 2, 5, 10, 20, 30, and 60 mg/cm2 were prepared in acetone as a solvent. There were four replicates for each treatment in addition to controls. One mL of each concentration was placed on the bottom of each Petri dish (9 cm diameter). After the acetone was evaporated, 10 adult of rice weevils were placed into each dish. The same procedure was used for the control treated with acetone. Mortality percentages were recorded after 72 h of treatment. The LC50 and LC90 values as well as the slope of lines were calculated according to Finney (1952). It should be noted that, according to the residual film technique used in the bioassay test, the concentration is attributed to the area Unit. A unit of area means a spread rate at which one milligram of a substance is spread over the area of one square centimeter, which used as a dose calculation unit.

Biochemical Assays

Homogenate preparation

After treatment with LC50 values of the tested oils, S. oryzae adults were separately weighted and homogenized in 10 volumes (w/v) of ice cold 0.1 M phosphate buffer (pH 7.2) using a Teflon glass tissue homogenizer. In the ALP assay, the samples were homogenized in 10 volumes (w/v) of ice cold 0.1 M phosphate buffer (pH 9.8). The homogenates were centrifuged at 5000 rpm for 30 min at 4 °C. The obtained supernatants were divided into small portions and stored at -20 °C. Three replicates were used for each treatment.

Total soluble protein

The total protein content was determined calorimetrically according to the method of Lowry et al. (1951) by using bovine serum albumin (BSA) as a standard.

AChE activity

Acetylcholinesterase (AChE) activity was determined according to Ellman et al. (1961). The reaction mixture contained 2.78 mL of 0.1 M phosphate buffer (pH 8.0); 0.1 mL of ten times diluted DTNB reagent solution (39.5 mg of 5, 5-dithiobis-[2-nitro-benzoic acid] and 15 mg of sodium bicarbonate dissolved in 10 mL of 0.1 M phosphate buffer, pH 7.2); 0.1 mL of the supernatant; and 0.02 mL of 0.075 M acetylthiocholine iodide. The yellow color formed was measured after 10 min at 412 nm. Specific activity was expressed as µmol acetylthiocholine hydrolyzed/min/mg protein.

Alkaline phosphatase activity

Alkaline phosphatase (ALP) activity (Klin 1972) was determined using a diagnostic kit (Diamond Co., Cairo, Egypt). In this method, 20 μL of the enzyme source was added to 1000 μL of 1.0 M diethanolamine buffer (pH 9.8) containing 0.6 mM magnesium ions and 1 mM p-nitrophenyl phosphate, mixed in the cuvette, incubated for 30 sec. With the stopwatch started simultaneously, the output was read again after exactly 1, 2, and 3 min at 405 nm using a spectrophotometer (Sequoia-Turner Model 340; Texas City, TX, USA). Specific activity was expressed as IU/mg protein/hr.

Transaminases activity

Aspartate aminotransferase (AST) and alanine aminotransferase (ALT) activities were determined as previously described (Reitman and Frankel 1957). In this method, 100 μL of enzyme source was added to 500 μL of 0.1 M phosphate buffer (pH 7.2) containing 80 mM L-aspartate as a substrate for AST or 80 mM D-L-alanine as a substrate for ALT, and 4 mM ά-ketoglutarate.

After incubating the mixture for 30 min at 37 °C, 500 μL of developing color reagent (4 mM 2, 4-dinitrophenylhydrazine) was added, and the solution was incubated for 20 min at room temperature. Lastly, 5 mL of 0.4 N NaOH was added, and the mixture was left at room temperature for 5 min. An assay mixture without enzyme source was used as the blank, and the absorption was measured at 546 nm. AST and ALT specific activities were expressed as IU/mg protein/hr.

GC–MS Analysis of the Oils

The chemical composition of the extracted oils was analyzed with a Trace GC Ultraho-ISQ mass spectrometer (Thermo Scientific, Waltham, MA, USA) with a direct capillary column TG–5MS (30 m × 0.25 mm × 0.25 µm film thickness). Oils were diluted in n-hexane solvent (3 n-hexane: 1 oil) before being injected to the GC–MS. The used carrier gas was He (flow rate of 1 mL/min).

The program and oven temperatures conditions can be found in previous work (Moustafa et al. 2021). The Xcalibur 3.0 data system (Thermo Fisher Scientific) with the Match factor from the GC–MS literature is a very intelligent tool to identify chemical constituents, where the value ≥650 is acceptable to confirm the compounds (El-Sabrout et al. 2019; Behiry et al. 2020; Salem et al. 2020; Abd-Elkader et al. 2021; Ali et al. 2021; Moustafa et al. 2021).

Statistical Analysis

Means were compared for significance using one-way analysis of variance (ANOVA) test with Least Significant Difference (LSD0.05) (2005). Mortality rates were corrected according to Abbott’s formula (Abbott 1925) and plotted against concentrations as log/probit regression lines. LC50, LC90 values, and the toxicity index, as well as the slope of the lines were calculated according to Finney (1952).

RESULTS AND DISCUSSION

Insecticidal Activity of the Extracted Oils

The effects of four natural plant oils from Cymbopogon citratus, Lantana camara, Artemisia camphorata, and Imperata cylindrica on the rice weevil Sitophilus oryzae adults were tested after 72 h from treatment by residual film technique. As shown in Table 1, among the four natural plant oils used, the L. camara EO exhibited the highest effect against the adults of S. oryzae (LC50 = 9.81 mg/cm2), followed by C. citratus EO (LC50 = 10.89 mg/cm2), A. camphorata EO (LC50 = 16.12 mg/cm2), and I. cylindrica HOE (LC50= 36.85 mg/cm2).

Table 1. Contact Toxicity of the Oils against S. oryzae Adults after 72 h from Treatment by Residual Film Technique

In vivo Effect of Oils on AChE and ALP Activities in S. oryzae Adults

The changes in AChE and ALP enzymes activities in S. oryzae are displayed in Table 2. There was no significant difference in the AChE activity of S. oryzae adults after being treated with LC50 of four EOs. The inhibition percentages were 38.8, 41.7, 35.0, and 27.2% after treatment with Eos from C. citratus, L. camara, A. camphorata, and I. cylindrica, respectively, relative to the control. The inhibition percentages in ALP activity were 42.4, 49.3, 28.1, and 18.7% after treatment with C. citratus, L. camara, A. camphorata, and I. cylindrica oils, respectively, relative to the control.

Table 2. Changes in Acetylcholinesterase (AChE) and Alkaline Phosphatase (ALP) Activities of S. oryzae Adults as Affected by LC50 of Extracted Oils

In vivo Effect of Oils on AST and ALT Activities in S. oryzae Adults

Table 3 shows the inhibitory effects in the activities of both AST and ALT in S. oryzae adults. The inhibition percentages of AST activity were 33.9, 38.7, 20.8, and 11.8% after being treated S. oryzae adults with C. citratus, L. camara, A. camphorata, and I. cylindrica oils, respectively, compared with the control. Meanwhile, the inhibition percentages of ALT activity were 22.7, 30.5, 14.6, and 9.6% after treating S. oryzae adults with the same previous extracts compared with the control.

Table 3. Changes in Aspartate Aminotransferase (AST) and Alanine Aminotransferase (ALT) Activities of S. oryzae Adults as Affected by LC50 of the Extracted Oils

Chemical Composition of the Oils

Table 4 presents the chemical components of the EO from C. citratus, where the highest abundant compounds were geranial (25.95%), nerylacetal (8.85%), neral (8.45%), linalool oxide (6.36%), 2-furanmethanol (5.4%), cis-linalool oxide (4.29%), 10-hydroxygeraniol (3.92%), cis-verbenol (3.85%), cis-linalool oxide (2.38%), 4-methyl-valeric acid (2.25%), and sobrerol 8-acetate (2.19%).

Table 5 shows the chemical composition of the EO from L. camara corresponding to the highest abundant compounds caryophyllene (12.2%), 3-elemene (8.89%), Germacrene D (5.46%),1,2-cyclopentanedione (3.73%), 5-hydroxymethylfurfural (3.57%), 2,3-dihydro-benzofuran (coumaran) (3.53%), geranyl vinyl ether (2.98%), 2,4-dihydroxy-2,5-dimethyl-3(2H)-furan-3-one (2.97%), adipic acid dimethyl ester (2.73%), 3,7-dimethyl-2,3-epoxy-6-octenol (2.71%), hydroxy-dimethyl ester butanedioic acid (2.69%), 2-furylmethanol (2.47%), 2(5H)-furanone (2.11%), and 1-ethylpentyl acetate (2.02%).

The chemical composition of the EO from A. camphorata is shown in Table 6, where the highest abundant compounds were germacrene D-4-ol (20.83%), borneol (19.47%), 1,8-cineole (7.71%), longiverbenone (5.15%), ascaridol (4.42%), camphor (4.16%), cyperotundone (2.86%), cedran-8-ol (2.53%), 7-epi-silphiperfol-5-ene (2.08%), (E)-isovalencenol (2.04%), β-dihydroagarofuran (2.03%), (Z)-piperitol (1.90%), and α-bisabolol oxide A (1.76%).

Table 4. EO Composition of Cymbopogon citratus leaves by GC-MS

Table 5. EO Composition from Lantana camara Leaves

Table 6. EO Composition from Artemisia camphorata Leaves

Table 7 presents the chemical compounds of I. cylindrica leaf HOE, where the highest abundant compounds were 5-phenylundecane (10.68%), 6-phenyldodecane (8.70%), 2-phenylundecane (5.84%), 4-phenylundecane (5.15%), 3-phenylundecane (5.08%), 2-phenyldodecane (4.98%), 3-phenyldodecane (4.65%), 6-ethyl-5-hydroxy-2,3,7-trimethoxynaphthoquinone (4.34%), 5-phenyldecane (4.31%), 6-phenyltridecane (4.09%), 4-phenyldecane (3.84%), 3-phenyldecane (3.76%), 2-phenyldecane (3.47%), 2-phenyltridecane (3.24%), 4-phenyltridecane (3.25%), and 3-phenyltridecane (3.13%).

Table 7. EO Composition from of Imperata cylindrica Leaves

Among botanical extracts, oil extracts have received much attention as pest control agents (Campolo et al. 2018). They are valuable characterized by less persistence in air, low toxicity to human and animals, high volatility, and toxicity to insect pests of stored products (Batish et al. 2008).

In the present work, essential oil from L. camara showed the highest potential insecticidal activity against the adults of S. oryzae, followed by C. citratus, A. camphorata, and I. cylindrica with LC50 values of 9.81 mg/cm2, 10.89 mg/cm2, 16.12 mg/cm2, and 36.85 mg/cm2, respectively.

Oil extracted from Cymbopogon species have shown potential repellent activity against storage insect pests, S. oryzae, Tribolium castaneum, Oryzaephilus surinamensis, and Sitophilus zeamais (Saljoqi et al. 2006; Olivero-Verbel et al. 2010; Hernandez-Lambraño et al. 2015). By GC-MS, β-citral (43.63%) and geranial (41.51%), were the highest abundant compounds in the oil from C. citratus grown in Egypt (Mansour et al. 2020); also geranial (42.2%) and neral (31.5%), were the main compounds of the oil from Algerian C. citratus (Boukhatem et al. 2014), and from Brazil, where geranial (α-citral) and neral were main compounds (Silva et al. 2008). In the present study, germacrene D-4-ol, borneol, 1,8-cineole, longiverbenone, ascaridol, and camphor were the highest abundant co compounds in A. camphorata EO. A. camphorata showed the major compounds germacrene D-4-ol, 1,8-cineole, ascaridole and borneol with percentages of 22, 12, 10, and 10 %, respectively in Brazilian plant (Seixas et al. 2018). A. camphorata EO showed the presence of davanone (20%) as major component in Tunisia plant (Mohsen and Ali 2009).

The most interesting finding in this study is the potent contact toxicity of Lantana camara oil against the rice weevil, S. oryzae followed by C. citratus oil then ending with A. camphorata and Imperata cylindrica oil by residual film technique. These findings were in accordance with the results of Morya et al. (2010), which demonstrated the efficacy of L. camara (L.) powdered leaves against the stored grain insect pest Corcyra cephalonica (Stainton). Also, L. camara hexane extract could be used as protectant against maize weevil, S. zeamais Motsch. and bean weevil, Callosobruchus maculatus (F.) (Ogunsina et al. 2011).

Both C. citratus powder and methanol extract were promised as insecticides and can be used effectively in the management of S. oryzae in storage (Uwamose et al. 2017). The powdered leaves of Spear grass (I. cylindrica) could serve as a safer alternative to synthetic insecticides for the maize weevil (S. zeamais) infestation in stored maize grains (Ehisianya et al. 2019). Extracts from L. camara leaf were used for the stored maize protection against the infestation of S. zeamais (Ayalew 2020). Recently, Loko et al. (Loko et al. 2021) showed the good potential of C. citratus as both antifeedant and fumigant toxic agent against Dinoderus porcellus Lesne.

Furthermore, Stefanazzi et al. (2011) stated that L. camara and C. citratus oils can be used in IPM programmes for S. oryzae control. Bhatt et al. (2014) showed the toxicity and the significant reduction in larval weight of 2nd and 6th day’s old larvae of Spodoptera litura as affected by L. camara (fruits). L. camara leaf extracts can be useful as a biorotional insecticide against Cadra cautella (Walker) eggs, larvae and adults (Gotyal et al. 2016). Furthermore, the aqueous extract of C. citratus Stapf leaf extract gave the highest protection on the maize grains against S. zeamais Motsch (Oboho et al. 2017).

Taxodium oil from trees grown in El-Beheira, Egypt, with its main compounds of α-pinene, thujopsene, and cedrol followed by carvone and oils from trees grown in Alexandria, Egypt, with its main compounds germacrene D (14.71%), borneol acetate (6.81%), ledene oxide-(II) (6.13%), and trans(β)-caryophyllene (6.13%) showed the potential LC50 against S. oryzae with values of <50, 28.52, and>50 μL/L, respectively (Abdelsalam et al. 2019). Recently, the acetone extract from leaves of peach trees treated with Ag NPs (15 mL/L) showed LC50 of 955.24 ppm against the rice weevils S. oryzae (Mosa et al. 2021). Essential oils from A. camphorata, A. absinthium, A. annua, A. dracunculus, and A. vulgaris showed potential activity against the melonworm Diaphania hyalinata (Linnaeus 1758) (Lepidoptera: Crambidae) larvae with different levels (Seixas et al. 2018).

The inhibitory effects of four tested oils on acetylcholinesterase (AChE) and alkaline phosphatase (ALP) activities in S. oryzae were examined in vivo. Several extracted oils have been described as inhibitors of AChE and ALP isolated from S. oryzae and other insect species (Kim et al. 2013; Abdelgaleil et al. 2016; Oboh et al. 2017). The inhibition of AChE activity may lead to the accumulation of acetylcholine at neuromuscular junctions, which ultimately led to insect mortality (Rajashekar et al. 2014).

Transaminases link the metabolism of amino acids, lipids, and carbohydrates. Exposure to the extracted oils caused significant decrease in the AST and ALT activities in S. oryzae adults. A similar pattern of reduction in AST (14.29%) and ALT (6.98%) level was observed when cotton leafworm, S. littoralis, was exposed to spinosad (Abd El-Mageed and Elgohary 2006).

From this perspective, the present findings could increase demand and experience in the use of these eco-friendly natural products, which could replace synthetic chemicals for management stored grain protection. Finally, these oils are recommended to be used as eco-friendly natural products and could replace synthetic chemicals for management stored grain protection.

CONCLUSIONS

  1. Cymbopogon citratus, Lantana camara, Artemisia camphorate, and Imperata cylindrica oil extracts showed insecticidal potency against the adults of Sitophilus oryzae.
  2. In terms of the enzymes inhibition percentages, L. camara essential oil gave the best results with AChE, ALP, AST, and ALT.
  3. According to contact toxicity of the oils against S. oryzae adults after 72 h, L. camara EO gave the highest effect followed by C. citratus EO.

ACKNOWLEDGMENTS

This research was funded by the Researchers Supporting Project (RSP-2021/123) King Saud University, Riyadh, Saudi Arabia.

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Article submitted: August 31, 2021; Peer review completed: September 28, 2021; Revised version received and accepted: September 29, 2021; Published: October 4, 2021.

DOI: 10.15376/biores.16.4.7767-7783