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Fall armyworm herbivory affects silica accumulation in corn and rice Flor E. Acevedo 1 , Michelle Peiffer 1 , Dawn Luthe 2 and Gary Felton 1 1 Department of Entomology, and 2 Department of Plant Science. The Pennsylvania State University, University Park, PA 16802. Silica accumulation is an inducible plant defense mechanism that causes wear on the insect mandibles and hampers their ability to feed on plant tissues. Insect herbivores are able to modify its development to feed on plants with high silica content. This research determined: 1) The effect of fall armyworm herbivory on the accumulation of silica in corn and rice plants. 2) The effect of silica on the fall armyworm mandible wear. 3) The fall armyworm adaptations to feed on plants with high silica content. Plants and insects We used the corn, Zea mays inbred line B73; and the rice, Oryza sativa japonica cultivar Nipponbare. The fall armyworm, Spodoptera frugiperda were obtained from a lab colony from the USDA-ARS in Gainesville Florida. Effect of insect feeding on plant silica accumulation Rice and corn plants were exposed to FAW herbivory for 24 hours or mechanically damaged with scissors in a complete randomized design; the new regrowth tissues were used to quantify the amount of silica with the molybdenum yellow method (3). Caterpillar mandible wear Mandible wear and mineral composition was determined by X-ray and scanning electron microscopy. Statistical analysis We used ANOVA to test for differences of treatment means, followed by the Tukey test at α=0.05. 2. Silica accumulated in plant leaves wears the fall armyworm mandibles. This is one of the \irst studies to explore the dynamic interplay between insect herbivores and silica accumulation in plants of economic importance. Three of the most important crops for human kind are grasses (silica-accumulating plants); a better understanding on how silica enhances crop resistance against insect herbivores is of global importance. Plants, especially grasses, accumulate large amounts of silicon dioxide in their leaves. Silica deposition plays a role in defense against abiotic and biotic stresses including herbivores. This research determined the effect of the fall armyworm (FAW) herbivory in the accumulation of silica in corn and rice plants, as well as the effect of silica on the caterpillar mandible durability. Our results show that FAW feeding induces silica accumulation in rice but not in corn plants. Silica wears the FAW mandibles affecting its feeding ability; in response the FAW plastically modi\ies its morphology and development to feed on silica- containing plants. 1. Hunt JW, Dean AP, Webster RE, Johnson JN, and Ennos AR. Annals of Botany 102: 653-656, 2008 2. Fürstenberg-Hägg J, Zagrobelny M, and Bak S. Int. J. Mol. Sci. 14, 10242-10297, 2013 3. Saito K, Yamamoto A, Sa T, and Saigusa M. Soil Sc. Plant Nutr., 51, 29-36, 2005 Fig. 1. Fall armyworm larva feeding on corn. Silicon accumulation increases plant resistance to herbivorous insects Among many other roles, silicon accumulation increases the abrasiveness of the plant leaves acting as a physical defense mechanism against phytophagous insects (1). Herbivorous insects face diverse plant defense mechanisms Insect herbivores adapt their morphology, physiology, development and behavior to counteract plant defenses (2). Research questions 1) Does insect feeding affect plant silica accumulation? 2) Does silica affects the herbivore feeding? 3) How do insects manage to feed on silica- containing plants? Model system Corn, rice and the polyphagous insect fall armyworm, Spodoptera frugiperda (Lepidoptera: Noctuidae) (Fig. 1). Results and discussion Rice and corn plants accumulate silica in the epidermis of their leaves and tips of trichomes (Fig. 2). Rice contain ~3 times more silica than corn. 3. The fall armyworm plastically modiOies its morphology and development to feed on tough silica-containing plants. 1. Fall armyworm feeding affects plant silica accumulation. Corn Rice Fig. 2. Backscatter electron image of silica bodies in rice and corn leaves. Fig. 3. Silica accumulation in rice (left) and corn (right) plants after mechanical damage and insect-feeding treatments. Different letters indicate signi\icant differences among treatment means (P<0.05). ArtiOicial diet-fed Rice-fed Corn-fed Fig. 5. Backscatter electron image of the fall armyworm mandibles from caterpillars fed on different diets. Whiter outer areas correspond to mineral-enriched regions. Zn Cl Na Fig. 4. X-ray mapping of Zn, Cl and Na of the FAW mandibles. Fig. 6. Head capsule size (left), mandibular muscles (center), and mandible size (right) of the fall armyworm last instar caterpillars reared on rice and corn leaves. Corn-fed Rice-fed 2.60 mm Rice-fed Corn-fed 2.92 mm Rice-fed Corn-fed When feeding on rice, the FAW undergoes an additional molt increasing the size of its mandibles and attached musculature (Fig. 6). Methods Rice Corn The FAW enhances the hardness of the cutting edges of its mandibles by incorporating minerals including zinc, chlorine and sodium (Fig. 4). Feeding on rice, a high silica- containing plant, destroys most of the mineralized area of the FAW mandibles (Fig. 5). Mandible wear hampers the caterpillar ability to feed on plant tissues. Conversely, in corn, the deposition of silica is the same for insect-fed plants and for untreated controls, both are lower than mechanically damaged plants (Fig. 3). The mechanical damage caused by the fall armyworm feeding induces silica accumulation in rice plants by ~37.2 % compared with untreated controls. USDA-ARS Gainesville, Fl. Penn State Microscopy facility. Penn State College of Ag. Sciences (CAS) grant. USDA-NIFA grant # 2011-67013-30352. NSF SigniOicance Conclusions References Acknowledgments Objectives Introduction Abstract

Fall armyworm herbivory affects silica accumulation in ... · Fall armyworm herbivory affects silica accumulation in corn and rice Flor E. Acevedo1, Michelle Peiffer1, Dawn Luthe2

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Page 1: Fall armyworm herbivory affects silica accumulation in ... · Fall armyworm herbivory affects silica accumulation in corn and rice Flor E. Acevedo1, Michelle Peiffer1, Dawn Luthe2

Fallarmywormherbivoryaffectssilicaaccumulationincornandrice

FlorE.Acevedo1,MichellePeiffer1,DawnLuthe2andGaryFelton11DepartmentofEntomology,and2DepartmentofPlantScience.ThePennsylvaniaStateUniversity,UniversityPark,PA16802.

•  Silica accumulation is an inducible plantdefense mechanism that causes wear on theinsect mandibles and hampers their ability tofeedonplanttissues.

•  Insect herbivores are able to modify itsdevelopment to feedonplantswithhigh silicacontent.

Thisresearchdetermined:1)  The effect of fall armyworm herbivory on the

accumulationofsilicaincornandriceplants.2)  The effect of silica on the fall armyworm

mandiblewear.3)  The fall armyworm adaptations to feed on

plantswithhighsilicacontent.

•  PlantsandinsectsWeusedthecorn,Zeamays inbredlineB73;andthe rice, Oryza sativa japonica cultivarNipponbare. The fall armyworm, Spodopterafrugiperda wereobtainedfromalabcolonyfromtheUSDA-ARSinGainesvilleFlorida.•  Effectofinsectfeedingonplantsilicaaccumulation

Rice and corn plants were exposed to FAWherbivory for 24hours ormechanically damagedwith scissors in a complete randomized design;the new regrowth tissues were used to quantifytheamountofsilicawiththemolybdenumyellowmethod(3).•  CaterpillarmandiblewearMandible wear and mineral composition wasdetermined by X-ray and scanning electronmicroscopy.•  StatisticalanalysisWe used ANOVA to test for differences oftreatment means, followed by the Tukey test atα=0.05.

2. Silica accumulated in plant leaves wears the fall armywormmandibles. This is one of the \irst studies to explore the

dynamic interplaybetween insectherbivoresandsilica accumulation in plants of economicimportance.Threeofthemostimportantcropsforhumankindare grasses (silica-accumulating plants); a betterunderstanding on how silica enhances cropresistance against insect herbivores is of globalimportance.

Plants, especially grasses, accumulate largeamounts of silicon dioxide in their leaves. Silicadeposition plays a role in defense against abioticand biotic stresses including herbivores. Thisresearch determined the effect of the fallarmyworm(FAW)herbivoryintheaccumulationofsilicaincornandriceplants,aswellastheeffectofsilica on the caterpillar mandible durability. Ourresults show that FAW feeding induces silicaaccumulation in rice but not in corn plants. Silicawears the FAW mandibles affecting its feedingability;inresponsetheFAWplasticallymodi\iesitsmorphology and development to feed on silica-containingplants.

1.  HuntJW,DeanAP,WebsterRE, JohnsonJN,andEnnosAR.AnnalsofBotany102:653-656,2008

2.Fürstenberg-HäggJ,ZagrobelnyM,andBakS.Int.J.Mol.Sci.14,10242-10297,2013

3.SaitoK,YamamotoA,SaT,andSaigusaM.SoilSc.PlantNutr.,51,29-36,2005

Fig.1.Fallarmywormlarvafeedingoncorn.

•  Silicon accumulation increases plantresistancetoherbivorousinsectsAmong many other roles, silicon accumulationincreases the abrasiveness of the plant leavesacting as a physical defensemechanism againstphytophagousinsects(1).

• Herbivorous insects face diverse plantdefensemechanismsInsect herbivores adapt their morphology,physiology, development and behavior tocounteractplantdefenses(2).• Researchquestions1)Doesinsectfeedingaffectplantsilica

accumulation?2)Doessilicaaffectstheherbivorefeeding?3) How do insects manage to feed on silica-

containingplants?• Modelsystem Corn,riceandthepolyphagousinsectfallarmyworm,Spodopterafrugiperda(Lepidoptera:Noctuidae)(Fig.1).

Resultsanddiscussion

Rice and corn plantsaccumulate silica intheepidermisoftheirleaves and tips oftrichomes (Fig. 2).Ricecontain~3timesmoresilicathancorn.

3. The fall armyworm plastically modiOies its morphology anddevelopmenttofeedontoughsilica-containingplants.

1.Fallarmywormfeedingaffectsplantsilicaaccumulation.CornRice

Fig.2.Backscatterelectronimageofsilicabodiesinriceandcornleaves.

Fig. 3. Silica accumulation in rice (left) and corn (right) plants aftermechanical damage and insect-feeding treatments. Different lettersindicatesigni\icantdifferencesamongtreatmentmeans(P<0.05).

ArtiOicialdiet-fedRice-fedCorn-fed

Fig.5.Backscatterelectronimageofthefallarmywormmandiblesfromcaterpillarsfedondifferentdiets.Whiterouterareascorrespondtomineral-enrichedregions.

Zn Cl Na

Fig.4.X-raymappingofZn,ClandNaoftheFAWmandibles.

Fig.6.Headcapsulesize(left),mandibularmuscles(center),andmandiblesize(right)ofthefallarmywormlastinstarcaterpillarsrearedonriceandcornleaves.

Corn-fedRice-fed

2.60mm

Rice-fedCorn-fed2.92mm

Rice-fedCorn-fed

Whenfeedingonrice,theFAWundergoesanadditionalmoltincreasingthesizeofitsmandiblesandattachedmusculature(Fig.6).

Methods

Rice Corn

The FAW enhances thehardness of the cuttingedges of its mandibles byincorporating mineralsincludingzinc, chlorineandsodium(Fig.4).

Feedingonrice,ahighsilica-containing plant, destroysmostofthemineralizedareaof the FAW mandibles (Fig.5). Mandible wear hampersthecaterpillarabilitytofeedonplanttissues.

Conversely, inc o r n , t h edeposition ofsilica is thes a m e f o ri n s e c t - f e dplants and foru n t r e a t e dcontrols, botharelowerthanmechanicallyd a m a g e dplants(Fig.3).

The mechanical damage caused by the fall armyworm feeding induces silicaaccumulationinriceplantsby~37.2%comparedwithuntreatedcontrols.

USDA-ARSGainesville,Fl.PennStateMicroscopyfacility.PennStateCollegeofAg.Sciences(CAS)grant.USDA-NIFAgrant#2011-67013-30352.NSF

SigniOicance

Conclusions

References

Acknowledgments

Objectives

Introduction

Abstract