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(Lepidoptera : Nolidae) in Relation to Nitrogen and Phenolics in Its Food

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(Lepidoptera : Nolidae) in Relation to Nitrogen and Phenolics in Its Food I WA¡iË INSTIT'UTÐ il.8.c6 tIt]IìARY THE PERFORMANCE OF U3ÀBA llgl\g WALKER (LEPIDOPTERÀ: NOLIDAE) IN RELÀTION TO NITROGEN ÀND PITENOLICS TN ITS FOOD. BY JÀNET DÀWN FARR B.Ag.Sc. (Hons. ) University of Àdelaide, South Australia. À thesis submitted. for the degree of Doctor of Philosophy in the Faculty of ÀgriculturaÌ Science at ' The University of Ad.elaide. Department of Entomology Waite Agricultural Re.search Institute. Septenber 1985 *n¿*nt{*"( //- ! ó'ç TO na n07q€t? DOR€€.IV AINITR8D 7AN]. Advìce fron a caterpiTTar - 'One side wÌ77 nake you g,row tal-ler¡ and the other side wì77 nake you grow shorter.û Lewis CarroTTt ATice's Adventures in tlonderTand. A late instar larvae of grgÞg lggggg Walk. and Ox¿opeq sp. (Oxyopiclae: Araneae) on Eucalypt us camal dulensis. !> i TABLE OF CONTENTS Page SUMMARY v DECLARATION vat ACKNOIVLEDGEMENTS vf-lr- CTTAPTER 1 LITERATURE REVIEW . TNSECT / EOST PLANT INTERREI,ATIONS 1 1.1 Introduction 1 1.2 Host Ptant Selection 3 1.3 Nutrition I 1.4 Nitrogen as a limiting nuÈrient 11 1.5 Feed.ing efficiencies of s¡lecialÍsts and generalists 20 1.6 Plant apparency or escape in space and time 22 t.7 Environment and plant physiologY 29 1.8 Environrnental stress and insect outbreaks 33 CTIÀPTER 2 INTRODUCTTON 38 2.1 The biology of Uraba log=g= Vla1k. 38 2.2 Eood. plants of g: lgggqg 40 i.¡ sro¡odkinrs survivar curves 4l 2.4 Survival of !L lggggg on different host plants in the context of Slobodkinrs survival curves 42 I i I t I I I I L I I I l l:- Page CHAPTER 3 THE INFLUENCE OF INDIVIDUAI. ÄMTNO ACIDS INCORPORATED TNTO ARTIFICIÀL DTETS ON TIIE BIOI.OGICAL PERFORMANCE OF U. LUGENS. 45 3.1 Introduction 45 3.2 Materials and methods 49 3.2.1 Diet 49 3.2.2 Insects 52 3.3 Results and discussion 54 3.3.1 Diet accePtance 54 3.3.2 Diet consumed 58 3.3.3 Growth 61 3.3.4 Survival 63 3.3.5 Potential fecunditY 72 CEAPTER 4 THE INFLUENCE OF VARIABI,E NTTROGEN ON THE PERFORMANCE OF U.I.UGENS FEEDING ON FRESH DIET. ?5 4.1 Introductíon 75 4.2 Materials and methods 7A 4.2.L Experiment I: The influence of vtater stress on the performance of u. Iggegg 7A 4.2"I.1 Food Plants 7A 4.2.L.2 Insects 79 4.2.2 Experiment fI: The influence of variation in nitrogen concentration in three different food plants on the survíval of first instar larvae for us tgqegg 80 lal Page 4.3 Results and discussíon 82 4.3"L ExPeriment I 82 4.3.1.1 Length of stadia 82 4.3.1.2 Survival 83 4.3.1.3 FecunditY 84 4.3.2 ExPeriment II 85 CHAPTER 5 THE INFLUENCE OF PHENOLS ON ÀSSIMILATION OF DIET BY U. LUGENS. 89 5.1 Introduction 89 5.2 Materials and methods 94 5.2.1 Diet 94 5.2.L.1 Stock solutions and mixtures 94 5.2.1.2 PreParation 95 fnsects 98 .5.2.2 5.2.3 Experimental Procedures 99 5.3 Results and discussion LO2 5. 3.1 Diet consurnPtion LO2 5.3.2 Utiiization of total di.et 104 5.3.3 Utílization of dietary nitrogen 105 CHAPTER 6 SEÀSONÀL AND ANNUAL VARIÀTION OF NITROGEN AND PHENOIJS PRESENT IN MATURE E' CAMALDULENSIS FOI,IAGE. 108 6.1 Introductíon 108 6.2 Materials and methods 111 6.2.L Determination of toÈal nítrogen 111 iv Page 6.2.2 Determination of t.otal phenols tL2 6.2.3 Determination and extraction of free amino acids 113 6.3 Results and discussion 11e 6.3.1 The va.riation of total nitrogen 118 6.3.2 Total phenols L20 6.3.3 Free amino acids L22 CHAPTER 7 GENERAL DISCUSSION 127 REFERENCES 137 APPENDTCES 168 v SUMMÀRY The performance of LEgÞê lggggg Walker was investigated in relation to the influence of nitrogen and phenolic components presenÈ in its food plants, using fresh and synthetic diets. The seasonal and annual varLation of these comPonents in Eucalyptus ="tnelclulgns i s, a good food ptant, for Ut lugens, was also examined. This was done in order to test the hypothesis that nitrogen is normally a limiting nutrient for phytophagous Ínsects, but maY become more available when its food plant is subjected to "lùater stresstr, leading to increased survival of early instars and therefore a population increase or "outbreaktr. Lar vae fed water sÈressed E. camaldulensís were potentially more fecund, although survival did not increase. For artificial diets incorporaÈing leaf powder of either good (E: g,egellglgggig) or poor (E, pfa!1pgs) food plants, the amino acids proline and valine increased Iarval performance compared with the respective base diets. Ilowevert although proline concentrations varied markedly in E. camaldulensis foliage, valine concentraÈions remaj-ned relaÈively constant throughout the study period. The najor inftuencing factor on the performance of U" lggens was the phenol quercetin, which decreased nitrogen assimilation to a much greater degree than other phenols studied (caffeic acid, chlorogenic acid and gat-lic acid). In Èhe food plant E. camaldulensis total niÈrogen and phenols v¡ere negatively correlated. TotaI nitrogen reached its maximum level in early spring and declineil to its minimum in winter but total phenols vi reached maximum levels in winter and minimum levels in early spring.The variation of individual amino acids in E' camaldul-ensis foliage was also examined. The results of this study are discussed in relation to existing hypotheses on insect host / plant inÈerrelaÈj-ons and population dynamics ând an alternative hypothesis is proposed. v1t- DECLARATION The work presented in this thesis is my own unless otherwise acknowledged, and has not previously been submitted to any university for the award of any degree or diplorna. This thesis may be made available for loan or photocopying provided that an acknowledgement is ¡nade in the instance of any reference to this work. Signed: (aI .D. Farr) September 1985 vLt-t ÀCKNOIVLEDGEMENTS I am extrernely graÈeful to ny supervisor, Dr. F.D. Morgan, for his untiring advice, criticism and encouragement given throughout the courae of this study. I am also indebÈed to Dr. À.C. Jennings who gave advice on the biochemi-cal aspects of this .project including the procedure for extraction of amino acids, and Mr. G.S. Taylor who assisted in all aspects of this study. Mr. D. Boeh¡n operaÈed the amino acid analyser and Mr. C. Rivers helped in the determination of total nitrogen. The experimental figures were photographed by Mr. B. Palk, and Mr. R. Coles phoÈographed the front plate. Dr. R. Laughlin introduced me to the "Apple" computer on which this thesis was typed and printed, and Mrs. C. SchonfeldÈ photocopied the thesis. To all these people I extend my appreciation of Èheir patience and excellenÈ help. I gratefully acknowledge the Univeristy of Adelaide for the arrrard of an UniversiÈy Research Grant Scholarship during this study. I also wish to thank my fellow students and members of the entomology department for their assisÈance, encouragement and friendship. Chapter 7 Insect / Host PTant ReTations- Literature Review. v,/AlTt ll'{5ilTU.IE l,'i!:ìAR'/ gEÀ3TEB 1 Insect / host p.Lant reTations - LÍterature Review. 1.1 INTROÐUCTION. Although insects are well known to be amongst the most abundant and successful of terresÈriaI anirnals, the evolutionary path of the phytophagous insect species has not been an easy one. Less than a third of insectan evolutionary pathways have achj-eved the ability to feed on the Spermatophyta (Southwood, 1973). DeÈhier Ã982) said: ' nÀs primary producersf green plants are the synthesizers and bankers of biochemical wealth. Herbivores are the have notst the pillagers of this biochemical wealth." This characterises well the relationship beÈween the phytophagous insect and its food plant. The insect/planÈ relationship may essentially be considered in terms of shelter, food and transport. Pl.anÈs provide food and,/or shelter for insects, while the rnobility of insects has led to the plantrs dependence on Ínsects for the transport of pollen and propagules. In utiÌizing green plants for food and shelter insects have had to overcome a number of problems: (a) survivíng the extremes of temperature and dessication; (b) attachment to, and penetration ofr what may be a tough, waxy, hairy or spiny surface; (c) locating a particular plant specíes amongst a great variety on which Èhe insect cannot survive and (d) ingestion, consumption and digestion of a food source which may be 2 suboptimal for nutritional or other reasons (SouÈhwood' 1973). In this revíew on insect/pì-ant, relations Èhe latter two "evolutionary hurdles", host selection and the provision of adequate nutrition¡ especially the role of nitrogenr will be considered with some emphasis on the physiology of the plant and its relationship to the growth and survival of insecÈ leaf feeders. Most adult insects are able to select their own diets, but many larval forms are Ínfluenced by the choice of food plant made by the ovipositing female. Such larvae however, in common wiÈh those that experience a greater or lesser degree of foraging are none-the-Iess endowed with the ability to identify a preferred diet. I'lhereas this ability may serve no apparent purpose in the more sedentary forms, it is as critical to free foraging larvae as to resultant dispersing adults. This is especialJ.y true in cases where the female may not oviposit on the food plant (Erlich and Raven, L964) directly but in its vicinity. Some species may oviposit in the vicinity of t.he preferred host (eg.
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