Journal of Agriculture and Food Research 12 (2023) 100627 Contents lists available at ScienceDirect Journal of Agriculture and Food Research journal homepage: www.sciencedirect.com/journal/journal-of-agriculture-and-food-research Investigation of amaranth production constraints and pest infestation reduction by basil intercropping Yèyinou Ginette Azandémè-Hounmalon a,*, Jhonn Logbo a, Gbèblonoudo Anicet Dassou b, Landry Lokossi a, Evrard Akpla c, Komi K. Mokpokpo Fiaboe d, Manuele Tamò e a Université Nationale d’Agriculture (UNA), Ecole d’Horticulture et d’Aménagement des Espaces Verts (EHAEV), BP 43 Kétou, Bénin b Université Nationale des Sciences, Technologies, Ingénierie et Mathématiques (UNSTIM), Ecole Nationale Supérieure des Biosciences et Biotechnologies Appliquées (ENSBBA), BP 14, Dassa, Bénin c Université d’Abomey-Calavi (UAC), Laboratoire de la Biomathématique et des Estimations Forestiéres (LABEF), 04 BP 1525, Cotonou, Bénin d International Institute of Tropical Agriculture (IITA), Messa, 2008, Yaoundé, Cameroon e International Institute of Tropical Agriculture (IITA), BP 08-0932, Cotonou, Bénin A R T I C L E I N F O A B S T R A C T Keywords: Amaranth (Amaranthus cruentus L.) is the most consumed leaf vegetable in Benin. A study carried out in south Yield Benin have shown that the production of this vegetable is severely limited by insect pest pressure. The present Pests study aimed to identify the major constraint limiting amaranth production in Ségbana municipality, located in Productivity the north of Benin and proposed sustainable agroecological solutions to farmers. Thus, a survey was conducted Vegetable leaf Intercropping system among 150 farmers in three villages of Ségbana (Lougou, Sokotindji and Piami) through well-structured ques- tionnaires to know the major constraint limiting amaranth production. Agroecological methods for managing pests of this vegetable were offered to farmers through an experimental trial consisting of a Fisher block with four treatments and five replications conducted in Sokoundji village. The control treatment (To) consisted of amaranth in pure culture; the treatment (T1), amaranth plants surrounded by basil plants; (T2) rows of amaranth alternated with basil rows and (T3), amaranth plants alternated with basil plants in all directions. Every 5 days, 5 plants were randomly selected per treatment and the numbers of S. recurvalis and P. basalis were counted. Pest damage and yield per treatment were also assessed. Almost all farmers surveyed (100%) reported pest attacks as the major constraint limiting amaranth production. The results obtained on the abundance and the herbivory rate showed powerful negative effects (Df = 3; P < 0.0001) of the different treatments. Moreover, the treatment T2 (amaranth plants alternated with basil rows) gave the highest yield (1.25 t/ha of fresh leaves) and differed significantly from the other treatments (Df = 3; p = 0.039). The association basil – amaranth reduced the abundance of P. basalis and S. recurvalis and also improved the amaranth productivity with LER = 1.16. The association basil – amaranth is more beneficial than pure cultures because it hosts less of the insect pests studied, provides higher yields and makes rational use of the growing space. 1. Introduction one of the priority sectors to be promoted in the "Plan Stratégique pour la Relance du Secteur Agricole (PSRSA)" and in the "Plan Stratégique de Agriculture is one of the main sectors contributing to economic Développement du Secteur Agricole (PSDSA)." It employs about 4% of development. In West Africa, and specifically in Benin, it employs more the active population, or 60,000 jobs [5]. Actual data indicate that the than 42.36% of the active population, contributes to 75% of export margin generated by market gardening can reach 16.395 million CFA earnings and provides around 70% of jobs [1,2]. The economic contri- francs/ha/year, or 4.31 billion on an area of 263 ha exploited in 2000 bution of agriculture is based on a number of activities; among these, [6]. The growing importance given to vegetable cultivation is due to the market gardening occupies a very important place and contributes primordial role it plays in the economies of most of these African significantly to food sovereignty, the fight against poverty and to family countries [7]. Indeed, vegetables are an important component of daily income [3,4,5]. In Benin current context, market gardening has become diets, and sources of income, particularly in urban and peri-urban areas * Corresponding author. E-mail address: fidelerocher@gmail.com (Y.G. Azandémè-Hounmalon). https://doi.org/10.1016/j.jafr.2023.100627 Received 17 October 2022; Received in revised form 30 April 2023; Accepted 5 May 2023 Available online 6 May 2023 2666-1543/© 2023 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by- nc-nd/4.0/). Y.G. Azandémè-Hounmalon et al. J o u r n a l o f A g r i c u l t u r e a n d F o o d R e s e a r c h 12 (2023) 100627 Fig. 1. Survey site. [4]. They can be grouped according to the part of the plant that is aromatic herb used as relief for blocked nose [29], delicious teas [30], consumed and/or sold. Thus, we find leafy vegetables (amaranth, mosquito repellence [31] and treatment of snake bites [32]. A literature nightshade, cabbage), fruiting vegetables (tomato, chilli) and root veg- survey has revealed that O. gratissimum has anti-microbial activity etables (carrot) which are all important sources of plant proteins, vita- against both gram positive and gram-negative bacteria [33], anti-fungal mins, and trace elements for human and animal nutrition [8]. Amaranth effect against Alternaria alternata, Alternaria tenuissima, TZ10.2.2 and (Amaranthus spp) is a traditional vegetable whose leaves and seeds are TZ8.2.2 [34], insecticidal and repellent effects against various insect consumed in various forms [9,10]. In Benin, there are several varieties, species [35–39]. Several studies have shown a repellent effect on pests but Amaranthus cruentus is the most cultivated and consumed, because when the crop was intercropped with Ocimum species [22,40–42]. leaves are rich in protein, vitamin C, beta-carotene, iron and calcium To increase the production of amaranth, the Government of Benin [11–13]. has set up the "Projet d’Appui au Développement du Maraîchage However, its production is limited by several challenges including Résilient aux Changements Climatiques et d’Amélioration des Revenus abiotic and biotic constraints. Climate change is not only creating des Exploitants" (PADCMaRCARE) in the Segbana municipality to sup- favourable conditions for the development of pests (leafminers, mites, port small-scale producers, mainly women, to improve their income. A whiteflies, aphids, thrips, moths, bugs) and diseases, but is also altering recent study has shown Spoladea recurvalis Fabricius and Psara basalis their modes of transmission [1,14,15]. According to Ref. [4], the dam- Walker as the mains pests encountered on amaranth in south Benin [43]. age caused by these pests can be estimated at 100% yield loss. Attempts The main goal of this study was to identify the real constraints limiting to control these pests consist of the frequent and unreasoned use of amaranth production in north Benin through a survey using synthetic chemical insecticides that can lead to chemical residues on well-structured questionnaires, and then to propose a sustainable ag- fruits and vegetables, biological imbalance, environmental contamina- roecological approach to control two main pests of amaranth, tion, intoxication of people and animals, as well as the appearance of S. recurvalis and P. basalis. pest resistance, the elimination of useful entomofauna, in particular parasitoids [4,16–20]. However, chemical control is not always effective 2. Materiel and methods in the short or medium term due to the use of unsuitable, poorly dosed or overused products. 2.1. First step: survey design and data description Environmental-friendly agricultural practices have retained agron- omists attention these latest years to solve the problems. An interesting The municipality of Segbana (north Benin) is located in the Alibori approach in this respect is the use of pesticide plants [21–23]]. The department and bounded: to the north by Malanville municipality, to the biocidal activities of various plant families used as extracts, essential oils south by Kalale municipality, to the east by the Federal Republic of or repellent odor sources have been demonstrated on various pests [24, Nigeria and to the west by Kandi and Gogounou municipalities (Fig. 1). 25]. Segbana is characterized by a monomodal rainy pattern, with rains Certain aromatic plants, according to their particular odors, can keep occurring May to October with annual rainfall generally between 800 away or eliminate harmful insects. Among these plants are those of the and 1200 mm3 [44] and dry season from October to May. The monthly genus Ocimum which are plants of the Lamiaceae family, rich in aro- relative humidity across the country ranges from a minimum of 44% to a matic essential oils. They are collectively known as the “basils” and are maximum of 99%. represented by more than 150 species cultivated and distributed In 2018, a sampling visit were conducted from 15th July to 30th throughout the tropical and temperate regions [26]. Different basil types August in Segbana municipality. During the survey, three (3) towns are commonly used, including holy basil (O. sanctum L.), sweet or Thai were visited: Lougou, Sokotindji and Piami. The survey was conducted basil (O. basilicum L.), lemon basil (O. citriodorum L.), and tree basil among 150 farmers. The choice of farmers was mainly based on the (O. gratissimum L.) [27,28]. Ocimum gratissimum is an erect hairy importance of amaranth production. A total of 21 questions were asked 2 Y.G. Azandémè-Hounmalon et al. J o u r n a l o f A g r i c u l t u r e a n d F o o d R e s e a r c h 12 (2023) 100627 Fig. 2. Schematic and picture illustration of the different treatments A: amaranth, B: basil; T0: control treatment with amaranth in pure culture; T1: amaranth plants bordered by basil plants; T2: amaranth plants alternated with basil rows; T3: amaranth plants alternated with basil plants in all directions. and the interviews took on average 15 min per farmer. After obtaining 2.3. Data collection information on the social situation, each farmer was asked about the amaranth production, the main pests causing damage to it. They were Sampling started 14 days after transplanting and every 5 days, 5 also asked about the major constraints encountered in amaranth pro- plants were randomly selected per treatment and the numbers of duction and the solutions applied. S. recurvalis and P. basalis were counted and recorded. The pest damages were also evaluated. The mean leaf damage index was identified 2.2. Second step: bioassay following [45] ranking: 0 = no damage; 1 = little damage (pinholes, and/or small holes, small leaf edge parts eaten, shot holes); 2 = medium 2.2.1. Plant material damage (some larger holes and/or larger leaf edge areas eaten); 3 = Ocimum gratissimum L. (basil) and Amaranthus cruentus L. were used heavy damage (many larger holes and/or larger leaf edge areas eaten) in this study. The seeds were purchased from approved shops in and 4 = total damage (destroyed, non-functional leaves). The harvest Cotonou. started 21 days after transplanting for amaranth and 30 days for basil. At harvest, the plants have been cut, counted and classified into three 2.2.2. Experimental design categories: plants totally attacked, plants totally healthy and plants The experimental design (Fig. 2) used in this study was the Fisher partially damaged (damaged but part of which is still marketable). block. Four subplots per bloc measuring 6 m × 1.2 m were made. The Leaves were then weighed using the Weiheng Brand Scale 10 Kg reach amaranth nursery was carried out one week after sowing the basil with 5 g. The Land Equivalent Ratio (LER) was also calculated using the ∑ plants. The nurseries of both crops were realized directly on the ground. formula LER = (Ypi/Ymi), where Yp is the yield of each crop or va- The portion of land delimited of that was disinfected with hot water to riety in the intercrop or polyculture, and Ym is the yield of each crop or kill any pests hidden in the ground. After 24 h, the ground was well variety in the sole crop or monoculture. For each crop (i) a ratio is plowed and mixed with compost; the beds were then dressed, watered calculated to determine the partial LER, then the partial LERs are and the sowing lines were drawn using a piece of wood, furrows spaced summed to give the total LER for the intercrop [46–48]. An LER value of with 1 cm of deep. For sowing, amaranth and basil seeds were mixed 1.0, indicates no difference in yield between the intercrop and the with a little fine sand and the blend was spread in the furrows. Next, the collection of monocultures [48–50]. Any Value greater than 1.0 in- nursery was covered with a thin layer of sand, watered it and cover it dicates a yield advantage for intercrop. The results obtained were esti- with straws then treated with fungicide Topsin M (10 g mixed in 11L of mated per hectare. water). Three days later for amaranth and ten for basil, the straws were removed and replaced by a shadehouse. The young seedlings with 3-4 2.4. Data analysis well-developed leaves were transplanted three weeks later for amaranth (with a density 20 cm × 20 cm) and four weeks for basil (with The data from the survey were analyzed by using analysis of variance a density 30 cm × 30 cm) on well-dressed beds. One week after trans- followed by the Bonferroni test according to the losses in production planting, plants were fertilized with a mixture of manure, chicken estimated by the farmers. We determined the pest abundances by droppings and cow purse all at the rate of 10 tones/hectare, i.e. 7.2 kg/ calculating the number of larvae collected per treatment. Generalized subplot. Because of the poorly state of soil, in addition to organic fer- Linear Models (GLMs) with the Poisson family were used to determine tilizer, Urea and NPK were also added the second week after trans- the variation in the pest abundances and vegetable yields according to planting at 75 kg/ha (i.e. 10 g/m2). The basil plants were transplanted 2 the different crop associations (treatments). GLMs with the Binomial weeks before the amaranth plants. A reasonable quantity of water was family were used to determine the variation in the herbivory rate ac- used to daily water the amaranth plants manually with watering can cording to the different treatments. To test the direct and indirect in- morning at 7 a.m. and afternoon at 5 pm. teractions between the crop associations and the pest abundances/ During the bioassay, no chemical was applied. Four treatments were herbivory rate, we used Structural Equation Modeling (SEM) with the tested: T0 (Control treatment with amaranth in pure culture); T1 ‘lavaan’ package [51]. Collected data were analyzed using the software (amaranth plants bordered by basil plants); T2 (amaranth plants alter- R version 3.4.2 [52]. nated with basil rows); T3 (amaranth plants alternated with basil plants in all directions) (Fig. 2). 3 Y.G. Azandémè-Hounmalon et al. J o u r n a l o f A g r i c u l t u r e a n d F o o d R e s e a r c h 12 (2023) 100627 Table 1 abundance and pest damage of P. basalis and S. recurvalis of the different Identifications and prioritization of constraints. vegetable associations. The abundances of the two species were higher N◦ Constraints reported Responds on the T0 and T1 treatments Fig. 4 (a and b). Moreover, Structural Production 1 No knowledge in pests control 100% 2 No knowledge in technical route 40% 3 Insufficient technical support 33.33% Conservation 4 Lack of storage store 20% 5 Lack of conservation equipment 25% Marketing 5 insufficient disposal market 60% Fig. 5. Effect of different associations on pest damages Fig. 3. Spoladea recurvalis damages on amaranth leaves. T0: control treatment with amaranth in pure culture; T1: amaranth plants bordered by basil plants; T2: amaranth plants alternated with basil rows; T3: 3. Results amaranth plants alternated with basil plants in all directions. 3.1. Identification and prioritization of constraints Table 2 Abundance relative of P. basalis and S. recurvalis on amaranth. The constraints mentioned by farmers have been prioritized and Treatments Df SE t-value Pr(>|t|) ranked into constraints of production, conservation and marketing. Almost all of the farmers (100%) reported the pest attacks (Table 1) as Psara basalis being the major constraint limiting the amaranth production, while 40% Intercept 116 0.4260 26.763 <0.0001*** T1 116 0.6024 − 4.427 <0.0001*** and 33.33% respectively reported the lack of technical route and sup- T2 116 0.6024 − 9.684 <0.0001*** port. In addition, 20% and 25% respectively estimated that the lack of T3 116 0.6024 − 10.56 <0.0001*** storage warehouse and the lack of conservation equipment are also Spoladea recurvalis limiting factors in amaranth production. The insufficiency of the market Intercept 116 0.7458 26.81 <0.0001*** T1 116 1.0547 − 3.856 0.00019 *** was also reported by a large number of farmers (60%) (Table 1). T2 116 1.0547 − 12.262 <0.0001*** T3 116 1.0547 − 12.262 <0.0001*** 3.2. Pests abundance and damage on amaranth T1: amaranth plants bordered by basil plants; T2: amaranth plants alternated with basil rows; T3: amaranth plants alternated with basil plants in all directions. The Fig. 3, Fig. 4 (a and b) and Fig. 5 showed respectively the relative Fig. 4. Effect of different associations on P. basalis (a) and S. recurvalis (b) abondances T0: control treatment with amaranth in pure culture; T1: amaranth plants bordered by basil plants; T2: amaranth plants alternated with basil rows; T3: amaranth plants alternated with basil plants in all directions. 4 Y.G. Azandémè-Hounmalon et al. J o u r n a l o f A g r i c u l t u r e a n d F o o d R e s e a r c h 12 (2023) 100627 Fig. 6. Structural Equation Models (SEM) showing the relationship between the different configurations of vegetable associations and the abundances of the two pests and then the relationship between the abundances of the pests and the herbivory rate. Fig. 7. Effect of different associations on vegetable yields T0: control treatment with amaranth in pure culture; T1: amaranth plants bordered by basil plants; T2: amaranth plants alternated with basil rows; T3: amaranth plants alternated with basil plants in all directions. Equation Models showed a significant difference between the different amaranth yield. The yield varied from 0.97 ± 0.04 (Control treatment) types of association (Df = 3; P < 0.0001) that exerted a powerful to 1.25 ± 0.09 (T2: amaranth plants alternated with basil rows) negative effect on the pest abundance and damage of P. basalis and (Table 3). The treatment T2 (amaranth plants alternated with basil S. recurvalis (Table 2) (see Fig. 5). rows) showed the highest yield (1.25 ± 0.09) followed by the T3 (amaranth plants alternated with basil plants in all directions) (1.11 ± 3.3. Relationship between pest abundance and pest damage 0.06) and T1 treatments (amaranth plants bordered by basil plants) (1.07 ± 0.1) (Fig. 7). The Generalized Linear Models (GLMs) showed a Fig. 6 showed on the one hand the relationship between the different significant difference in vegetable yields across the different associations associations of vegetables and the relative abundances of the two pests (Df = 3; P = 0.039). and on the other hand the relationship between the abundances of the pests and the rate of herbivory. Structural Equation Models showed 4. Discussion significant effects of the abundance of P. basalis (Df = 3; P < 0.0001) and S. recurvalis (Df = 3; P < 0.0001) on herbivory rate. In addition, the This study carried out in northern Benin have showed constraints abundance of pests has significant positive effects but is weak on the relative to the amaranth production and the use of environmental- herbivory rate (Fig. 6). friendly agricultural practices such as approach in this respect like plants pesticide to control pests [21–23]. Our results showed there are 3.4. Effects of different associations on the amaranth productivity many factors that hinder the production of this vegetable. As limiting constraints, few farmers reported the lack of technical route and support Fig. 7 showed the effect of different cropping systems on the while other estimated that the lack of storage warehouse and the 5 Y.G. Azandémè-Hounmalon et al. J o u r n a l o f A g r i c u l t u r e a n d F o o d R e s e a r c h 12 (2023) 100627 Table 3 insect pests [64–67]. Yield agronomic evaluation. These properties of the genus Ocimum have been demonstrated on Treatments Yield (t/ha) Land Equivalent Ratio (LER) various arthropods [68–70]. In fact, the biocidal effect of these plant species to control pests would be linked to phenol, which is a phyto- Amaranth Basil Total Amaranth Basil Total chemical compound present in the plant [71,72]. The low infestation of T0 0.97 ± – 0.97 ± 1 – – the insect pest population would therefore be linked to these egg-laying 0.04 0.04 T0’ – 1.41 ± 1.41 disturbances and to the phenol [73]. Thus, the volatile compounds ± – 1 – 0.12 0.12 emitted by basil plants would have disturbed the oviposition of P. basalis T1 0.47 ± 0.6 ± 1.07 ± 0.49 0.41 0.90 and S. recurvalis on amaranth plants. This could then explain on the one 0.03 0.07 0.1 hand the low numbers of pests recorded in treatments T2 (amaranth T2 0.51 ± 0.74 ± 1.25 ± 0.65 0.51 1.16 plants alternated with basil rows) and T3 (amaranth plants alternated 0.03 0.06 0.09 T3 0.43 ± 0.68 ± 1.11 ± 0.44 0.47 0.90 with basil plants in all directions). On the other hand, the activity of the 0.02 0.04 0.06 natural enemies of these pests could also influence. Indeed, in pear or- Df 3 chards associated with O. basilicum, [74] demonstrated that O. basilicum P 0.039 emits volatile substances attractive to auxiliary insects. Thus, basil T0: Control treatment with amaranth in pure culture; T0’: Control treatment would have attracted the natural enemies of these pests in the associated with basil in pure culture T1: amaranth plants bordered by basil plants; T2: crop plots. These natural enemies would have contributed to reduce the amaranth plants alternated with basil rows; T3: amaranth plants alternated with number of pests in plots of associated amaranth crops. The so-called basil plants in all directions. natural enemy hypothesis of [75,76] that natural regulation by para- sitism, predation, etc. is more effective in intercropping systems than in conservation equipment follow by insufficiency of the market. Almost pure cropping systems is verified. all of them (100%) reported the pest attacks as being the major The best yields were obtained on T2 (amaranth plants alternated constraint in the amaranth production. Our results confirms those of with basil rows) and T3 (amaranth plants alternated with basil plants in [43,53,54] who have shown pest attack as the important factor limiting all directions) while the lowest were on T0 (Control treatment with the production of this vegetable. Apart from pests attack [53], reported amaranth in pure culture) and T1 (amaranth plants bordered by basil certain constraints related to production, processing, distribution and plants). The Structural Equation Models showed significant negative marketing who hinder its development and maintain it in the status of a effects of pest abundances on intercropping but the effects are weaker neglected and underused species. It is therefore necessary to develop this for T2 and T3 treatments. This confirms the hypothesis that Ocimum production and diversify amaranth products for national and interna- plants secrete volatile substances that repel pests [77,78], so crop as- tional markets, knowing that it is a very well vegetable appreciated in sociations would have harbored natural enemies [79]. By these SEM, we Beninese dishes. understood that the two pests increase the rate of herbivory but not Environmental-friendly agricultural practices like crop association more important than in monoculture. These results confirm the insec- appears to be an interesting approach [21–23] that offer major potential ticidal and biocidal properties of basil since the treatments that have in market gardening, in reducing the use of chemical products while been the most effective on pest populations are also those that have maintaining a satisfactory level of yield in terms of quality and quantity induced the best yields of amaranth. [55]. In the second part of this study, we evaluated the effect of basi- The LER in the T2 (amaranth plants alternated with basil rows) = l/amaranth to control P. basalis and S. recurvalis, two major pests 1.16 is greater than 1. This means that the association amaranth plants limiting amaranth production [54]. The results showed that treatments alternated with basil rows have a potential interest in rationalizing the T2 (amaranth plants alternated with basil rows) and T3 (amaranth cultivable space. The amaranth plants alternated with basil rows (T2) plants alternated with basil plants in all directions) harbored fewer pests could save 16% of the area compared to the pure culture of basil and overall than T1 (amaranth plants bordered by basil plants) and T0 amaranth. Our results are similar to those obtained by Refs. [80,81] on (Control treatment with amaranth in pure culture) treatments. Indeed, the cowpea/maize association, [82] on the cabbage/basil association several studies have shown that cabbage plots in association with basil and [59] on gboma/basil. According to these authors, the LER obtained plants show less infestation by Spodoptera littoralis Boisd, Plutella xylos- were respectively 1.46, 1.07, 1.2, 1.11 et 1.40. The amaranth/basil tella Linnaeus and Hellula undalis Fabricius than those of cabbage in pure cropping association would be a cropping system that allows better culture [22,56]. Moreover [57], reported less damage of H. undalis to space management in the of increasing population context with conse- cabbage grown in association with pepper or onion. Similar results were quent pressure on land resources. also reported by Ref. [58] who found a low density of mites Poly- phagotarsonemus latus Banks in the gboma field by using an aqueous 5. Conclusions extract of basil, Neem oil and Sunpyrifos. Other study carried out by Ref. [59] in the cropping system gboma/basil showed also low density of From this study, we can conclude that the amaranth production is aphids, Selepa docilis, Bemisia tabaci and mealybugs. Our result could mainly limited by pest attacks and others constraints relative to the also be explained by the insecticidal and repellent properties of basil production, conservation and marketing. As agroecological practice to [34–39]. Indeed, the essential oil of the basil was principally composed control S. recurvalis and P. basalis, the amaranth plants alternated with by the aromatic ether estragol (74.0%) and the terpene with alcohol basil rows are the best cropping system. This spatial arrangement of group linalool (17.8%), which are known to have repellent and toxic these two vegetable crops provides a high yield of amaranth and shelters activities against stored product insect [60]. The repellent effect of ar- natural enemies for their conservation and the natural regulation of omatic plants on insects is generally attributed to their volatile organic vegetable pests. We therefore suggest to continue investigation of other compounds (VOCs) shown to disturb the oviposition of P. xylostella and important vegetables more sensitive to arthropod pests in order to Pieris brassicae Linnaeus on cabbage plants in association with clover explore the potential of basil to really control these pests of vegetable [61]. According to the same authors, this disturbance is much more crops. linked to the presence of repellent compounds than to the physical appearance of the clover. In fact, volatile organic compounds (VOCs) Funding emitted by aromatic plants greatly influence the process of localization of host plants by pests [62,63]. The genus Ocimum, has been also This work was supported by the Ecole d’Horticulture et d’Am- investigated with regard to its insecticidal properties against diverse énagement des Espaces Verts (EHAEV) of Université Nationale 6 Y.G. Azandémè-Hounmalon et al. J o u r n a l o f A g r i c u l t u r e a n d F o o d R e s e a r c h 12 (2023) 100627 d’Agriculture (UNA) and International Institute of Tropical Agriculture Environnement-Santé et Développement Durable, Centre Interfacultaire de of Benin (IITA-Benin) for their financial supports. Formation et de Recherche en Environnement pour le Développement Durable, Université d’Abomey-Calavi, 2010, p. 74. [19] C. Ahouangninou, B. Fayomi, T. Martin, Evaluation des risques sanitaires et Declaration of competing interest environnementaux des pratiques phytosanitaires des producteurs maraîchers dans la commune rurale de Tori-Bossito (Sud-Bénin), Cah. Agric. 20 (3) (2011) 216–222. [20] C. Ahouangninou, T. Martin, P. Edorh, S. Bio-Bangana, S. Onil, L. St-Laurent, The authors declare that they have no conflict of interest. S. Dion, B. Fayomi, Characterization of health and environmental risks of pesticide use in market-gardening in the rural city of Tori-Bossito in Benin, West Africa, Data availability J. Environ. Protect. 3 (3) (2012) 241–248. [21] C.A. Amoatey, E. Acquah, Basil (Ocimum basilicum) intercrop as a pest management tool in okra cultivation in the Accra plains, Ghana J. Hortic. 8 (2010) 65–70. The authors do not have permission to share data. [22] B.B. Yarou, F. Assogba-Komlan, E. Tossou, A.C. Mensah, S. Simon, F.J. Verheggen, F. Francis, Efficacy of basil-cabbage intercropping to control insect pests in Benin, West Africa Comm, Appl. Biol. Sci., Ghent University 82/2 (2017). Acknowledgements [23] A. Mkindi, N. Mpumi, Y. Tembo, P.C. Stevenson, P.A. Ndakidemi, K. Mtei, R. Machunda, S.R. Belmain, Invasive weeds with pesticidal properties as potential The authors are grateful to Ecole d’Horticulture et d’Aménagement new crops, Ind. Crop. Prod. 110 (2017) 113–122, https://doi.org/10.1016/j. indcrop.2017.06.002. des Espaces Verts (EHAEV) of Université Nationale d’Agriculture (UNA) [24] S.E. Halbert, D. Corsini, M. Wiebe, S.F. Vaughn, Plant-derived compounds and and International Institute of Tropical Agriculture of Benin (IITA-Benin) extracts with potential as aphid repellents, Ann. Appl. Biol. 154 (2009) 303–307, for their financial and technical supports. https://doi.org/10.1111/j.1744-7348.2008.00300.15. [25] N. Zapata, E.J.M. Van Damme, M. Vargas, L. Devotto, G. Smagghe, Insecticidal activity of a protein extracted from bulbs of Phycella australis Ravenna against the References aphids Acyrthosiphon pisum Harris and Myzus persicae Sulzer, Chil. J. Agric. Res. 76 (2016) 188–194, https://doi.org/10.4067/S0718-58392016000200010. [1] ACEDP (Agricultural Competitiveness and Export Diversification Project), Project [26] A.K. Pandey, P. Singh, N.N. Tripathi, Chemistry and bioactivities of essential oils of ID, 2020, P168132. some Ocimum species: an overview, Asian Pac. J. Trop. Biomed. 4 (2014) 682–694. [2] FAO Cultures Maraîchères au Bénin, Atelier de validation de fiches et manuels [27] T. Juntachote, E. Berghofer, S. Siebenhandl, F. Bauer, The antioxidative properties techniques de production maraîchère, 2015. of Holy basil and Galangal in cooked ground pork, Meat Sci. 72 (2006) 446–456. [3] R. Kahane, L. Temple, P. Brat, H. De Bon, Les légumes feuilles des pays tropicaux: [28] N. Mahajan, S. Rawal, M. Verma, M. Poddar, S. Alok, A phytopharmacological Diversité, richesse économique et valeur santé dans un contexte très fragile. overview on Ocimum species with special emphasis on Ocimum sanctum, Biomed. Colloque Angers 7-9 septembre 2005-03-14 : "Les légumes : un patrimoine à Prev. Nutr. 3 (2013) 185–192. transmettre et ̀a valoriser Thème III : Utilisation et perception". CIRAD département [29] J.O. Kokwaro, Medicinal Plants of East Africa, second ed., Kenya Literature Bureau, Flhor, Bd de la Lironde, 34398 Montpellier cedex 5, 2005, 9p. Nairobi, 1993. [4] B. James, C. Atcha-Ahoué, I. Godonou, H. Baimey, G. Goergen, R. Sikirou, M. Toko, [30] FAO, Mixed crop-livestock farming: a review of traditional technologies based on Gestion intégrée des nuisibles en production maraichère : guide pour les agents de literature and field experience, FAO Anim. Prod. Health Pap. (2001) 152. vulgarisation en Afrique de l’Ouest, Ibadan, Nigéria, 2010. IITA. [31] C.W. Githinji, J.O. Kokwaro, Ethnomedicinal study of major species in the family [5] I. Yolou, I. Yabi, F. Kombieni, P.G. Tovihoudji, J.A. Yabi, A.A. Paraïso, F. Afouda, Labiatae from Kenya, J. Ethnopharmacol. 39 (1994) 197–203. Maraîchage en milieu urbain à Parakou au Nord-Bénin et sa rentabilité [32] B.O. Owuor, B.A. Mulemi, J.O. Kokwaro, Indigenous snake bite remedies of the Luo économique, Int. J. Innov. Sci. Res. 19 (2) (2015) 290–302. of western Kenya, J. Ethnobiol. 25 (2005) 129–141. [6] H.F. Kakai, S.G. Kakai, A.G. Tohouegnon, Agriculture urbaine et valorisation des [33] L.G. Matasyoh, J.C. Matasyoh, F.N. Wachira, M.G. Kinyua, A.T.M. Muigai, T. déchets au Bénin: une approche de développement durable, [VertigO] La revue K. Mukiama, Chemical composition and antimicrobial activity of the essential oil of électronique en sciences de l’environnement 10 (2) (2010). URI: https://id.erudit. Ocimum gratissimum L. growing in eastern Kenya, Afr. J. Biotechnol. 6 (2007) org/iderudit/045516ar. 760–765. [7] E. Adango, A. Onzo, R. Hanna, P. Atachi, J. Braima, Inventaire de la faune des [34] A. Marco, S. Santos, J. Caetano, M. Pintado, E. Vieira, P.R. Moreira, Basil essential acariens sur Amaranthus cruentus (Amaranthaceae), Solanum macrocarpon et oil as an alternative to commercial biocides against fungi associated with black Solanum aethiopicum (Solanaceae) dans le Sud Bénin, Int. J. Trop. Insect Sci. 26 stains in mural painting, Build. Environ. 167 (2020), 106459. (2006) 155–165, https://doi.org/10.1079/IJT2006115. [35] S.M. Kéita, C. Vincent, J.P. Schmidt, Efficacy of essential oil of Ocimum basilicum L. [8] PNUD, Renforcement des capacités des maraîchers des Communes d’Adjohoun, and O. gratissimum L. applied as an insecticidal fumigant and powder to control Aplahoué Bopa, Malanville Ouaké, Sô-Ava sur la protection phytosanitaire aux fins Callosobruchus maculatus (Fab.) (Coleoptera: Bruchidae), J. Stored Prod. Res. 37 de l’adaptation aux changements climatiques : Programme intégré d’adaptation (2001) 339–349. pour la lutte contre les effets néfastes des changements climatiques sur la [36] K. Murugan, P. Murugan, A. Noortheen, Larvicidal and repellent potential of production agricole et la sécurité alimentaire au Bénin (PANA1), in: Dépôt légal Albizia amara Boivin and Ocimum basilicum L. against dengue vector, Aedes aegypti n◦7754 du 03/02/2015 1er trimestre Bibliothèque Nationale, 2015, p. 80, 978- (Insecta: Diptera: Culicidae), Bio Technol. 98 (2007) 198–201. 99919-0-363-7. [37] P. Singh, R.H. Jayaramaiah, P. Sarate, H.V. Thulasiram, M.J. Kulkarni, A.P. Giri, [9] B. Písaříková1, S. Kráčmar, I. Herzig1, J. Czech, Amino acid contents and biological Insecticidal potential of Defense Metabolites from Ocimum kilimandscharicum value of protein in various amaranth species, Anim. Sci. 50 (4) (2005) 169–174. against Helicoverpa armigera, PLoS One 9 (2014), e104377. [10] M.W. Mburu, N.K. Gikonyo, G.M. Kenji, A.M. Mwasaru, Nutritional and functional [38] M.L. Bhavya, A.G.S. Chandu, S.S. Devi, Ocimum tenuiflorum oil, a potential properties of a complementary food based on kenyan amaranth grain (Amaranthus insecticide against rice weevil with antiacetylcholinesterase activity, Ind. Crop. cruentus), Afric. J. Food Agric. Nut. Dev. 2 (2) (2012) 1–19. Prod. 126 (2018) 434–439. [11] A.S. Adekambi, P.Y. Adegbola, Analyse des systèmes de production des légumes : [39] Á. Rodríguez-González, S. Álvarez-García, Ó. González-López, F.D. Silva, P. Rapport d’étude, Ministère de l’Agriculture de l’Elevage et de la Pêche), 2008, A. Casquero, Insecticidal properties of Ocimum basilicum and Cymbopogon p. 33p. winterianus against Acanthoscelides obtectus, insect pest of the common bean [12] E.G. Achigan-Dako, E.D. Olga, P.M. Sogbohossou, Current knowledge on (Phaseolus vulgaris L.), Insects 10 (2019) 151. Amaranthus spp.: research avenues for improved nutritional value and yield in [40] G.B. Tang, B.Z. Song, L.L. Zhao, X.S. Sang, H.H. Wan, J. Zhang, Y.C. Yao, Repellent leafy amaranths in sub-Saharan Africa, Euphytica (2013), https://doi.org/ and attractive effects of herbs on insects in pear orchards intercropped with 10.1007/s10681-014-1081-9. aromatic plants, Agrofor. Syst. 87 (2013) 273–285, https://doi.org/10.1007/ [13] A.D. Wouyou, E.A. Ahissou, C.B. Gandonou, F. Assogba-Komlan, A. Houngbèmè, F. s10457-012-9544–2. A. Gbaguidi, H. Ahissou, L. Lagnika, S.A. Zanklan, S. Lutts, Salinity increased [41] S. Beizhou, G. Tang, X. Sang, J. Zhang, Y. Yao, N. Wiggins, Intercropping with vitamins concentration in Amaranthus cruentus leaves, Afr. J. Biotechnol. (2017) 6. aromatic plants hindered the occurrence of Aphis citricola in an apple orchard [14] Y. Trottin Caudal, D. Grasselly, P. Millot, Gestion de la protection des plantes. Serre system by shifting predator–prey abundances, Biocontrol Sci. Technol. 23 (2013) et tomate protégée, Centre Technique Inter-professionnel des Fruits et Légumes, 381–395, https://doi.org/10.1080/09583157.2013.763904. Paris (France), 1995. [42] B.B. Yarou, A.H. Bokonon-Ganta, F.J. Verheggen, G.C. Lognay, F. Francis, Aphid [15] G.G. Kennedy, Tomato, Pests, parasitoids and predators: tritrophic interactions behavior on Amaranthus hybridus L. (Amaranthaceae) associated with Ocimum spp. involving the genus Lycopersicon, Annu. Rev. Entomol. 48 (2003) 51–72. (Lamiaceae) as repellent plants, Agro 10 (2020) 736, https://doi.org/10.3390/ [16] B. Ndakidemi, K. Mtei, P.A. Ndakidemi, Impacts of synthetic and botanical agronomy10050736. pesticides on beneficial insects, Agric. Sci. 7 (2016) 364–372, https://doi.org/ [43] A.C.G. Mensah, R. Sikirou, F. Assogba Komlan, B.B. Yarou, G.S.-K. Midingoyi, 10.4236/as.2016.76038. J. Honfoga, M.-E. Dossoumou, G.N. Kpéra, A.K.A. Djinadou, Guide pratique pour la [17] A.I. Sankoh, R. Whittle, K.T. Semple, K.C. Jones, A.J. Sweetman, An assessment of culture de l’amarante (Amaranthus cruentus) au Bénin. Référentiel Technico- the impacts of pesticide use on the environment and health of rice farmers in Sierra Economique (RTE). MAEP/INRAB/FIDA/ProCar/PADMAR/World Vegetable Leone, Environ. Int. 94 (2016) 458–466, https://doi.org/10.1016/j. Center/Bénin. Dépôt légal N◦ 11557, du 26/08/2019, Bibliothèque Nationale (BN) envint.2016.05.034. du Bénin, 3ème trimestre (2019) 52, 978-99982-53-17-9. [18] G.Y. Azandémè, Evaluation de l’efficacité de l’appât protéique GF-120 appliqué sur [44] S. Simon, F. Assogba-Komlan, L. Adjaïto, A. Mensah, H.K. Coffi, M. Ngouajio, deux espèces de plantes de perchoir dans la lutte contre les mouches des fruits T. Martin, Efficacy of insect nets for cabbage production and pest management téphritides des cucurbitacées. Mémoire d’étude pour l’obtention du Master-II en 7 Y.G. Azandémè-Hounmalon et al. J o u r n a l o f A g r i c u l t u r e a n d F o o d R e s e a r c h 12 (2023) 100627 depending on the net removal frequency and microclimate, Int. J. Pest Manag. 60 [65] S.M. Kéita, C. Vincent, J.P. Schmit, J.T. Arnason, A. Bélanger, Efficacy of essential (2014) 208–216, https://doi.org/10.1080/09670874.2014.956844. oil of Ocimum basilicum L. and O. gratissimum L. applied as an insecticidal fumigant [45] S. Toepfer, P. Fallet, J. Kajuga, D. Bazagwira, I.P. Mukundwa, M. Szalai, T. and powder to control Callosobruchus maculatus (Fab.) (Coleoptera: Bruchidae), C. Turlings, Streamlining leaf damage rating scales for the fall armyworm on J. Stored Prod. Res. 37 (2001) 339–349. maize, J. Pest 94 (4) (2021) 1075–1089. [66] M.D. Lopez, M.J. Jordan, M.J. Pascual-Villalobos, Toxic compounds in essential [46] W.A.T. Herrara, R.R. Harwood, The effect of plant density and row arrangement on oils of coriander, caraway and basil active against stored rice pests, J. Stored Prod. productivity of com rice intercrop, Paper presented at the 5th annual convention of Res. 44 (2008) 273–278. the crop science of Philippines 16–18 (1974). Nagar City. [67] J.O. Ogendo, M. Kostyukovsky, U. Ravid, J.C. Matasyoh, A.L. Deng, E.O. Omolo, S. [47] D.C. Pathick, M.L. Malla, Study on the Performance of Crop Legume under T. Kariuki, E. Shaaya, Bioactivity of Ocimum gratissimum L. oil and two of its Monoculture and Intercrop Combination, Sixth annual maize development constituents against five insect pests attacking stored food products, J. Stored Prod. workshop, Nepal, 1979. Res. 44 (2008) 328–334. [48] T. Kurata, A study on farming system in USSA, Quarterly J. Agro. Eco. 26 (1986) [68] S. Del Fabbro, F. Nazzi, Repellent effect of sweet basil compounds on Ixodes ricinus 179–205. ticks, Exp. Appl. Acarol. 45 (3–4) (2008) 219–228. [49] D. Mazaheri, M. Oveysi, Effects of intercropping of two corn varieties at various [69] E.T. Oparaocha, I. Iwu, J.E. Ahanaku, Preliminary study on mosquito repellent and nitrogen levels, Iranian J. Agro. (2004) 71–76. mosquitocidal activities of Ocimum gratissimum (L.) grown in eastern Nigeria, [50] R.L. Agrawal, Emerging trends in cropping system, Indian Farmers Digest 10 J. Vector Borne Dis. 47 (2010) 45–50. (1995) 20–23. [70] T. Kazembe, D. Chauruka, Mosquito repellence of Astrolochii hepii, Cymbopogon [51] Y. Rosseel, lavaan: an R package for structural equation modeling and more. citratus and Ocimum gratissimum extracts and mixtures, Bull. Environ. Pharmacol. Version 0.5–12 (BETA), J. Stat. Software 48 (2012) 1–36. Life Sci. 1 (8) (2012) 60–64. [52] R Core Team, A Language and Environment for Statistical Computing, 2017. http: [71] A. Attou, Contribution à l’étude phytochimique et activités biologiques des extraits //www.R-project.org. de la plante Ruta chalpensis (Fidjel) de la région d’Ain Témouchent. Mémoire du [53] A.P. Agre, A.F. Sanoussi, I. Dossou-Aminoni, A. Dassou, A. Dansi, P. Rudebjer, diplôme de magister en biologie à l’Université d’Abou Bekr Belkaid Tlemcen R. Hall, R. Vodouhe, Plan d’actions stratégiques pour la promotion de la chaine de Algérie, 2011, p. 119. valeur de l’Amarante (Amaranthus spp) au Bénin, Int. J. Neglected Underutilized [72] C.R. Dossoukpevi, M.C. Djaboutou, C.C. Gnimadi, G. Cacaï, G.A. Mensah, C. Species 2 (2016) 33–41. Ahanhanzo, Fiche technique : Détermination des principes actifs des huiles [54] D.M. Mureithi, K.K.M. Fiaboe, S. Ekesi, R. Meyhöfer, Important arthropod pests on essentielles des feuilles des vitroplants et des plantes mères de Ocimum basilicum et leafy amaranth (Amaranthus viridis, A. tricolor and A. blitum) and broad-leafed Ocimum gratissimum cultivés au Bénin. Dépôt légal N 9037 du 15 novembre 2016 African nightshade (Solanum scabrum) with a special focus on host, Afr. J. Hort. Sci. Bibliothèque Nationale (BN) du Bénin, 4ème trimestre 2016, ISBN : 978-99919-2- 11 (2017) 1–17, 2017. 603-2. [55] S. Gaba, F. Lescourret, S. Boudsocq, J. Enjalbert, P. Hinsinger, E.P. Journet, M. [73] S. Puri, S. Singh, S.K. Sohal, Oviposition behaviour and biochemical response of an L. Navas, J. Wery, G. Louarn, E. Malézieux, E. Pelzer, M. Prudent, H. Ozier- insect pest, Zeugodacus cucurbitae (Coquillett) (Diptera: Tephritidae) to plant Lafontaine, Multiple cropping systems as drivers for providing multiple ecosystem phenolic compound phloroglucinol, Comp. Biochem. Physiol. C Toxicol. services: from concepts to design, Agron. Sustain. Dev. 35 (2) (2015) 607–623. Pharmacol. 255 (2022), 109291. [56] F. Assogba-Komlan, B.B. Yarou, A. Mensah, S. Simon, Les légumes traditionnels [74] S. Beizhou, Z. Jie, H. Jinghui, W. Hongying, K. Yun, Y. Yuncong, Temporal dans la lutte contre les bioagresseurs des cultures maraichères : associations dynamics of the arthropod community in pear orchards intercropped with aromatic culturales avec le Tchayo (Ocimum gratissimum) et le Yantoto (Launaea plants, Pest Manag. Sci. 67 (2011) 1107–1114. taraxacifolia). Fiche technique, Cotonou, Bénin, INRAB, 2012. [75] R.B. Root, Organization of a plant-arthropod association in simple and diverse [57] E. Asare-Bediako, A.A. Addo-Quaye, A. Mohammed, Control of icapitata using habitats: the Fauna of collards (Brassica Oleracea), Ecol. Monogr. 43 (1) (1973) intercropping with non-host crops, Am. J. Food Technol. 5 (4) (2010) 269–274. 95–124. [58] E. Adango, A. Onzo, C.O.G.W. Daoudou, Evaluation de l’activité acaricide de [76] E.P. Russel, Enemies hypothesis: a review of the effect of vegetational diversity on quelques biopesticides sur l’acarien tarsonème, Polyphagotarsonemus latus Banks predatory insects and parasitoids, Environ. Entomol. 18 (4) (1989) 590–599. (Acari: Tarsonemidae) infestant l’aubergine gboma (Solanum macrocarpon L.) au [77] F.K. Matu, L.K. Murungi, S. Mohamed, E. Deletre, Behavioral response of the Sud-Bénin, Eur. Sci. J. 16 (15) (2020) 442–463, https://doi.org/10.19044/ greenhouse whitefly (Trialeurodes vaporariorum) to plant volatiles of Ocimum esj.2020.v16n15p442. basilicum and Tagetes minuta, Chemoecology 31 (1) (2021) 47–62. [59] D. Houadakpode, Valorisation des plantes aromatiques dans la gestion intégrée des [78] B.B. Yarou, A.H. Bokonon-Ganta, F.J. Verheggen, G.C. Lognay, F. Francis, Aphid principaux insectes ravageurs de la Grande Morelle au Sud-Bénin: cas de Ocimum behavior on Amaranthus hybridus L. (Amaranthaceae) associated with Ocimum spp. gratissimum et O. basilicum. Mémoire de Master, Liège Université, Gembloux Agro- (Lamiaceae) as repellent plants, Agro 10 (5) (2020) 736. Bio Tech, 2018, p. 69p. [79] T.F. Marcos, L.B. Flor, A.R. Velilla, K.G. Schoenly, J.O. Manalo, O.M. Ofilas, S. [60] V. Rozman, I. Kalinovic, Z. Korunic, Toxicity of naturally occurring compounds of R. Obien, Relationships between pests and natural enemies in rainfed rice and Lamiaceae and Lauraceae to three stored products insects, J. Stored Prod. Res. 43 associated crop and wild habitats in Ilocos Norte, Philippines, in: T.W. Mew, (2007) 349–355. E. Borromeo, B. Hardy (Eds.), Exploiting Biodiversity for Sustainable Pest [61] S. Finch, M. Kienegger, A behavioural study to help clarify how under sowing with Management: Proceedings, Los Baños, Laguna (Philippines), IRRI, 2001, pp. 23–42. clover affects host-plant selection by pest insects of brassica crops, Entomol. Exp. [80] K.E. N’Goran, K.E. Kassin, G.P. Zohouri, M.F.P. N’Gbesso, G.R. Yoro, Performances Appl. 84 (2) (1997) 165–172. agronomiques des associations culturales igname-légumineuses alimentaires dans [62] T.J.A. Bruce, L.J. Wadhams, C.M. Woodcock, Insect host location: a volatile le Centre ouest de la Côte d’Ivoire, J. Appl. Biosci. 43 (2011) 2915–2923. situation, Trends Plant Sci. 10 (6) (2005) 269–274. [81] K. Coulibaly, Analyse des facteurs de variabilité des performances agronomiques et [63] T.J.A. Bruce, J.A. Pickett, Perception of plant volatile blends by herbivorous insects économiques des cultures et de l’évolution de la fertilité des sols dans les systèmes - finding the right mix, Phytochem 72 (13) (2011) 1605–1611. culturaux intégrant les légumineuses en milieu soudanien du Burkina Faso: [64] V.N. Souza, C.R.F. de Oliveira, C.H. Cysneiros Matos, D.K.F. de Almeida, approche expérimentale chez et par les paysans, Thèse de doctorat unique, Fumigation toxicity of essential oils against Rhyzopertha dominica (F.) in stored Université Polytechnique de Bobo Dioulasso, Burkina Faso, 2012, p. 139p. maize grain, Revista Caatinga 29 (2016) 435–440. [82] B.B. Yarou, Le tchayo (Ocimum gratissimum L.) : un légume traditionnel contre les ravageurs du chou, Poster PCM/INRAB, 2013. 8