Insecticidal Activity of Seed Extracts of Carica Papaya (L.) Against the Fall Armyworm Spodoptera Frugiperda (J.E

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Insecticidal activity of seed extracts of Carica papaya (L.) against the fall armyworm Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae)

Rodolfo Figueroa-Brito, Arturo Huerta-de la Peña, Ignacio Pérez Moreno, Vicente Santiago Marco Mancebón and Jesús Francisco López-Olguín

SUMMARY

The present study shows that natural products from Carica giperda. The response variable was insect mortality. Extracts papaya can be considered as a valid alternative to control from seeds of the Maradol, Mammee and Yellow cultivars of pests in agriculture. The insecticide properties of the seed ex- C. papaya, followed by extracts from seeds of the Hawaiian tracts of four cultivars of C. papaya (Maradol, Mammee, Yel- cultivar, applied at concentrations of 10, 100 and 1000ppm, low and Hawaiian) were added to an artificial insect diet. Bio- were toxic on S. frugiperda larvae (50-70% corrected mortal- assays were conducted with hexanic, acetonic and methanolic ity rate). The acetonic extracts of the Maradol and Mammee extracts at concentrations of 10, 100 and 1000ppm. All tests cultivars at 10ppm were the most effective, causing mortalities were performed with the first larval stage of Spodoptera fru- of 73.6 and 62.8% of the larvae, respectively.

Introduction Brazil (12×106ha); Mexico posed primarily of yellow 2002). In Mexico it causes (7.5×10 6ha); and India maize for livestock feed from crop losses from 20 to 100% Maize (Zea mays Linnaeus (6×106ha). the USA (García-Raño and (Del Rincón et al., 2006). 1753, Gramineae) is one of In a global ranking of di- Keleman, 2007). For control, either seed the most important cereal rect human consumption of One of the factors of the treatment with systemic insec- crops in the world, contribut- maize, Mexico is at the top, losses in maize crop in Mexi- ticides or the application of ing to the well-being of mil- exceeded only by Bosnia-Her- co and other maize-growing granulated insecticides scat- lions of poor farmers. It is a zegovina. In 2006, per capita areas of the American Conti- tered on the ground are used. globally important crop and maize consumption in Mexico nent is pest insects, among It has been shown that the preferred staple food for more was ~260kg/person, and the which Spodoptera frugiperda indiscriminate use of these than a billion people in Sub- country dedicated ~12×106ton (J.E. Smith, 1797) (Lepidop- compounds leads to a major Saharan Africa and Latin of maize grain to direct hu- tera: Noctuidae) is the most deterioration of the environ- America, where animal pro- man consumption. Combining important. It attacks during ment and can be harmful for tein sources are not affordable human consumption with all the stages of plant growth the health of both producers by the common people. Of maize dedicated to livestock and it can cause yield reduc- and consumers, and also en- the 1.4×10 8ha grown globally and national reserves, the to- tions of up to 10% (Pingali courages the appearance of with maize, ~96×106ha are tal annual consumption of and Pandey, 2001). This loss resistant pest populations, located in developing coun- maize in Mexico hovers percentage is higher in tropi- eliminating their natural ene- tries. Four countries account around 26×10 6ton. Of this cal and subtropical regions of mies and facilitating a resul- for more than half of the de- amount, ~20×106ton were pro- Latin America, with over 35% tant growth in the impact of veloping world’s maize culti- duced in the country, and an- damage recorded in Colombia secondary pests, which results vation area (Pingali and Pan- other 6-7×10 6ton were im- (Torres y Cotes, 2005) and in a cost increase in the dey, 2001): China (26×106ha); ported, the latter being com- 40% in Cuba (Fernández, growing of maize.

KEY WORDS / Biological Activity / Caricaceae / Maize / Plant Extracts / Toxicity / Received: 06/05/2010. Modified: 09/16/2011. Accepted: 09/19/2011.

Rodolfo Figueroa Brito. Ph.D. Politécnica de Madrid, Spain. Vicente Santiago Marco Mance- Spain. Professor-Researcher, in Strategies for Regional Ag- Professor-Researcher, COL- bón. Doctor Ingeniero Agróno- Benemérita Universidad Autó ricultural Development, Cole- POS, Puebla, México. mo, Universidad Politécnica de noma de Puebla (BUAP), Méxi- gio de Postgraduados (COL- Ignacio Pérez Moreno. Ph.D. in Madrid, Spain. Professor-Re- co. Address: Departamento de POS), México. Professor-Re- Agronomy, Universidad de searcher, Universidad de La Agroecología y Ambiente, In- searcher, Instituto Politécnico Navarra, Spain. Researcher Rioja. Logroño, Spain. stituto de Ciencias, BUAP. Av. Nacional, Morelos, México. and Professor, Universidad de Jesús Francisco López-Olguín. 14 Sur 6301, Edif. 103-A, CU, Arturo Huerta de la Peña. Ph.D. La Rioja, Logroño, Spain. Ph.D. in Agronomy, Universi- 72570-Puebla, México. e-mail: in Agronomy, Universidad dad Politécnica de Madrid, [email protected]

752 0378-1844/11/10/752-05 $ 3.00/0 OCT 2011, VOL. 36 Nº 10 Actividad insecticida de extractos de semillas de Carica papaya (L.) contra el gusano cogollero Spodoptera frugiperda (J.E. Smith) (LEPIDOPTERA: NOCTUIDAE) Rodolfo Figueroa-Brito, Arturo Huerta-de la Peña, Ignacio Pérez Moreno, Vicente Santiago Marco Mancebón y Jesús Francisco López-Olguín RESUMEN

Este estudio muestra que los productos naturales de Carica respuesta fue la mortalidad del insecto. En los resultados se papaya pueden ser considerados como una alternativa válida registró que los extractos de semillas de los cultivares Mara- para el control de insectos nocivos en la agricultura. Se es- dol, Mamey y Amarilla de C. papaya, seguidos por los extrac- tudiaron las propiedades insecticidas de extractos de semillas tos de semillas de el cultivar Hawaiana, aplicados en concen- de cuatro cultivares de C. papaya (Maradol, Mamey, Amarilla traciones de 10, 100 y 1000ppm, resultaron tóxicos en las lar- y Hawaiana) incorporados a una dieta artificial para insectos. vas de S. frugiperda (50-70% de mortalidad corregida). Entre Se realizaron bioensayos con extractos hexánicos, acetónicos ellos, los extractos acetónicos de los cultivares Maradol y Ma- y metanólicos a concentraciones de 10, 100 y 1000ppm. To- mey a 10ppm fueron los más efectivos al causar un porcentaje das las pruebas se realizaron con larvas de primer estadio del de mortalidad de 73,6 y 62,8% de las larvas, respectivamente. gusano cogollero del maíz Spodoptera frugiperda. La variable

ATIVIDADE INSETICIDA DE EXTRATOS DE SEMENTES DE Carica papaya (L.) CONTRA A LAGARTA-DO- CARTUCHO Spodoptera frugiperda (J.E., Smith) (LEPIDÓPTERA NOCTUIDAE) Rodolfo Figueroa-Brito, Arturo Huerta-de la Peña, Ignacio Pérez Moreno, Vicente Santiago Marco Mancebón e Jesús Francisco López-Olguín RESUMO

Este estudo mostra que os produtos naturais de Carica papaya perda. A variável resposta foi a mortalidade do inseto. Nos resul- podem ser considerados como uma alternativa válida para o con- tados ficou registrado que os extratos de sementes dos cultivares trole de insetos nocivos na agricultura. Estudaram-se as proprie- Maradol, Mamey e Amarela de C. papaya, seguidos pelos extratos dades inseticidas de extratos de sementes de quatro cultivares de de sementes do cultivar Havai, aplicados em concentrações de 10, C. papaya (Maradol, Mamey, Amarelo e Havai) incorporados a 100 e 1.000ppm, resultaram tóxicos nas larvas de S. frugiperda uma dieta artificial para insetos. Realizaram-se bioensaios com (50-70% de mortalidade corrigida). Entre eles, os extratos acetô- extratos hexânicos, acetônicos e metanólicos em concentrações de nicos dos cultivares Maradol e Mamey a 10ppm foram os mais 10, 100 e 1000ppm. Todas as provas foram realizadas com larvas efetivos ao causar uma porcentagem de mortalidade de 73,6 e de primeiro estágio da lagarta militar do milho Spodoptera frugi- 62,8% das larvas, respectivamente.

It is therefore important to a greater respect for beneficial the Maradol, Yellow and Ha- tions in Yautepec, Morelos, foster the implementation of insect fauna (Bahena et al., waiian cultivars (Franco et al., Mexico, during July 2005. integrated pest management, 2003) due to their selectivity. 2006) in concentrations of 10, Using entomological brushes improving the processes of Figueroa-Brito (2002) evaluat- 15 and 20% were highly toxic and tweezers, 153 larvae at sampling and monitoring as a ed the effects of powders and caused 100% mortality different stages were collected basis for correct decision-mak- from different plants applied rates of larvae of S. frugiper- from the maize plants (~15 ing and promoting the rational as 15% of an artificial diet of da in less than 96h. Based on days of growth). The larvae integration of other alternative S. frugiperda. The author not- these antecedents, the aims of were then taken to the labora- tools, so as to reduce the ad- ed that the leaves of Pithecel- the present study were: a) to tory and fed individually on verse effects mentioned previ- lobium dulce Benth (Fabace- evaluate the toxic effect of an artificial diet (Burton and ously. One of such tools is the ae) and Crescentia alata hexanic, acetonic and metha- Perkins, 1987) and kept in use of products from vegetable H.B.K (Bignoniaceae) and the nolic extracts of seeds of the closed, cylindrical plastic con- sources. Plants contain a wide seeds of Jacaratia mexicana Mammee, Maradol, Yellow tainers, 3cm high by 3.5cm diversity of secondary metabo- A. DC. (Caricaceae) acted as and Hawaiian cultivars of C. diameter. The larvae under- lites, some of which exert an a deterrent, and that the papaya upon neonate larvae of went pupation in these same important role in their defense leaves of Prosopis juliflora S. frugiperda, and b) to com- vials and when they emerged against pathogens and herbi- (Sw.) DC. (Fabaceae) and the pare the relative toxicity of the as adults, they were placed in vores (Ware and Whitaker, seeds of Carica papaya (L.) different cultivars and the con- brown paper containers with a 2004). In various cases, it has (Caricaceae) and Bromelia centrations of the extracts. volume of 3 liters, containing been demonstrated that the use hemisphaerica Lam. (Brome- a 10cm-diameter plastic Petri of products of plant origin re- liaceae) proved toxic for first Materials and Methods dish with cotton wool damp- duces the application of syn- instar larvae of the pest. In ened in a 10% sugar solution thetic agrochemicals with a addition, powder from C. pa- Collection and rearing of to feed them. Mating and lay- resulting lower level of resis- paya cultivar Mammee seeds S. frugiperda ing of eggs took place in tance development among pest (Figueroa-Brito, 2002; these containers. For the tests, populations, thanks to their Figueroa-Brito et al., S. frugiperda larvae were the second generation of neo- differing modes of action and 2002a,b), as well as those of collected from maize planta- nate larvae were used.

OCT 2011, VOL. 36 Nº 10 753 Plant extracts Table I ways: 1) mortality means of Mortality (±SE) of first larval stage of each cultivar, in order to C. papaya fruits were ac- Spodoptera frugiperda fed with artificial diet know which of them, in gen- quired from different markets. CONtaining different concentrations of eral, was the most toxic; and The Yellow and Hawaiian hexanic, acetonic and methanolic extracts 2) by comparing all the ex- cultivars in the state of Oaxa- from seeds of four cultivars OF Carica papaya tracts, in order to know which ca, and the Mammee and Mortality ±SE (%) were more active. Raw data is Maradol cultivars in the state Extract presented and the mortality of Morelos. The seeds were 10ppm 100ppm 1000ppm data is corrected data. Statis- separated by cultivar and left tical analysis was performed Maradol Hexane 50.6 ±3.1 c 38.7 ±3.1 c 44.0 ±4.3 cd to dry in the shade for 15 Acetone 73.6 ±2.1 a 63.5 ±2.8 a 71.7 ±3.4 a using SigmaStat (2004). days. Once dry, the seeds Methanol 46.1 ±3.9 c 47.6 ±1.9 b 48.1 ±1.3 c were ground and sieved with Results Mammee Hexane 50.1 ±2.8 c 51.1 ±3.5 b 57.0 ±1.0 b an lka Wearke electric grinder Acetone 62.8 ±3.9 b 57.0 ±2.5 ab 48.7 ±1.6 c (model MF 10 Basic, GMBH Methanol 57.1 ±7.3 bc 51.2 ±5.0 b 52.6 ±4.2 bc In the absence of C. papa- & Co., Germany) using a ya, mortality in the case of 0.25mm mesh. From the pow- Yellow Hexane 50.3 ±2.9 c 59.7 ±1.4 a 56.1 ±1.1 b the controls with hexane and Acetone 54.0 ±1.2 c 62.3 ±1.2 a 58.9 ±2.2 b der obtained, 500g of each Methanol 54.7 ±1.6 c 42.3 ±1.5 c 33.1 ±1.9 e methanol was <10% and with cultivar were weighed out and the acetone control it was placed in 2 liter Erlenmeyer Hawaiian Hexane 20.6 ±3.0 e 49.7 ±2.2 b 42.2 ±6.3 cd 10.5% (Table I). At a concen- Acetone 38.5 ±3.9 d 49.5 ±2.1 b 43.3 ±5.1 cd flasks with 1.5 liters of hex- Methanol 25.3 ±2.6 e 24.1 ±1.4 d 36.4 ±6.7 de tration of 10ppm, the Maradol ane for a first maceration. and Mammee cultivars were Control Hexane 8.9 ±0.1 f 8.9 ±0.1 e 8.9 ±0.1 f The mixture obtained was Acetone 10.5 ±0.4 f 10.5 ±0.4 e 10.5 ±0.4 f the most active with an av- stirred lightly in the recipient Methanol 9.6 ±2.3 f 9.2 ±2.3 e 9.2 ±2.3 f erage larval mortality rate of and left in extraction for 72h 56.7% in both cases, show- Means in the same column followed by different letters are significantly at ambient temperature. After different (Tukey test, p<0.05). SE: Standard error. ing a significant difference this time, the mix was vacu- compared to the Yellow and um filtered using Whatman® Hawaiian cultivars, which Nº 5 filter paper. The same diet and 1ml of hexane, ace- random way in a breeding also showed significant dif- procedure was followed using tone or methanol. Diet ingre- chamber (Precision Incubator ferences (p<0.001) between either acetone or methanol as dients and the concentrated 818, model FFU20FC4CW0 themselves (Figure 1). Re- a solvent (Figueroa-Brito, extracts were mixed following 18, Electrolux Home Products, garding the extracts, the mean 2002). The hexane, acetone or the method suggested by USA) at 27 ±1ºC, 60 ±5% for mortality obtained with methanol, as applicable, were Franco et al. (2006). Of the relative humidity and photo- acetone (47.8 ±3.2%) was sig- removed from the solutions prepared mixture, 5ml were period of 12:12h. Mortality nificantly greater (p<0.001) obtained by reduced pressure placed in cylindrical plastic was recorded for the life span than the means in those with distillation using a rotavapor containers measuring 3cm of the larvae, which the test methanol and hexane, without (Büchi model R-114), so as to high by 3.5cm diameter. lasted, and was corrected ac- there being significant differ- obtain the hexanic, acetonic Once the diet-extract or di- cording to the Abbot method ences between the latter two and methanolic extracts of et-solvent gelled, a neonate (Abbot, 1925). The mortality (Figure 2). The three extracts each cultivar, and these were larva was placed in each con- data were transformed to arc- from Yellow and Mammee dried in a laminar flow cabi- tainer with the aid of a fine, sine before carrying out cultivars, and hexanic and net, to be used in the bioas- camel-hair brush. This pro- ANOVA at a significance lev- acetonic extracts from cultivar says. cess was repeated 3 times, el of 5% (α=0.05). The mor- Maradol, were the most toxic using 30 larvae for each rep- tality (%) means were com- ones, as they caused mortality Bioassay lication, and the containers pared using the Tukey test rates ≥50%. In particular, the were arranged in a totally (Steel et al., 1997), in two acetonic extract from cultivar For the preparation of 250g Maradol was the most active of artificial diet, the following one (p<0.001), with a cor- components proposed by Bur- rected mortality of 73.6 ±2.1% ton and Perkins (1987), were (Table I). used: beans (30g), wheat germ At a concentration of (13.75g), brewers’ yeast 100ppm, the Yellow and (8.75g), ascorbic acid (0.87g), Mammee cultivars were the sorbic acid (0.27g), methyl most toxic for the insect, with parahydrobenzoate (0.55g), no significant mean mortality formaldehyde at 10% (2.5ml), difference (Figure 1) noted water for beans (116ml) and between them (54.7 ±6.3 and water for agar (90ml). These 53.1 ±3.3%, respectively). Of ingredients were mixed with the extracts, those obtained the hexanic, acetonic or meth- with acetone were the most anolic extract of C. papaya active (p<0.005), leading to seeds of each cultivar to 48.5 ±2.1% mortality (Figure reach concentrations of 10, Figure 1. Effects of papaya cultivars on mortality of first larval stage of 2). Various extracts caused a 100 and 1000ppm. Controls Spodoptera frugiperda fed with artificial diet treated with different significant mortality rate of consisted on only the artificial concentrations of extracts from seeds. over 50% (Table I). The ace-

754 OCT 2011, VOL. 36 Nº 10 tonic extracts from cultivars Yucca periculosa Baker (Tor- Maradol and Yellow, and res et al., 2003), Iostephane hexanic from cultivar Yellow heterophylla (Cav.) Benth. ex extracts were the most active Hemsl. (Figueroa-Brito et al., (63.5, 62.3 and 59.7% mortali- 2006), acetonic extracts of P. ties, respectively) and with a dulce (Figueroa-Brito et al., statistically significant differ- 2007) and aqueous extracts of ence (p<0.001) with respect to Ricinus communis L. (Trujillo the rest of the extracts. and García, 2001) and Trichil- At the 1000ppm concentra- ia pallens C. DC. (Bogorni tion, the Maradol and Mam- and Vendramim, 2005). mee cultivars were the most Knowledge of biologically toxic, with no significant dif- active plant products which do ference between them, causing not cause damage to the envi- mortality rates of 54.4 ±4.6 Figure 2. Effects of different concentrations of hexanic, acetonic and ronment, with new biodegrad- and 52.8 ±5.6% respectively methanolic extracts from seeds of the Carica papaya on mortality of first able structures and which pre- (Figure 1). According to the larval stage of Spodoptera frugiperda fed with treated artificial diet. serve biodiversity, contributes extracts, the acetonic treat- to the development of less ments caused the highest damaging strategies than the (p<0.01) mortality in the insect of the four cultivars in the Mammee, are mixed with ac- use of chemical insecticides. (Figure 2), with 45.9 ±1.2%. In form of extracts in this study, etone, the most active compo- Such is the case of papain and the concentrations of 10, 100 lower mortality rates than nents are extracted. cystecine from the latex of C. and 1000ppm, various extracts those reported for seed powder It is worth pointing out that papaya against Spodoptera li- showed significant mortality were obtained. This can be the acetonic extracts of the tura (F.) (Konno et al., 2004), (>50%), among which the ac- explained by the fact that the Maradol and Mammee culti- or cystein protease inhibitors etonic extract from cultivar high toxic activity of the C. vars at 10ppm caused mortal- like papain on Coleoptera and Maradol was again the most papaya seeds in powder form ity rates of 73.6 and 62.8% Hemiptera insects, as well as toxic (p<0.01) for S. frugiper- may be due to a possible syn- respectively. These concentra- on phytopathogen nematodes da larvae with 73.6 ±2.1, 63.5 ergy, in which the active com- tions are significantly lower (Blanco-Labra and Aguirre, ±2.8 and 71.7 ±3.4% mortality, ponents act in a complemen- than those tried by Figueroa- 2002). Other research has respectively (Table I). tary way on S. frugiperda. Brito (2002), who found that shown a similar effect to that Despite these lower concen- the acetonic extract of seeds of the C. papaya seeds; such Discussion trations, in extract the seeds of the Mammee cultivar at is the case with the mixture of of C. papaya continue to ex- 5% led to a 100% mortality. epiperic phytogedunines or This study shows that vari- ercise high toxic effects on There are many studies of acetates of phytogedunines and ous extracts of the C. papaya larvae of S. frugiperda. toxicity carried out with other compounds: ligedunin from seed of the Maradol, Yellow Figueroa-Brito (2002) evalu- plants that reflect a similar Cedrela spp. (Céspedes et al., and Mammee cultivars, and to ated hexanic, acetonic or behavior on S. frugiperda (50- 2000), piplartin, 4´-desmetilpi- a lesser extent of the cultivar aqueous extracts of leaves, 100% mortality), such as those plartin and cenocladamin Hawaiian, are toxic to S. fru- seeds and flowers of C. pa- with macerations of Trichilia (Dyer et al., 2003) and piperin giperda larvae, achieving cor- paya cultivar Mammee in havanensis (Jacq.) by López- from Piper cenocladum C. rected mortality percentages fresh and powder form, at Olguín (1994), of Cabralea DC. (Batista-Pereira et al., of between 50.0 and 73.6% in concentrations of 5, 10, 15 canjerana (Vell.) and Cedrela 2006), and acetogenin from concentrations of 10, 100 and and 20%, applied to maize fissilis Vell. by Rodríguez and Annona cherimolia Mill. (Ál- 1000ppm. leaf discs on S. frugiperda Vendramim (1996), of Azadi- varez et al., 2007) on S. frugi- In previous studies with larvae and the results showed rachta indica (A. Juss.) and perda. These compounds do powdered C. papaya seeds, that the acetonic extracts of Cedrela odorata L. by Rodrí- not affect its predators and Figueroa-Brito (2002) reported the seeds (fresh and pow- guez and Vendramim (1997), parasitoids (Bahena et al., 100% larval mortality at con- dered) were the most active, and of Melia azedarach L. and 2003), as is the case with centrations of 10, 15 and 20% causing between 50 and 100% Trichilia pallida Sw. by Rodrí- Doru taeniatum (Dohrn) and with powder of the cultivar mortality in the insect at all guez and Vendramim (1998). Ectatomma ruidum Roger Mammee. The same happened the concentrations tested. In Many of the Meliaceae spe- (Schmutterer, 1990), and simi- with powder from the Marad- the current study, a similar cies continue to be active in larly in Chrysoperla carnea ol, Yellow and Hawaiian culti- effect was obtained with the the form of ethanolic extract, (Stephen) and Trichogramma vars which, at the same con- acetonic extracts of the Mara- as in the case of M. azedar- spp., with NeemAzal-T/S, PC centrations, led to mortality in dol Yellow and Mammee cul- ach (Mikolajczak et al., 1989) 05, Blank and PC Blank pro- all the larvae after 24h (Gar- tivars, which were the most and of aqueous extracts, as ducts (El-Wakeil et al., 2006; cía, 2004). Franco et al. active ones, causing corrected occurs with A. indica (Mikola- Aggarwal and Brar, 2006). (2006) studied mortality relat- mortality percentages of 50- jczak and Reed, 1987) and T. Considering the results ob- ed to time, noting that the 70%, the most effective of pallida (Bogorni and Vendra- tained in this study together powder of the four cultivars at them all proving to be the mim, 2005). In addition to with the experimental data al- 10 and 15% caused corrected acetonic extract of the Mara- these plants, there are other ready gathered in the literature, mortality rates in the insect of dol cultivar. These results plants whose extracts have it would be of interest to con- over 90% after 72 and 96h may be due to the fact that been reported as having insec- tinue with further studies with respectively. In evaluating the when the different cultivars ticide effects on S. frugiperda, the purpose of fractioning and insecticide effect of the seeds tested, especially Maradol and such as methanolic extracts of purifying the compounds found

OCT 2011, VOL. 36 Nº 10 755 in the seeds of the tested culti- nandes JB, Vieira PC, Da Silva ae) seeds. Int. J. Exp. Bot. 69: and survival of Spodoptera fru- vars of C. papaya, in order to MFGF, Corrêa AG (2006) In- 97-99. giperda (J. E. Smith). J. Chem. Ecol. 15: 121-128. study whether they continue to secticidal activity of synthetic Figueroa-Brito R, Calderón PJS, amides on Spodoptera frugi- Pérez-Amador MC, Muñoz V, Pingali PL, Pandey S (2001) World be effective against S. frugiperda. Z. Naturforsch. 61c: Hernández RMC, Valdés EME, maize needs meeting: techno- perda in the form of pure com- 196-202. Aldana LlL (2002b) Toxicity logical opportunities and priori- pounds or their mixtures, as Blanco-Labra A, Aguirre MC (2002) and growth inhibitory effects of ties for the public sector. En has already been shown with Proteínas involucradas en los extracts and some fractions Pingali PL (Ed.) CIMMYT the mixtures of compounds mecanismos de defensas de las from Carica papaya against 1999–2000 World Maize Facts plantas. Universidad de Guana- Spodoptera frugiperda (Lepi- and Trends. Meeting World from the seeds of the Maradol, juato. Acta Univ. 12: 3-28. doptera: Noctuidae). Rev. Lati- Maize Needs: Technological Op- Yellow and Hawaiian cultivars Bogorni PC, Vendramim JD (2005) noam. Quim. 30: 98-102. portunities and Priorities for the (Figueroa-Brito et al., 2002a,b). Sublethal effect of aqueous ex- Figueroa-Brito R, Aldana LlL, Public Sector. CIMMYT. Mexi- It is also necessary to carry out tracts of Trichilia spp. on Gutiérrez OM, Lotina HB, co, D.F. Part 1, p. 9. www.cim- evaluations of these mixtures Spodoptera frugiperda (J. E. King DB, Aguilar MI (2006) myt.org/Research/Economics/ map/facts_trends/maizeft9900/ and pure compounds in semi- Smith) (Lepidoptera: Noctuidae) Actividad bioinsecticida de ex- development on maize. Neotrop. tractos vegetales contra Spo html/maizeft9900_contents.htm. field and in field conditions to Entomol. 34: 311-317. doptera frugiperda (Lepidop- Rodríguez HC, Vendramim JD obtain basic knowledge in or- Burton LR, Perkins WD (1987) tera: Noctuidae). Entomol. Mex. (1996) Toxicidad de extractos der to establish their possible Rearing the corn earworm and 5: 677-681. acuosos de Meliaceae en incorporation into the integrat- fall armyworm for maize resis- Figueroa-Brito R, Valdés ME, Al- Spodoptera frugiperda (Lepi- ed management of the pest. tance studies. Proc. Int. Symp. dana LlL, Hernández RMC, doptera: Noctuidae). Man. Integ. on Methodologies for Develop- Gutiérrez OM (2007) Actividad Plagas 42: 14-22. ing Host Plant Resistance to insecticida de semillas de Rodríguez HC, Vendramim JD Conclusions Maize Insects. CIMMYT. Pithecellobium dulce. En (1997) Avaliação da bioatividade Mexico. pp 35-37. López-Olguín JF, Aragón GA, de extratos aquosos de Melia- Hexane, acetone and metha- Céspedes CL, Calderón JS, Lina L, Rodríguez HC, Vázquez GM ceae sobre Spodoptera frugi- nol extracts of seeds of the Aranda E (2000) Growth in- (Eds.) Agricultura sostenible: perda (J.E. Smith). Rev. Agric. hibitory effects on fall army- substancias naturales contra 72: 305-318. Maradol, Mammee and Yel- plaga. Benemérita Universidad worm Spodoptera frugiperda Rodríguez HC, Vendramim JD low cultivars of C. papaya, of some limonoids isolated Autónoma de Puebla / Colegio de Postgraduados / Universidad (1998) Uso de índices nutricio- followed by extracts from from Cedrela spp. (Meliaceae). nales para medir el efecto in- seeds of the Hawaiian culti- J. Agric. Food Chem. 48: de Guadalajara. Mexico. pp. 136-141. sectistático de extractos de var, applied at concentrations 1903-1908. Meliáceas sobre Spodoptera of 10, 100 and 1000ppm, Del Rincón CMC, Méndez LJ, Franco ASL, Jiménez PA, Luna LC, frugiperda. Man. 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