enhanced b c 13 7,8 incrusted with lignin. Arlete A Soares, cellulose microfibrils a According to Aoyama and Labinas, 9–12 Correspondence to: NSAgroindústria Dias-Pini, Tropical, Laboratório Rua60511-110, Brazil. de Doutora E-mail: [email protected] Entomologia, Sara Embrapa Mesquita 2270,Laboratório de Fortaleza, Entomologia, Embrapa Agroindústria CE, Tropical, Fortaleza, Brazil Laboratório de Morfologia e Anatomia Vegetal, Universidade Federal doFortaleza, Ceará, Brazil Laboratório de Entomologia, Embrapa Algodão, Campina Grande, Brazil Plant resistance to whiteflies has been associated with a number and Cherre S Bezerra Da Silva ∗ c a b genotypes expressing traitscould be valuable associated for breeding with and IPM programs. insect resistance of factors, including increased epidermalof thickness, type trichomes, and production size ofthe secondary metabolites, leaves. and color of epidermal thickness confers rigidity tonumbers leaves of owing overlapping to increased Sap-sucking insects need to insertinto their plant mandibular tissues stylets deep ina order thickened to leaf access epidermis the hinders vascular this bundles, and process by increasing Celli R Muniz, a , is an important pest of cashew in Brazil. The use of resistant plants 4,5 a* CW is 1 L.) and is ) linked to morphological Curtis (Hemiptera: Aleurodicus cocois ; leaf glandular trichomes; leaf phenolic compounds; leaf lamina; leaf surface; Aleyrodidae; Antixeno- Nivia S Dias-Pini, Anacardium occidentale a

Francisco C Vidal-Neto Aleurodicus cocois b

: 464–471 www.soci.org © 2019 Society of Chemical Industry 2,3 little is known about their resistance to CW 76 6 2020; Anacardium occidentale

Aleurodicus cocois Despite the agricultural losses caused by CW, there is a scarcity of cashew-specific pesticides registeredof in Agriculture the Brazilian to Ministry the control adoption this of pest. aninvolving Under integrated these resistant pest circumstances, management cashewThe (IPM) genotypes strategy advantages is of anpest such populations an appealing to approach levels option. positive include that do impacts the not reduction on causepest of the economic damage, environment, control compatibility methods, with cost-effectiveness, other and lasting results. Pest Manag Sci Despite the plethoraavailable of in cashew Brazil, clones that are commercially Aleyrodidae), is a sap-sucking insect thatcrops causes by direct puncturing damage to leaves duringby feeding, promoting and the proliferation indirect of damage sooty molds and viruses. 1The INTRODUCTION cashew whitefly (CW),

Keywords: and histochemical characteristics of leaves may be an effective strategy for the controllevels of of this resistance pest. to In CW. a Here, preliminary we assay, hypothesized weleaves that found such and that resistance dwarf-cashew their is clones content associated show of with different morphological phenolic characteristics compounds. of cashew RESULTS: We determined (i) theleaf attractiveness morphology and and suitability chemistry for of ovipositiongreenhouse those multiple-choice of assays, clones, PRO143/7 five and and dwarf-cashew CCP76 (iii) showed, respectively, clones the theand lowest towards eggs. relationship and Scanning CW, highest between electron (ii) counts leaf microscopy of (SEM) the both characteristics analysis CWand revealed and adults that lowest resistance PRO143/7 numbers and to of EMBRAPA51 CW. have, leaf respectively, In the glandularsurface highest trichomes. of We found cashew a leaves negativesulfate and correlation indicated between CW number a oviposition. of higher In trichomes accumulationDwarf-cashew in addition, of the clones confocal abaxial did phenolic microscopy not compounds analysis significantly inleaf and differ lamina the based histochemical and resistant on the tests the clone epidermal with layer. number PRO143/7 ferrous of relative leaf to epicuticular the striations,CONCLUSION: other and The the clones. resistance thickness of of dwarf-cashew both lation plants of to high CW levels of is phenolicslamina/epidermal associated layer in with are leaves. insignificant. an Additionally, the elevated contribution number© of 2019 of epicuticular Society trichomes striation of and density Chemical Industry accumu- and thickness of leaf sis; deterrence Abstract BACKGROUND: The cashew whitefly (CW), Elaine SS Goiana, James C Alves, ( Dwarf-cashew resistance to whitefly (wileyonlinelibrary.com) DOI 10.1002/ps.5531 Research Article Received: 12 February 2019 Revised: 14 June 2019 Accepted article published: 25 June 2019 Published online in Wiley Online Library: 27 July 2019 and respective mechanisms of resistance. Identification of cashew a major pest ofwidespread dwarf-cashew in all ( growing areas of northeastern Brazil,the occurring in form of episodic but intensive outbreakslosses that in cause productivity. significant

464 465 )and 1 − m). μ areas of abaxial 2 m), and the thicknesses of the ) in 95% ethanol], which binds 1 μ − wileyonlinelibrary.com/journal/ps aminoethyl diphenylborinate (10 g L = 0.0207. Boxes with different letters significantly - P Treated and non-treated samples were mounted = 3.70, 21 4,20 , in greenhouse, multiple-choice tests. (A) Number of CW adults counted. Seedlings were used asthis an assay exception because to it required adultwould plant not plants be confinement in feasible in with cages, adult plants.plants which However, were adult used 3-year-old in the next three microscopy experiments. 2.3 Scanning electronMorphological microscopy analyses weremicroscopy carried (SEM) using out a Tescan by (SãoVega Bernardo scanning do 3 Campo, electron Brazil) microscope underexperimental an acceleration design voltage wasreplicates of each 15 completely involving kV. five The leaf randomizeddifferent fragments individuals removed of with from each dwarf-cashew three three clone.collected The from leaves 3-year-old were adult plants inof the Embrapa Agroindústria experimental Tropical station (Pacajus, CE, Brazil). Plantrial mate- was treated withtimes Karnovsky (10 min fixative each) forosmium with tetroxide 48 for h, 1 phosphate h, washed washed buffer, intreatment post-fixed three distilled with water, with dehydrated ethanol by solutions 1% of increasing30, concentration 40, (20, 50, 60, 70, 80,times 90%) (15 (15 min min each) each) andto finally in washed critical 100% three point ethanol. drying,platinum, Samples and mounted examined were by on SEM. submitted stubs, For eachthe sputter sample, number we coated determined of with glandular trichomes in 1.05 mm epidermal layers on the abaxial and adaxial surfaces ( 2.4 Confocal microscopy Histochemical analyses of leavesclones were performed of in order the to detect selectedIn the presence dwarf-cashew the of phenols. first assay,the six leaf clones fragments and were submittedtioning. removed Samples to from were each paradermic separated of andwas into two retained longitudinal groups, as sec- one non-treatedwith of Neu’s control reagent which [2 and the other was treated surface, the number ofthe epicuticular thickness striations of between the stomata, leaf lamina ( to specific chemicalflavonoids. groups and enhances the fluorescence of polyethylene glycol 4000 (50 g L = 0.0262. Eggs: F A. P Aleurodicus cocois = 3.47, 4,20 20 while the release of = 0.05). 𝛼 15–17 = 5. Adults: F N 18,19 were selected. on 25 dwarf-cashew genotypes A. cocois A. cocois : 464–471 © 2019 Society of Chemical Industry 76 Moreover, an increase in the number of tector 14 2020; Resistance of dwarf-cashew clones to cashew whitefly (CW), . This selection was based on preliminary experiments where Here, we hypothesized that the resistance of cashew plants differ from each other according to Tukey’s test ( to CW is associatedand with morphological their characteristics content ofthe leaves of relative phenolic attractiveness compounds. anddwarf-cashew We suitability clones for determined towards oviposition CW, (i) chemistry (ii) of of those the five clones, leaf andcharacteristics morphology and (iii) resistance and the to CW. relationship between leaf Pest Manag Sci Figure 1. on leaves. (B) Number of CW eggs on leaves. was assessed in thesusceptible field. or The resistant clones to that stood out as the most 2.2 Attraction andThe oviposition assays objective of thischaracteristics multiple-choice with assay attraction was and oviposition tofemales. preferences correlate of The leaf CW experimentalwith design five was replicates, completely eachfeaturing randomized mature involving leaves one (120 days dwarf-cashewstudied old) selected seedling clones. from The each of1.0 plants m the were cube screen distributed cageto randomly and prevent contact spaced inside between at the leaves. least a Onewhiteflies 15 hundred cm adult (20 apart female insects in per order plant)the were attraction released of into thecounting, insects cage with to the and aid plants ofon was a the mirror, evaluated the abaxial after number surfaces ofremoved 24 of insects h in the present by tubes. leaves. The Subsequently, insects the were eggs carefully laid on leaves were the infestation rates of cocois 22.1 MATERIAL AND METHODS Plant material The dwarf-cashew clonesand CCP76, PRO143/7, originating BRS226, from EMBRAPA51, the germplasm BRS274 Agroindústria bank of Tropical Embrapa (Fortaleza,investigation CE, of the Brazil), characteristics associated with were resistance to selected for Mechanisms of dwarf-cashew resistance to whiteflyenergy demand. www.soci.org trichomes on the leaflimiting the mobility surface of nymphs and may larvae, act as a physical barrier by secondary compounds from glandular trichomes maypreference, oviposition, affect and food survival. et al. = 0.0089. : 464–471 P 76 magnification). × = 6.21, 2020; 4,10 Pest Manag Sci = 0.0001. Striations: F P magnification). (B) and (F) Epicuticular striations × = 18.11, 4,10 = 0.05). 𝛼 magnification). Micrographs: C.R. Muniz. × samples weremounted washed onto withmicroscopy slides distilled using under a water,B digital Leica water cross-sectioned, microscope (Wetzlar, with and lightcoupled Germany) emitting with a diode examined model digital (LED) camera. illumination DM4000 by light 2.6 Statistical analysis After confirmation ofPearson, and normality Kolmogorov–Sminorv’s (Shapiro–Wilk,ticity tests) (Barllet’s D’Agostino and and & homoscedas- Brown–Forsythe’s tests), one-way ANOVA www.soci.org ESS Goiana © 2019 Society of Chemical Industry SEM) of leaves of five dwarf-cashew clones with different levels of resistance to the cashew whitefly, ) for 48 h to determine ± 1 objective, an ultraviolet − × Treated and non-treated (control) 22 . (A) and (E) Density of glandular trichomes on the abaxial leaf surface (267 magnification). (C) and (G) Cross-section of the leaf lamina with leaf thickness indicated by yellow line (1460 × = 3 plants, each donating five leaf fragments for analysis. Trichomes: F N , from scanning electron micrographs. (A) Number of glandular trichomes on the abaxial leaf surface. (B) Number of epicuticular striations Morphological characteristics (mean Scanning electron micrographs of leaves of dwarf-cashew clones that are susceptible (A–D, CCP76) and resistant (E–H, PRO143/7) to the cashew Aleurodicus cocois Columns with different letters significantly differ from each other according to Tukey’s test ( the presence of total phenols. 2.5 Optical microscopy In the second assay, leaffixed fragments in from ferrous each sulfate of solution the (0,002 g clones L were filter (450–490 nm) andviewing. a barrier filter (520 nm) for fluorescence wileyonlinelibrary.com/journal/ps on semi-permanent slidesa and Zeiss examined/photographed (Oberkochen, using scanning Germany) microscope (CLSM) model with a LSM710 40 confocal laser Figure 3. whitefly, between stomata (2000 (D) and (H) Abaxial epidermis with epidermal thickness indicated by yellow line (1930 Figure 2. Aleurodicuscocois between stomata.

466 467 = 0.0089). Striations P = 0.0001). PRO143/7 and P = 6.21, = 0.0207) on leaves. PRO143/7 = 0.0257. Abaxial epidermal layer: P P 4,10 wileyonlinelibrary.com/journal/ps = 18.11, = 4.43, = 3.70, 4,10 4,10 4,20 = 0.05). 𝛼 )( r = 0.0262) and eggs (F P were highest and lowest, respectively, in EMBRAPA51 and BRS226. had significantly fewer adults and eggsicant than CCP76, differences but were no found signif- betweenand either of all these two the clones and eggs. other tested clones (Fig. 1) regarding3.2 both adults Scanning electronDensities microscopy ofdwarf-cashew leaf clones trichome (F were strongly different among EMBRAPA51 showed the highest and lowest trichomeall densities tested of clones, respectively. However, nowere significant found among differences EMBRAPA51, CCP76, and BRS226 (Figs3(A,E)). 2(A) The and number of epicuticular striations between stomata was affected by dwarf-cashew clone (F ) r = 3.47, 4,20 = 0.1843. Columns with different letters significantly differ from each other according to P = 2.02, 4,8 SEM) of leaves of five dwarf-cashew clones with different levels of resistance to the cashew whitefly, ± = 3 plants, each donating five leaf fragments for analysis. Leaf lamina: F . (A) CW adults. (B) CW eggs. Pearson’s correlation coefficient ( N = 0.05) was applied to test the effects 𝛼 : 464–471 © 2019 Society of Chemical Industry 76 , from scanning electron micrographs. (A) Leaf thickness across the leaf lamina. (B) Thickness of the abaxial epidermal layer. (C) Thickness 2020; Aleurodicus cocois = 0.05). NS = non-significant difference among clones. = 0.1982. Adaxial epidermal layer: F 𝛼 P Morphological characteristics (mean Correlation between number of glandular trichomes on the abaxial leaf surface of dwarf-cashew plants and number of individuals of cashew = 1.93, = 0.05). All analyses were performed using GraphPad Prism 4,8 𝛼 version 8.0.2 for Mac OS X (GraphPad Software, La Jolla, CA, USA). F Tukey’s test ( ( Pest Manag Sci 3RESULTS 3.1 Attraction andWe oviposition observed assays significant differences among the fiveclones dwarf-cashew tested regarding both number of CW adults (F of dwarf-cashew clone onthe the number number of ofthe leaf CW thickness adults trichomes of and andthe eggs, leaf adaxial epicuticular lamina, epidermal striations, the layer.tions and Additionally, between abaxial we numbers epidermal of testedtrichome CW layer, for adults density and correla- and using eggs versus the the Pearson’s plant correlation coefficient ( whitefly (CW), followed by Tukey’s test ( Figure 5. Figure 4. of the adaxial epidermal layer. Aleurodicus cocois Mechanisms of dwarf-cashew resistance to whitefly www.soci.org et al. : 464–471 76 2020; Pest Manag Sci Confocal laser scanning micrographs of the longitudinal sections Figure 6. of leaves ofresponding dwarf-cashew non-treated clones controls. Clone treatedtreated; BRS226, with BRS274, (A) (C) Neu’s non-treated non-treated reagentand and and (F) (B) and (D) treated; cor- treated; EMBRAPA51,(I) CCP76, (G) non-treated (E) non-treated and non-treated and (J) (H)clones treated. treated; and The PRO143/7, with greenish higher fluorescence intensitygraphs: observed in C.R. PRO143/7 in Muniz. is all typical of phenols. Micro- B. 12,23–25 0.6198; www.soci.org ESS Goiana − = 0.1887, 2 = r r © 2019 Society of Chemical Industry 0.4344; − = r = 0.1982) or in the adaxial P thus a high density of glandular 1.428, intercept = 64.85) (Fig. 5). 6 − = 1.93, Our morphological and histochemical 4,8 reported that varieties of eggplant that 20 = 0.0257). BRS274 and EMBRAPA51 showed 11 P et al. = 0.1843) surface (Figs 3(D,H) and 4(B,C)). There = 0.0137, slope = = 4.43, P P 4,10 . The abaxial surfaces of cashew leaves are preferred for = 2.02, = 0.3841, The presence of glandular trichomes in plants decreases attack 4,8 = 0.1056). However, we observed a negative correlation between 2 trichomes in this areaOur represent data on a oviposition physical andthis barrier glandular hypothesis as trichome for it density the shows support thatdecreases pest. the as number trichome of density CW increases. eggs on leaves wileyonlinelibrary.com/journal/ps Hasanuzzaman 4 DISCUSSION With the attraction and oviposition assays,the we dwarf-cashew demonstrated that clone PRO143/7CCP76 is in resistant semi-field to conditions, CWin as relative it a to had preliminary been assay. observed in field However, we found no significantthese differences two between clones either and3(B,F)). of all In addition, the we otherleaves observed tested that of clones the the (Figs vascular 2(B)lignified system resistant and cell in PRO143/7 walls the compared comprised tocant broad effect the of susceptible and dwarf-cashew CCP76. clone highly Alamina was signifi- (F found on thickness of leaf (F P the number of eggs and trichome density (Pearson analysis revealed that the leaves ofhigher such a glandular resistant clone trichome featured density,phytochemicals as (aromatic well amino as acids, highercompounds) flavonoids, intensity relative and of to phenolic the othercharacteristics clones, are showing associated that with these resistance of leaf PRO143/7 to CW. by herbivores and indicates resistance to whiteflies. was no significanton correlation plants and between trichome density the (Pearson number of adults contained highaffected densities the landing, of feeding, and glandular oviposition trichomes preferences of negatively tabaci feeding and oviposition by CW, the highest and lowest thickness, respectively. However, weno found significant differences among eitherall of the these other two tested clonesclone clones did and not (Figs significantly 3(C,G) affect andeither the 4(A)). thickness in of Dwarf-cashew epidermal the layer, abaxial (F 3.4 Optical microscopy Microscopic examination of cross-sections ofhad leaf been fragments that treatedof with phenols ferrous in sulfate theas confirmed epidermal demonstrated the and by mesophyll presence the(Fig. layers black–brown 7). of areas The all in clones, reactionPRO143/7 the (Fig. 7(J)) micrographs was in comparison with the more other clones,CCP76 especially (Fig. intense 7(F)). in leaf samples from 3.3 Confocal microscopy Longitudinal sections of leaf fragments that had beenNeu’s treated reagent with exhibited a greenish fluorescence,epidermal especially cells in (Fig. the 6), when examinednation, by while CLSM under non-treated UV samples illumi- showedAlthough no we such observed fluorescence fluorescence. in the samples ofintensity all was clones, particularly its strong in PRO143/7 (Fig.from 6(J)). In CCP76 samples and EMBRAPA51,(Fig. the 6(F,H)). fluorescence was less intense r

468 469 ,(G) Soares wileyonlinelibrary.com/journal/ps : 464–471 © 2019 Society of Chemical Industry 76 2020; Optical micrographs of cross-sections of leaves of dwarf-cashew clones treated with ferrous sulfate solution and corresponding non-treated Pest Manag Sci Figure 7. controls. Clone BRS226, (A) non-treated andnon-treated and (B) (H) treated; treated; BRS274, PRO143/7, (C) (I) non-treated non-treatedand and and C.R. (J) (D) Muniz. treated. treated; The CCP76, black–brown (E) color non-treated indicates and the (F) presence of treated; phenols. EMBRAPA51 Micrographs: A.A. Mechanisms of dwarf-cashew resistance to whitefly www.soci.org et al. : 464–471 76 2020; Sistema de produção :309–328 (2012). Pest Manag Sci Pragas do cajueiro, Agronegócio caju: 57 , 2nd edn. Embrapa Agroindústria Tropical, systematics: how many species are there? . Embrapa, Brasília, pp. 195–215 (2013). :176–186 (2012). In a breeding program that aims at incorporat- 11 Annu Rev Entomol 46 Bemisia tabaci whitefly JIntegrAgric práticas e inovações do caju: Pragas do cajueiro Fortaleza (2016). arthropods. The present study focused on the role of physical and chemi- 1 Liu S, Colvin J and De Barro PJ, Species concepts as applied to the 2 Mesquita A and Braga Sobrinho R, 3 Mesquita ALM, Dias-Pini NS and Braga Sobrinho R, 4 Smith CM and Clement SL, Molecular bases of plant resistance to carried out in our study revealed that thehere five dwarf-cashew tested clones differ incompounds, their with contents the of amino resistantcontents acids clone relative and PRO143/7 to phenolic showing the susceptible higher worth CCP76 noting and that EMBRAPA51. EMBRAPA51 and Itby CCP76 is are farmers extensively in grown northeasternattacked Brazil, by where CW. the latter is intensively ing mechanisms of resistance todiseases, PRO143/7 biotic arises factors as a such source as ofthat pests interesting will characteristics and contribute to theof development dwarf-cashew. of CW-resistant varieties cal barriers in the previouslyCW. reported First, resistance we of confirmed PRO143/7ond, to that we demonstrated PRO143/7 that the ischomes presence resistant and of epicuticular to large striations, numbers as CW. of wellhigh Sec- as tri- levels the of accumulation of phenolics inCW the landing leaves, are and factors oviposition, thatof consequently discourage PRO143/7 conferring to resistance CW. Third,ina we and showed thickness that thickness of of epidermalto such leaf layer a lam- of resistance. leaves Hence, dochomes the not and presence epicuticular of contribute striations, large as numbers wellhigh of as levels of tri- the phenolics accumulation in the of leaves, should betance considered as factors resis- in future breeding programstionally, of dwarf-cashew. the Addi- resistance ofmost PRO143/7 to appropriate CW candidate makes forgenetic this further improvement clone investigation program the within at our cal, Embrapa especially Agroindústria if Tropi- wePRO143/7 consider produces that better besides qualitytrichome/phenolic-based the cashew resistance resistance nuts to approach than CW, investigated could CCP in PRO143/7 be 76. with the A further aim ofthus creating reducing healthier pesticide plants usage, although thethis commercialization genotype of still depends on the improvement of itsand agronomical industrial attributes. REFERENCES ACKNOWLEDGEMENTS We would like toTechnological thank Development the (CNPq) Brazilian forfellowship awarded to Council a NSDP,and the for Brazilian research Coordination Scientific forImprovement productivity the of and Higher Education Personnel (CAPES) for aresearch Masters scholarship to ESSG. AUTHOR CONTRIBUTION STATEMENT ESSG, CRM, and NDSDP conceivedCRM, and JCA, designed and research. ESSG, AAScontributed with reagents conducted and/or analytical experiments. tools. CSBDS NDSDP analyzed data. and ESSG, FCVN CRM, NDSDP,authors and read and CSBDS approved the wrote manuscript. the manuscript. All ), Pre- Betula www.soci.org ESS Goiana towards B. tabaci 26,27 44 ) In our study, the glandu- 11 © 2019 Society of Chemical Industry 30 In our study, confocal Vigna radiata Cuticle and cell wall thick- 35,36 and eggplant. 13 39 ), species release exudates contain- Populations of the whitefly 37 . In the young shoots of birch ( may be as important in birch resistance However, in agreement with the morpho- and mung beans ( 45 Indeed, the presence of high concentrations Vignamungo 31,32 40 41–43 Lycopersicon It is possible that phenolics in the glandular tri- According to Simmons and Gurr, Thrips tabaci 33,34 cotton, B. pubescens 10,28,29 12 black gram ( . Within this class of natural compounds, the condensed 38 Another possible mechanism of resistance afforded by glan- Epidermal thickness can be augmented by the deposition of Phenolic compounds are the most common group of plant logical analysis previously discussed, the fluorescence technique dular trichomes istures chemical can protection excrete since lipids, these terpenoids, leaf and struc- polyphenols. wileyonlinelibrary.com/journal/ps and optical microscopy analysis showedlates that higher PRO143/7 accumu- amounts ofother phenolic aromatic compounds amino relative acids,which is to flavonoids, consistent with CCP76 the and higher and densityobserved of in EMBRAPA51, glandular the trichomes resistant clone (PRO143/7). However, wea observed lack ofchome correlation density, between while dwarf-cashew number cloneof of affected females both and CW number of trichomes, females asthat previously and trichome-related discussed. chemicals This tri- implies mostrole likely in the play repellent potential an of dwarf-cashew insignificant clones toand CW suggests adults, that the negative impact of trichomes ontion CW is oviposi- based on deterrence by chemical and physical mechanisms. lignin, cellulose, andmented hemicellulose, epidermal hence layers are leaf farresistant cells more to the rigid with attack and, of aug- therefore, herbivores. more nesses, as well as epicuticular wax deposition, acters by as hindering physical movement, barri- feeding, and oviposition of herbivores that feed directly from thestructural epidermal characteristics cells. were In associated with onion resistance plants,ent of these cultivars differ- to phenolics from the glandularnisms trichomes and have act important asinsect roles defense pests. in mecha- the plant’s resistance against lar trichomes of ing ketones, suchact as to 2-tridecanone deter and orstore repel 2-undecanone, volatile arthropod which terpenoids pests.a-terpinene, Glandular such and trichomes a-phellandrene, as which can tance 7-epizingiberene, confer antixenotic to p-cymene, resis- whiteflies. vious reports have showndular trichome negative density correlations and betweenwhiteflies. attraction/oviposition glan- preference of B. tabaci tannins (polyphenols) have significantties since insect they resistance bind to proper- proteins,growth inhibiting of digestion and the delaying herbivores.rich sources In of tannins. this regard, cashew plants are very of phenolics has been negatively correlatedeggplant, with the resistance of chomes of the five dwarf-cashew clonesboth tested abaxial showed and similar adaxial thickness epidermal of ferent layers levels despite of displaying dif- resistanceoviposition to trials. In CW addition, basedon the on thickness impact of both leaf of lamina attraction dwarf-cashew andfered epicuticular and clone from striations the strongly clone dif- impact onon number leaves. of adults Therefore, and the eggs thicknessesdermal found of layer, as leaf well lamina assignificantly and to epicuticular the the striations, resistance epi- of do dwarf-cashew not clones contribute to CW. defense secondary metabolites andphenolic include acids, simple hydroxycinnamic phenols acid derivatives, and andall of flavonoids, which play an importantherbivorous role insects. in host plant resistance against have been positively correlated with the thickness ofcotton, leaf lamina in against herbivores as foliar phenolics.

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